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CONTENTS

PIN ge bt haw

Volume 119, Number 1

March 28, 2005

ISSN 0028-1544

Kevin J. Roe

Paul D. Hartfield

Russell L. Minton

Steven P. Savarese, Jr.

Frank Kohler

Matthias Glaubrecht

John Slapcinsky

Angel Valdés

Kelvin Barwick

Claude Vilvens

Hamiota, a new genus of freshwater mussel (Bivalvia:

Unionidae) from the Gulf of Mexico drainages of the

southeastern United States

Consideration of genetic relationships in management

decisions for the endangered Anthony’s riversnail, Leptoxis

crassa anthonyi (Redfield, 1854) (Gastropoda:

Pleuroceridae)

Fallen into oblivion—the systematic affinities of the

enigmatic Sulcospira Troschel, 1858 (Cerithioidea:

Pachychilidae), a genus of viviparous freshwater gastropods

TVOMY, JAVA: 2 pidatsie syooae dh oH 4 48 4 Hamed gd Gea dalaed. aetnhignna. a2

Six new species of Paryphantopsis (Gastropoda: Pulmonata:

Charopidae) from the Papuan Peninsula of New Guinea...

First record of Akera Miiller, 1776, from the eastern

Pacific, with the description of a new species ............

Description of Calliotropis pulvinaris new species

(Gastropoda: Trochidae: Eucyclinae: Calliotropini) from

West Madagascar

Pe ee al

4 Batre de UW Gaal Been 50

Sponsored in part by the State of Florida, Department

of State, Division of Cultural Affairs, the Florida Arts

Council and the National Endowment for the Arts.

NATIONAL

ENDOWMENT

FOR THE ARTS

THE NAUTILUS L19(1):1-10, 2005

Page 1

Hamiota,

(Bivalvia:

Kevin J. Roe

Delaware Museum of Natural History U.S

4840 Kennett Pike

Wilmington, DE 19807 USA

[email protected]

Paul D. Hartfield

. Fish and Wildlife Service

6578 Dogwood View Parkway, Suite A

Jackson, MS 39213 ,

[email protected]

a new genus of freshwater mussel

Unionidae) from the Gulf of Mexico drainages

of the southeastern United States

USA

ABSTRACT

Hamiota, a new genus of freshwater mussel containing four

species formerly assigned to the genus Lampsilis Rafinesque,

1820, is described. In addition to the genus Lampsilis, mem-

bers of Hamiota had previously been placed in the genera Vil-

losa Frierson, 1927, and Ligumia Swainson, 1840. Several char-

acters including the packaging of their larvae in a superconglu-

tinate lure to attract host fishes, placement and shape of the

marsupia, and release of glochidia through the excurrent si-

phon, support the recognition of these species as a distinct

genus.

INTRODUCTION

Following the discovery in 1988 by Robert Butler of the

first superconglutinate lure ensnared on a snag in a trib-

utary of the Choctawhatchee River, a sumber of publi-

cations (e.g., Haag et al., 1995; Hartfield and Butler,

1997: O’Brien and Brim Box, 1999: Blalock-Herod et al.,

2002) have confirmed through direct observation the

supposition that these unique Tures are produced by four

species of freshwater mussels endemic to the Gulf of

Mexico drainages of the southeastern United States.

Herein, we confirm earlier published suggestions that

these four species represent a distinct genus of fresh-

water mussels (Fuller and Bereza, 1973; O’Brien and

Brim Box, 1999). The recognition of this genus is based

on several characters including the production of the su-

perconglutinate lure, and the unique shape and place-

ment of the marsupia (the region of the demibranchs

where female unionoid mussels brood developing lar-

vae), and is supporte od by mole cular evidence (Roe et al.,

2001). Use of marsupié al { eatures is consistent with pre-

vious designations of unionid “divisions” initiated by

Simpson (1900a) and continued by Heard and Guckert

1970). The designation of Hamiota increases the num-

ber of North American unionids genera to 50 (Turgeon

et al., 1998S).

MATERIALS AND METHODS

A list of specimens examined is included in Appendix 1.

Acronyms used in the text are: Academy of Natural Sci-

ences of Philadelphia (ANSP), Delaware Museum of

Natural History (DMNH), Florida Museum of Natural

History (UF), Unite sd States National Museum (USNM),

University of Alabama Unionid Collection (UAUC), and

Mississippi Museum of Natural Science (MMNS). Ab-

breviated synonymies are presented for bas taxon and

include novel combinations and publications with illus-

tration. Measurements were taken to the nearest 0.05

mm using dial calipers.

SYSTEMATICS

Family Unionidae Rafinesque, 1520

Tribe ‘Lampsilini von Ihering, 1901

Hamiota new genus. Type species: Hamiota subangulata (Lea,

1840) by original designation

Diagnosis: A monophyletic group of freshwater bi-

valves (Roe et al., 2001) in which all of the glochidia are

released simultaneously encased in mucous packages

that are referred to as superconglutinates (Haag et al.,

1995, fig. 1). The supe rconglutinate lure exits the aaile

cavity via the excurrent opening and is encased within a

transparent mucous tube (Hartfield and Butler, 1997;

O’Brien and Brim Box, 1999). When acted upon by wa-

ter currents the superconglutinate mimics the move-

ments of a swimming fish, and has been shown to elicit

attacks from fishes (Haag and Warren, 1999). The mar-

supium is restricted to the ventral portion of the outer

demibranchs of female mussels (Figure 1). The precise

shape and pigmentation of the marsupia, as well as the

degree of posterior mantle margin development, varies

across species.

Description: Members of this genus are small- to me-

dium-sized freshwater bivalves, and adult valves gener-

ally are between 45-100 mm in length. Shells range

from ovate to elliptical in outline, and are somewhat

Page 2

THE NAUTILUS, Vol. 119, No. 1

Figures 1, 2.

Inner mantle and outer demibranchs of gravid superconglutinate-producing mussels in the genus Hamiota and

inner mantle and outer demibranchs typical of non-supe »rconglutinate- -producing mussels of the genera Lampsilis, Ligumia, and

Villosa. 1. Female Hamiota australis. 2. Female Villosa vibex.

compressed to moderate ly inflated. Shell thickness rang-

es from heavy to thin. Sexes display some degree of ae

morphism in ‘shell sh: ape. Shells of male musse cis are typ-

ically more acutely pointed posteriorly, whereas shells of

female mussels disy slay an ante posterior margin,

Periostracum is typically smooth, but can be very glossy

in some species. Background color ranges from. dark-

brown and black through chestnut-brown to straw-yel-

low. Black to bright green rays of variable width are of-

ten present and may be limited to the posterior slope,

or cover the entire disk. Nacre color is typically white

although other colors such as salmon or blue may be

seen as well, particularly in the beak cavities or the pos-

terior margins. The m: arsupia are often asymmetrical in

shape; the anterior portion is typically broadest, tapering

toward the posterior end. The ventr: dm: wein of the mar-

supium is darkly pigmented in gravid fe males, Pigmen-

tation of marsupia varies across species and populations

and colors may include purple, red, black, or white. In

females, the mantle margins anterior to the branchial

opening are elaborated to varying degrees. The mor-

phology of the glochidial valves of qwember of Hamiota

is similar to that of members of Lampsilis or Villosa

(O’Brien and Brim Box, 1999).

Etymology: Hamiota = angler. Derived from the

Greek word hans, meaning hook. This name refers to

the means by which members of this genus attract host

fishes by packaging their parasitic larvae in a lure that

mimics a small fish.

Remarks: Species of Hamiota generally have been

treated as Lampsilis ce to the similarity in shape and

coloration of their shells. Some authors also have in-

cluded these species in the genus Villosa or Ligumia,

due to shell shape, thickness, and/or ornamentation of

the mantle flap. The shells of Hamiota species are in-

K. J. Roe and P. D. Hartfield, 2005

» a |

Page 3

deed similar to the shells of these genera, and the man-

tle flap may exhibit characters of both Lampsilis and

Villosa. However, in Hamiota, the marsupia is restricted

to the ventral half of the posterior portion of the outer

demibranchs, while in Lampsilis, Ligumia, and Villosa

the marsupium fully occupies the water tubes of the pos-

terior portion of the outer demibranchs (Figure 1). Un-

like members of Lampsilis, which release larvae through

pores in the ventral edge of the demibranchs, members

of Hamiota release the superconglutinate lure through

the excurrent opening. The placement and shape of the

marsupium and the extraordinary method of glochidial

release and host fish attraction, the superconglutinate,

are uniquely derived characters in the Lampsilini that

warrant Petite recognition.

Hamiota altilis (Conrad, 1834) new combination

Unio altilis Conrad, 1834; Conrad, 1834: 43, pl. 2, fig. 1;

Chenu, 1845: 21, pl. 1, fig. 1; Reeve, 1865; pl. 23, fig. 109.

Margarita (Unio) altilis (Conrad, 1834); Lea, 1836; 24,

Margaron (Unio) altilis (Conrad, 1834); Lea, 1852a: 27.

Lampsilis altilis (Conrad, 1834); Simpson, 1900a: 529; Par-

malee and Bogan, 1998: 125, pl. 47.

Unio clarkianus Lea, 1852; Lea, 1852b: 251; Lea, 1852c: 273,

pl. 21, fig. 30; Lea, 1852d: 29, pl. 21, fig. 30.

ieee eon (Unio) clarkianus (Lea, 1852); Lea, 1852a: 27.

Lampsilis clarkianus (Lea, 1852); Simpson, 1900a: 532.

Unio gerhardtii Lea, 1862; Lea, 1862a: 168; Lea, 1862b: 208,

pl. 31, fig. 277; Lea, 1862c: 30, pl. 31, fig. 277.

Margaron (Unio) gerhardtii (Lea, 1862); Lea, 1870: 35.

Lampsilis (Lampsilis) gerhardtii (Lea, 1862); Simpson, 1900a:

532.

Unio doliaris Lea, 1865; Lea, 1865: 88; Lea, 168: 260, pl. 32,

fig. 75; Lea, 1869: 20, pl. 32, fig. 75.

Margaron (Unio) doliaris (Lea, 1865); Lea, 1870: 42.

Lampsilis (Lampsilis) doliaris (Lea, 1865); Simpson, 1900a:

533.

Description: Described by Conrad (1834) as sub-oval,

thin, and inflated. The periostracum was described as

“rugose” and “blackish” with rays and “numerous short

vermicular lines on the posterior slope,” and the nacre

as whitish and iridescent. The periostracum of speci-

mens of H. altilis is typically brown to chestnut-brown

in color with a variable number of dark green rays. The

left valve has two heavy, spatulate pseudoc -ardinal teeth,

the smaller above the larger. The right valve has two

nearly equally sized triangular teeth, the larger anterior

to the smaller. The later an teeth are short but blade-like,

two in the right valve, and one in the left

The posterior mantle margin of the Saale is expand-

ed into a well-developed flap with papillae along the bor-

der. Coloration, number, and size of papillae vary some-

what between populations. In general, however, the in-

terior mantle flap is colored red to dark red or brown

with darker spots, while the exterior of the flap is brown

to black, often with vertical lighter bars, and with a

small, but prominent dark “eye spot” on the posterior

end. Small papillae are present along the mantle flap,

usually becoming more robust anteriorly. In males, the

mantle flap is typically not expanded and is reddish in

color. Mar supia of H. altilis are finely tapered at each

end when immature, becoming bro: idly rounded on the

ends in most populations, tapering anteriorly in others.

Marsupia color is a dark reddish- Dione or black along

the margin and white above. The anus is usually pig-

mented red and the incurrent and excurrent siphons are

reddish-brown to black. The glochidia of H. altilis are

described by Haag et al. (1999).

Type Material: Unio altilis Conrad, 1834, Lectotype

ANSP 56419 (Figures 3, 4) here designated. Type lo-

cality: Alabama River, near Claiborne [Monroe Co., Al-

abama|.

hela clarkianus Lea, 1852, Type not found. Type local-

: Williamsport, [Maury Co.], Tennessee; Georgia or

Unio doliaris Lea, 1865, Lectotype USNM 84936, here

designated. Type locality: Etowah River, Georgia.

Unio gerhardtii. Lea, 1862, Holotype USNM 25711 by

monotypy. Type locality: Chattanooga, Georgia.

Remarks: The most variable species included in Ham-

iota, H. altilis, is endemic to the Mobile River Basin.

Some of the conchological variation is undoubtedly eco-

phenotypic in nature, although the extent and nature of

the variation in shell shape and pigmentation has not

been adequately expl ored.

Conrad (1834) in his original description did not iden-

tify a primary type. Johnson and Baker (1973) identified

ANSP 56419 as the figured holotype, although the spec-

imen label indicates the locality as “Ogeeche re R., Ga.”

Johnson and Baker (1973) state that the label is in error,

and “probably was mixed with ANSP 46415, which is

labeled ‘Claiborne, Alabama’, by error.” Conrad (1534)

clearly indicated that the specimen(s s) of U. altilis de-

scribed were collected from the “Alabama River, near

Claiborne.” Conrad (1834) did not specifically designate

a holotype and according to the ICZN recomme ndation

73F and Article 74.5 the holotype designation of John-

son and Baker (1973) is deemed inv alid. In order to

preserve stability of nomenclature, we herein designate

specimen ANSP 56419 as the le sctotype of U. altilis. In

accordance with ICZN Article 74.7, we herein designate

the specimen USNM $4936 as the lectotype of U. do-

liaris Lea in order to maintain taxonomic stability and

because this specimen appears to be that figured by Lea

(1868). Hamiota altilis is considered threate ned by the

United States Fish and Wildlife Service (USFWS, 1994).

Life History: Mature gravid females have been re-

ported from March through June. Hamiota altilis have

also been observed releasing glochidia in a superconglu-

tinate (Haag et al., 1999). Large centrarchid fishes, in-

cluding Micropterus coosae Hubbs and Bailey, ee M.

punctulatus (Rafinesque, 1S19), M. salmoides (Lace-

péde, 1802), and Lepomis cyanellus Rafinesque, 1a18,

have been confirmed as suitable hosts (Haag et al.,

1999).

Range: Hamiota altilis was historically reported

SS SESS ES SE

Page 4 THE NAUTILUS, Vol. 119, No. 1

Figures 3-10. Type material of species of Hamiota Photographs are of the interior of left valve and exterior of right valve. 3,

1. Lectotype of H. altilis ANSP 56419. 5, 6. Holotype of H. australis USNM 150473. 7, 8. Lectotype of H. perovalis ANSP 56416

9, 10. Lectoty pe ol H. subangulata USNM 8580]

K. J. Roe and P. D. Hartfield, 2005

Page 5

throughout the Mobile River Basin, including the Tom-

bigbee, Black Warrior, Cahaba, Alabama, Tallapoosa, and

Coosa River drainages in Alabama, Georgia, Mississippi,

and Tennessee. The species is currently restricted to lo-

calized portions of the Cahaba, Coosa, and Tallapoosa

rivers and some of their tributaries (USFWS, 2003).

Hamiota perovalis (Conrad, 1834) new combination

Unio perovalis Conrad, 1834; Conrad, 1834; 43, pl. 2, fig. 2:

Chenu, 1845: 21, pl. 1, fig. 2; Kiister, 1861: 257, pl. 87,

fig. 2; Reeve, 1866: pl. 38, fig. 209.

Margarita (Unio) perovalis (Conrad, 1834); Lea, 1836: 24.

Margaron (Unio) perovalis (Conrad, 1834); Lea, 1852a: 27.

Lampsilis perovalis (Conrad, 1834); Simpson, 1900a; 531.

Unio spillmanii Lea, 1861; Lea, 1861: 39, Lea, 1862d: 98, pl.

15, fig. 246; Lea, 1862e: 102, pl. 15, fig. 246, Reeve, 1868:

pl. 82, fig. 435.

Margaron (Unio) spillmanii (Lea, 1861); Lea, 1870: 42.

Lampsilis (Lampsilis) spillmani (Lea, 1861); Frierson, 1927: 69

[misspelling].

Description: Conrad (1834) described this species as

oval and inflated with a moderately thick shell. He noted

two color varieties, one in which the periostracum was

olivaceous and obscurely rayed with white nacre and an-

other in which the periostracum was reddish-brown with

“rose colored” nacre. The periostracum of specimens of

H. perovalis is generally lighter in color than H. altilis

and range from straw -yellow to light brown. The number

of rays is variable and can cover fhe entire disk. The left

valve contains two robust equal sized pseudocardinal

teeth. The right valve has two pseudocardinals and the

anterior tooth is smaller than the posterior tooth. The

lateral teeth are elongate, two in the left, one in the

right.

The mantle margins of female H. perovalis are ex-

panded into well-developed flaps, pigmented red on the

interior and darker red to brown or black on the exterior.

No eyespot is present and short papillae are present

along the mantle edge. Males possess a rudimentary

mantle margin with w eak pigmentation and few papillae.

The marsupia of H. perovalis are pisciform in shape,

broader anteriorly and narrowly tapering posteriorly.

The marsupium is reddish or darker along the margin,

often with a darker spot of pigment on thie broader an-

terior end that resembles an eyespot in the supercon-

glutinate lure. The anus can be pigmented red and

black, and the incurrent and excurrent siphons are usu-

ally reddish or brown in color.

Type Material: Unio — Conrad, 1834, Lecto-

type ANSP 56416 (Figures 7, 8), here designated. Type

locality: Alabama River, at Clabore [Monroe Co., Al-

abama].

Unio spillmanii Lea, 1861, Lectotype USNM 84925

here designated. Type locality: Luxpalila Creek, near Co-

lumbus, Mississippi.

Remarks: There has been some question as to the dis-

tinctiveness of H. perovalis from H. altilis, perhaps be-

cause both were described from practically the same lo-

cality. An examination of mitochondrial DNA sequences

by Roe et al. (2001) recovered these two taxa as a clade,

but failed to resolve them into re ciprocally monophyletic

groups. Hurd (1974) considered perovalis a junior syn-

onym of altilis, as did Burch (1975). Frierson (1927)

considered U. doliaris (Lea, 1865) a synonym of perov-

alis, although Parmalee and Bogan (1998) include U. do-

liaris as a synonym of altilis. Based on the collection

locality and ‘the appearance of the type specimen, we

place U. spillmanii Lea, 1861 as a synonym of H. per-

ovalis. As with U. altilis, ron §34) did not specifi-

cally designate a holotype for U. perovalis, therefore,

accor ding to the ICZN Article a 5 and recommendation

73F the holotype designation of Johnson and Baker

(1973) is deemed to be in error. In an effort to maintain

nomenclatural stability we herein designate ANSP 56419

the lectotype for U. perovalis. Accor ding to ICZN Arti-

cle 74.7, the specimen USNM 84925 is here des ignated

as the lectotype of U. spilmanii in order to fix the name

and maintain nomenclatural stability. This species is list-

ed as threatened by the United States Fish and Wildlife

Service (USFWS, 1994).

Life History: Discharge of superconglutinates was

first observed in H. perovalis (Haag et al., 1995). Glo-

chidia mature and are discharged between March and

June, with releases concentrate din early April (Hartfield

and Butler, 1997). Micropterus coosae, M. punctulatus,

and M. salmoides have been identified as suitable host

fishes for the orange-nacre mucket (Haag and Warren,

1999).

Range: Hamiota perovalis was historically known from

the Mobile Basin’s Alabama, Tombigbee, Black Warrior,

and Cahaba rivers and their tributaries in Alabama and

en eae The species has apparently become extir-

pated from the main channels of the larger rivers, but

continues to survive in some tributaries of “all four drain-

ages (USFWS, 2003).

Hamiota subangulata (Lea, 1840) new combination

Unio subangulatus Lea, 1840; Lea, 1840: 287; Lea, 1$42a: 209,

pl. 13, fig. 23; Lea, 1$42b: 47, pl. 5, fig. 23; Kiister, 1861:

278, pl. 94, fig. 2; Simpson, 1892: 415, pl 58, fig. 1; Reeve,

1868. pl. 65, fig. 327.

Margaron (Unio) subangulatus (Lea, 1840); Lea, 1$52a: 29.

Lampsilis subangulatus (Lea, 1840); Simpson, 1900a: 556;

Clench and Tumer, 1956: 196, pl. 2, fig. 2.

Ligumia subangulata (Lea, 1840); Haas, 1969: 443.

Villosa subangulata (Lea, 1840); Heard, 1979: 44.

Unio kirklandianus S$. H. Wright, 1897; S. H. Wright, 1S97:

136.

Lampsilis kirklandianus (S. H. Wright, 1897); Simpson, 1900a:

557; Simpson, 1900b: 76, pl. 1, fig. 7.

Description: A medium-sized mussel that reaches ap-

proximately $5 mm in length (Brim Box and Williams,

2000). Specimens are gene ae elongate; the posterior

ridge is rounded and the posterior slope is usué uly con-

Page 6

THE NAUTILUS, Vol. 119, No. 1

cave. Periostracum color is variable in this species. In-

dividuals range from straw-yellow to chestnut- brown in

color with a variable number of black to bright emerald

green rays of variable width. These rays emanate from

the umbo and radiate across the disk. Most shells are

shiny, although some populations exhibit an extremely

glossy periostracum, This species has been described as

one of the most beautiful of all North American fresh-

water mussels (S. H. Wright, 1897; Clench and Turner,

1956).

Wright (1897) described U. kirklandianus from the

Ochlockonee River in Leon County, Florida and re-

marked that his specimens were more polished and had

broader rays than typical of U. subangulatus, and that

the shells were “deeper and broader.” The right valve

has two somewhat spatulate pseudocardinal teeth, the

smaller nearly directly above the other. The left valve

has two pseudocardinal teeth, the anterior one much

larger than the other. Lateral teeth are thin, but not

dekeate. two in the left and one in the right valve. Ex-

amination of specimens from across the range of this

species reveals substantial variation in shell color and

size. The mantle margins in females are only slightly ex-

panded into a sncdlesk flap. The flaps are light brawn in

coloration on the interior and freckled- brown on the ex-

terior, and no eyespot is present. Short papillae are pres-

ent along the margin of the flap, becoming larger ante-

riorly, In males, the mantle is only slightly expande od,

without pigment, and with very short papillae. The mar-

supia are pisciform in shape, broader anteriorly and ta-

pered behind, and darkly pigmented along the mar gin

often with a darker spot of pigment adtenoaly The anus

is unpigmented and the siphons are brownish. Glochidia

were figured and described by O’Brien and Brim Box

(1999).

Type Material: Unio subangulatus Lea, 1840, Lecto-

type USNM 85801 (Figures 9, 10) designated by Clench

and Turner (1956). Type locality: @hatahooskes River,

Columbus, [Muscogee Co.,] Georgia.

Unio kirklandianus S. H. Wright, 1597, Paratype USNM

149648. Type locality: Oclocknee River, Leon Co., Flor-

ida.

Remarks: Hamiota swhangulata is listed as an endan-

gered species (USFWS, 1998). The analysis of Roe et al.

(2001) resolved this taxon as monophyle tic and in a clade

with H. australis.

Life History: Discharge of superconglutinate lures

has been documented by O’Brien and Brim Box (1999)

from late May through early June. Micropterus punc-

tulatus and M. salmoides appear to be primary hosts for

the species (O’Brien and Brim Box, 1999).

Range: Hamiota subangulata was found throughout

the Apalachicola River Basin and the Ochlockonee River

drainage (Brim Box and Williams, 2000). Currently the

species continues to survive in some small streams and

headwater rivers (USFWS, 1998).

Hamiota australis (Simpson, 1900) new combination

Lampsilis australis ou ou 1900; Simpson, 1900a: 544; Simp-

son, L900b: 75, pl. 2, fig. 2; Clench and Tumer, 1956: 199,

pl. 2, fig. 3

Ligumia australis (Simpson, 1900); Haas, 1969: 432.

Villosa australis (Simpson, 1900); Heard, 1979: 44.

Description: Simpson (1900b) described the shells of

a species as “long and elliptical . .. moderately inflat-

* The periostracum was described as “smooth, shin-

ng greenish yellow, rayed with green” and the nacre

“bright bluish white.” Specimens examined were el-

liptical to elongate oval and often terminate in a blunt

point. Specimens are almost always stained black, with

some green rays visible on the posterior slope. The per-

iostracum is often glossy. The right valve contains two

pseudocardinal teeth, the smaller above the larger. The

left valve has two compressed pseudocardinal teeth. Lat-

eral teeth are elongate and slightly curved, two in the

left valve and one in the right. Posterior mantle flaps are

poorly developed with a aieal OF ved.ctain along the

margins. There is no eyespot on the mantle flap, and

only: a few very small papillae. The marsupia are pisci-

form, broadly rounded anteriorly and tapering behind

(Figure 1). Color of the marsupium is white along the

margin and black above. The anus is unpigmented and

the incurrent and excurrent siphons are reddish in color.

Glochidia were described and figured by Blalock-Herod

et al. (2002).

Type Material: Lampsilis australis Simpson, 1900,

Holotype USNM 150473 by original designation (Fig-

ures 5, 6). Type locality: Little Patsiliga Creek, south-

eastern Alabama.

Remarks: Fuller and Bereza (1973) stated that this

species represented an “undescribed lampsiline genus”

and that its marsupium allied it closely with Ptychobran-

chus. This species was incorrectly synonymized with Pty-

chobranchus jonesi (van der Schalie, 1934) by Clench

and Turner (1956). The phylogenetic analysis of Roe et

al. (2001) clearly placed this species with ‘the other su-

perconglutinate } yroducers.

Life History: Superconglutinate releases have been

documented in this species by Blalock-Herod et. al.

(2002). Micropterus spp. are likely hosts (Blalock-Herod

et al., 2002).

Range: Hamiota australis was known historically from

the Escambia, Yellow, and Choctawhatchee River sys-

tems. It continues to survive in some river and stream

segments within these systems. This species is not cur-

re sntly protected under the End: angered Species Act.

ACKNOWLEDGMENTS

We express thanks to the many individuals who have

worked with this unique group of mussels and directly

or indirectly assisted in the production of this work. Sev-

eral museum curators provide sd access to the spe cimens

K. J. Roe and P. D. Hartfield, 2005

Page 7

in their care including the National Museum of Natural

History (Dr. Robert Hershler), The Florida Museum of

Natural History (Dr. Fred Thompson), The Academy of

Natural Sciences of Philadelphia (Dr. Dan Graf), The

University of Alabama Malacology Collection (Dr.

Charles Lydeard), and the Mississippi Museum of Nat-

ural Science (Dr. Robert Jones and Leann Staton). The

comments of several individuals greatly improved this

manuscript: We are grateful to Art Bogan, Jettrey Gar-

ner, Dan Graf, Jeanne Serb, and Jim Williams. Gary

Bloomer (DMNH) assisted with figures.

LITERATURE CITED

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APPENDIX 1

SPECIMENS EXAMINED

Hamiota altilis

ALABAMA

Alabama River: USNM 25948; Alabama River, near Clai-

borne: ANSP 56419 [Lectotype U. altilis Conrad, 1834],

ANSP 56415; Big Swamp Creek, Macon Co.: USNM

361723; Jackson Shoals, Choccolocco Creek: ANSP

103834, ANSP 103871; Beaver Creek: ANSP 103863;

ieee River: ANSP 41120, DMNH 130623; Coosa Riv-

, Coosa Valley: ANSP 103771; Higgin’s Ferry, Coosa

oe Chilton Co.. USNM 218118: McGowen’s Ferry,

near Wilsonville, Coosa River: USNM 521359: Weduska

Shoals, Coosa River: ANSP 48001, DMNH 075252,

DMNH_ 150037, DMNH 150038, USNM 348970,

USNM 452169; Coosa River, [incomprehensible hand-

writing] Shoals, Shelby Co.: ANSP 341399; Shoal Creek,

Pine Glen Recreational Area, Cleburne Co.: MMNS

7743, MMNS S0S4, MMNS 8085, UAUC 120, UAUC

121, VAUC 125; Yellowleaf Creek, Jumbo, Chilton Co.:

ANSP. 89031; Cane Creek, West of CR 65, 2 mi. West

CR 7S Jet., T15S, RIIE, sec 3: UAUC 3292, MMNS

SOS1; Little Cane Creek, at CR 78, East of Edwardsville:

UAUC 3293; Chewacla Creek at CR 22, ~4 mi. East of

Tuskeegee, Macon Co.: UAUC 246, UAUC 247, UAUC

248; Uphappee Creek, 0.5 mi. upstream of Hwy. 29,

Macon Co.: MMNS 8082: Cahaba River: USNM

152026: ue River: oe 126054; Little Cahaba

River, 0.5 mi. below Cahaba Beach Rd. bridge, Jefferson

Co.; UAUC 149: Coosa River, Weduska Shoals: Shelby

Co.; UF 3955: Coosa River at Fort William Shoals, Tal-

ladega Co.: UF 65420; Hurricane Creek, Cherokee Co.:

UF 175098; Chewacla Creek, 8 mi. ESE of Tuskeegee

Co. Rd. 22, Macon Co.: UF 202249; Shoal Creek, St.

Clair Co.: UF 245989; Tuskeegee National Forest, Ma-

con Co.: UF 266048; Cahaba River: UF 269576; Shoal

Creek, St. Clair Co.. MMNS 8083.

GEORGIA

Etowah River: USNM 84936 [Lectotype, U. doliaris

Lea, 1865]; Chattooga River: ANSP $9102; Chattooga

K. J. Roe and P. D. Hartfield, 2005

Page 9

River, Chattooga Co.: USNM 59527; Chattanooga:

USNM 25711 [I iolonpert ’. gerhardtii, ‘Lea, 1862]; Fish

Creek at Highway 278/ GA State Rt. 6, ~3.9 air mi.

West of Rockm: art, Polk Co.: UAUC 538, UAUC 539:

Conasauga River at Tibbs Bridge Murray CR 109 (Whit-

field CR1OO), Murray/Whitheld Co’s.: UAUC 515,

MMNS S092; Conasauga River, Muskrat Midden, Tri-

togonia Shoals (CRM 46.70), Whitfield/Murray Co’s.:

UAUC 376; Conasauga River, south of state line, Murray

Co.: MMNS S090; Conasauga River: DMNH_ 150124,

USNM $4937, USNM 348969: Etowah River: USNM

123202.

TENNESSEE

Conasauga River: DMNH 014683; Conasauga River,

Conasauga: ANSP 341305, ANSP 347949; Conasauga

River, Polk Co.;: MMNS SO91.

Hamiota perovalis

ALABAMA

Alabama River: ANSP 56416 [Lectotype U. perovalis

Conrad, 1834], ANSP 333496; Alabama River, Clai-

borne: USNM 84938: Coosa River: ANSP 56415; Coosa

River, Gadsen: ANSP 126051; Coosa River, Talladega

Co.: ANSP 126048; Black Warrior River: ANSP 88483:

Mulberry River [Fork, Black Warrior River]: ANSP

88485: Rush Creek [Black Warrior River Dr.], FS Rt.

245, Winston Co.: MMNS 7745, MMNS 8088, UAUC

426; Flannigan Creek at FS Rd. 229, Lawrence Co.:

MMNS 7744, UAUC 423, UAUC 424, UAUC 425;

Toadvine, Valley Creek, Black Warrior River Dr., Jeffer-

son Co.: UF 65302, UF 65304; Forks of the Warrior

River, Walker Co.: UF 65305; North River, near Hagler’s

Mill, Black Warrior Dr., Tuscaloosa Co.: UF 65306; Sip-

sey Fork at N.F. 234, Bankhead National Forest, Win-

ston Co.: UF 79069, UF 79072, UF 79082, UF 79085,

UF 79136, UF 79137; Brushy Creek above Brushy Lake

Recreational Area, Bankhead National Forest, Winston

Co.: UF 79094: Capsey Creek, 50 mi. from Jct. with

Brushy Creek, Bankhead N.F., Winston Co.: UF 79115;

Sipsey Fork at mouth of Hurricane Creek, Bankhead

National Forest, Winston Co.: UAUC 95; Brown Creek,

Bankhead National Forest, Winston Co.: UAUC 1774:

Alabama: ANSP 126049: North River, near Samantha

[Black Warrior River], Tuscaloosa Co.. UAUC 107; But-

tahatchee River, Hamilton: ANSP 100657, DMNH

075231: Tuscaloosa Co. Alabama: DMNH 146496; Black

Warrior River, Jefferson Co.: UF 269609; Squaw Shoals,

Black Warrior River, Jefferson Co.: UF 65298, UF

65299. UF 65300. UF 65303. UF 65307. UF 65429. UF

269515: Garden City, Mulberry Fork: UF 69207, UF

244558: Banks of Brushy Creek, at N.F. Rd. 255, Bank-

head N.F, Winston Co.: UF 69269, UF 79177, UF

79178, MMNS 7748, MMNS SOS9: Sipsey Fork, 1 mile

N. of AL Hwy. 33 crossing, Winston Co.: UF 69279;

Brushy Creek at N.F. Rd., Bankhead N.F., Winston Co.:

UF 79076; Sipsey River at Sipsey Recreational Area,

Bankhead N.F., Winston Co.: UF 79089: Borden’s

Creek, 1 mi., upstream of from Jct. Sipsey Fork, eo y

Wilderness, Bankhead National Forest, Winston: | os o.

UF 79092, UF 79151; Sipsey Fork at N.F. Rd.,

Bankhead National Forest, Winston Co.: UF ee

Limestone Creek, 6.3 mi WNW of Monroeville, Monroe

Co.: UF 197636; Blackwater Creek upstream from Har-

ris bridge, Walker Co.: UF 266369; Sipsey River, 1.6 mi

N. of Pleasant Ridge, Greene Co.; UF 197671; North

River at Co. Hwy. 30, Fayette Co.: UF 197686; Sipsey

River, 200 m. below Co. Hwy 23, Greene ho UF

197552; Sipsey River, 4-6 mi. below Co. Hwy. 2, Greene

Co.: UF 197566; Sipsey River at CR 2, Lanta of

boat ramp, Pickens Co.: UAUC 156; Tombigbee River:

ANSP 126053; Elrod, Sipsey River, Tombigbee R.:

DMNHEL 146493; Lubbub Creek, at CR 24, 3.25 mi.

Northeast of Aliceville, Pickens Co.: UAUC 67: Tombig-

bee River: USNM 159989: Lubbub Creek, 1.8 mi SSE

of Aliceville above Hwy 14: UF 197619, UF 197632

Sipsey River, 3.6 mi. W. of Jena downstream of CR 2,

Greene Co.: UF 197697: Sipsey River near confluence

with C varpe nter’s Creek, Greene Co.: UF 197801; Sipsey

River, 5.7 mi. NNE of Mantua, Greene Co.: UF 197857,

UF 197862; Sipsey River, near Elrod, Tuscaloosa Co.:

UF 269559, UF 65301; Trussels Creek at CR 19 bridge,

Greene Co.: MMNS SOS7.

MISSISSIPPI

Luxpalila Creek, near Columbus: USNM 84925 [Lec-

totype, U. spillmanii Lea, 1561], USNM 123279; Colum-

bus, Lowndes Co.: UF 269560.

Hamiota subangulata

ALABAMA

Uchee Creek, Russell Co.; UAUC 116.

FLORIDA

Chipola River, Look-Tremble Falls near Alpha, Calhoun

Co.: ANSP 175750; Chipola River, near Pole Bluff land-

ing, Calhoun Co.: ANSP 175751; Chipola River: ANSP

84324; Ochlockonee River: DMNH 150098; Ochlocko-

nee River, Leon Co.: USNM 149648 [Paratype, U. kirk-

landianus 8. H. Wright, 1897]; Ochlockonee River, Tal-

lahassee, Leon Co.: ANSP 156892, ANSP 341307; Och-

lockonee River, 7 mi. west of Tallahassee: ANSP 157553:

Ochlockonee River, 10 mi. west of Tallahassee, Leon

Co.: ANSP 159126; Ochlockonee River, 11 mi. north-

west of Tallahassee: DMNH 119506; Ochlockonee Riv-

er, 2 mi. west of Bloxham, Liberty Co.: ANSP 360553;

Spring Creek, Marianna: ANSP 160210; State Rt. 167,

1 mi. north of Marianna, Chipola River, Jackson Co.:

ANSP : seen Spring Creek, Reynoldsville, Seminole

Co.: UF 177: 1 mi. north Marianna, C Jhipola River: UF

390; 3.5 mi. ane of Quincy, Little River: UF 415: ca. 2

Page 10

mi. east of Clarksville, Chipola River, Calhoun Co.: UF

418; Chipola River, 9.2 km ENE Kinard, 12.5 km NW

Lewis, 16.4 km N. Ida, Calhoun Co.: MMNS 8099.

GEORGIA

Chattahoochie River: ANSP 56477; Chattahoochie Riv-

er, Columbus: USNM 85081 [Lectotype, U. subangula-

tus Lea, 1840]; ANSP 126272: Cooleewahee Creek, 0.9

mi. NE of Newton, Baker Co.: USNM 853746; Coolee-

wahee Creek at GA Rt. 91, Baker Co.: MMNS 8095;

Abram’s Inlet, Flint River: ANSP 190294; Mill Creek,

Flint River, several mi. north of Albany: ANSP 267572;

Kinchafoonee Creek at GA Rt. 41 crossing, just south of

Preston, Webster Co.: UAUC591:; Kinchafoonee Creek

at GA Rt. 49 bridge ~9 air mi. northeast of Dawson,

Terrell/Sumter Co’s.. UAUC 602, UAUC 603, UAUC

604; Kinchafoonee Creek at GA Rt. 32, Lee Co., GA:

MMNS._ 8096; Chickasawhatchee Creek at CR 130

bridge ~4 air mi. SW of Chickasawhatchee, Terrell Co.:

UAUC 1753; Muckalee Creek at GA Rt. 195 bridge

~3.5 air mi. Northeast of Leesburg, Lee Co.: UAUC

312; Whitewater Creek on Morgan Mill Rd., Fayette

Co.: UAUC 645; Ochlockonee River: DMNH 173390;

Ochlockonee River, 7 mi. S. of Cairo: ANSP 194640,

DMNH_ 048538, UF 412; Mimsville: ANSP 47892,

DMNH 075151; Georgia: ANSP 126273; Calvary: ANSP

47891; Ochlockonee River, Thomas/Grady Co's. Georgia:

MMNS S101; Spring Creek at GA Rt. 54, Decatur Co.:

MMNS 8094, MMNS S100; Line Creek at GA Rt. 85/

THE NAUTILUS, Vol. 119, No. 1

74 bridge, Coweta/Fayette Co.: MMNS 8097; Ichaway-

nochaway Creek at GA Rt. 216, Baker Co.: MMNS

SO9S.

Hamiota australis

ALABAMA

Andrews fish trap, Pea River, Barbour Co.: UF 65309;

7 mi. east of Brundidge, Pea River, Pike Co.: UF

123284; Bozemans landing, Conecuh River, near Cren-

shaw Co. line, Covington Co.: UF 65313; Lightwood

Knot Creek, 1.6 mi. west of Opp, Covington Co.:

ANSP#; Little Patsaliga Creek: USNM_ 150473 [Holo-

type, Lampsilis australis, Simpson, 1900]; West Fork

Choctawhatchee River at Blue Spring State Park, Bar-

bour Co.; UAUC 134, UAUC 511, UAUC 512, UAUC

513, UAUC 514: Little Choctawhatchie Creek, near

Drian bridge, Houston Co.: UF 229532; Conecuh River

on CR 28 ~1 mi. east of Goshen, Pike Co.: UAUC 510;

Flat Creek at AL Hwy 153, near Flat Creek Church,

Geneva Co.: UAUC 547: Five Runs Creek, Conecuh N.

F., Covington Co.: MMNS SOS6.

FLORIDA

Shoal River at Hwy 85, 1 mi. south of I-10 Jct. in Crest-

view, Okaloosa Co.: UVAUC 550, UAUC 551, VAUC 552.

UAUC 643, UAUC 644; Shoal Creek, ca. 1 mi. above

U.S. Highway 90, Okaloosa Co.: UF 261852; Limestone

Creek, Walton: MMNS 7746, MMNS 7747.

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Page 11

Consideration of genetic relationships in management decisions

for the endangered Anthony’s riversnail, Leptoxis Crassa

anthonyi (Redfield, 1854) (Gastropoda: Pleuroceridae)

Russell L. Minton

Museum of Natural History

University of Louisiana at Monroe

Monroe, LA 71209-0504 USA

[email protected]

Box 870345

Steven P. Savarese, Jr.

University of Alabama

Tuscaloosa, AL 35487 USA

ABSTRACT

Anthony’ riversnail, Leptoxis crassa anthonyi, is a federally en-

dangered pleurocerid restricted to three natural populations in

the Tennessee River drainage. Recovery plans organized the

three populations into two management units, and called for

specific numbers of populations for downlisting or delisting.

Given that nothing was known about the genetic structure of

these populations and that individuals were being randomly

transplanted, we examined each population using COI mtDN A

sequences. All three populations possessed unique sequence

haplotypes, and the two units identified in the recovery plan

did not group the populations accurately in a phylogenetic con-

text. Potential management decisions in light of our findings

are discussed.

INTRODUCTION

Anthonys riversnail, Leptoxis crassa anthonyi (Redfield,

1854), is a pleurocerid gastropod currently listed as en-

dangered by the United States Fish and Wildlife Service

(USFWS) under the End: mgered Species Act of 1973 as

amended (USFWS, 1994). Adult shells of L. crassa an-

thonyi are large compared to sympatric pleurocerids,

globose to ovate, greenish to greenish-brown in color,

often with purple bands. The body whorl may be sculp-

tured with low, indistinct tubercles, giving the she il a

bumpy appearance. The aperture is ovate with a thin

outer lip, and the columellar lip is fees d so that it

covers a prominent umbilicus (Tryon, LS7: 3). Historically,

L. crassa anthonyi had been documented aa the Ten-

nessee River drainage in Alabama, Georgia and Tennes-

see, including the lower reaches of some of the larger

tributaries (Burch and Tottenham, 1980; Bogan and Par-

malee, 1953). Most populations of the species were ex-

tirpated when much of the Tennessee River and its trib-

utaries were impounded by the Tennessee Valley Au-

thority. Natural populations of L. crassa anthonyi persist

at three localities: a small, scattered population in the

main channel of the Tennessee River near the Alabama

and Tennessee state line; a large population in Lime-

stone Creek, Limestone County, Alabama; and a small

population limited to a single stretch of the Sequatchie

River, Marion County, Tennessee (Garner, 1994; Jenkin-

son, 1994; USFWS, 1997; Figure 1).

Two morphologically similar snails have occurred

within the historic range of » Leptoxis crassa anthonyi.

The species often occurs sympatrically with L. praerosa

(Say, 1821), and adults of L. praerosa are easily confused

with juveniles of L. crassa anthonyi (Dillon and Ahls-

tedt, 1997). The other taxon, L. crassa crassa (Halde-

man, 1841), was described as being similar to L. crassa

anthonyi but with larger, more prominent tubercles on

the body whorl, is presumed extinct (Turgeon et al.,

1998). Leptoxis crassa anthonyi shells are most easily

recognized as juveniles, given their saucer shape and the

presence of a heavy carina that disappears with age (Dil-

lon and Ahlstedt, 1997). The distinctness of L. crassa

anthonyi has been supported by both allozyme (Dillon

and Ahlstedt, 1997) and mitochondrial sequence data

(Holznagel and Lydeard, 2000).

In 1997, USFWS published their recovery plan

(USFWS, 1997) for L. crassa anthonyi. The plan iden-

tified two “populations” of L. crassa anthonyi: individ-

uals from the Sequatchie and Tennessee Rivers as one

population; and individuals from Limestone Cre eek as the

other. Recovery criteria for L. crassa anthonyi were to

protect the extant populations and to successfully re-es-

tablish other populations. [f a total of four “poy sulations”

could be established, the species could be downlisted to

threatened status; if six “populations” could be estab-

lished, the species would be delisted entirely. Though

the plan treated the three localities as two populations,”

no effort had been made to determine if they were ge-

netically homogeneous (i.e., a single gene ee population )

or genetically variable ( two or three separate pop-

ulations). In order to oo the stated goals of the re-

covery plan, the number of genetically distinct Lt -

tions within the species needed to be determined.

working baseline of genetic information about each ae

ulation was even more important due to reports of snails

Page 12

THE NAUTILUS, Vol. 119, No. 1

Tennessee \ ~""G

a Alabama *&

Figure 1. Map showing the locations of the three extant nat-

ural populations of Leptoxis es anthonyi. L. = Limestone

Creek; S = Sequatchie River; T = Tennessee River.

being translocated into existing populations and moved

to new localities in the Tennessee River drainage (J. Gar-

ner and D. Hubbs, pers. comms.). In this study, we used

mitochondrial DNA sequence data to conduct a com-

parative genetic analysis of extant L. crassa anthonyi

populations to examine the degree of genetic differen-

tiation among the three populations. Knowledge of the

genetic structure would help direct management efforts

in determining which populations could and should

serve as sources for augmentations and reintroductions

if desired.

MATERIALS AND METHODS

Adult specimens of L. crassa anthonyi from the three

natural populations were collected under an endan-

gered/threatened species subpermit (SA99-13). The

maximum allowed number of specimens were collected,

twenty from Limestone Creek, and five each from the

Sequatchie and Tennessee Rivers (see Appendix 1

). For

Figure 2.

Leptoxis crassa anthonyi specimens identified as in Figure

position of Leptoxis and Lithasia clades are given in Table

3remer support >]

Strict consensus of two most parsimonious trees (TL =

L. Jackknife values >

|; both clades were monophyletic with jackknife support >50% and

comparative purposes, a reduced data set from previous

analyses (Minton and Lydeard, 2003) was used to place

L. crassa anthonyi in the proper systematic context. Fif-

teen individuals of L. crassa anthonyi from Limestone

Creek and five each from the other two populations

were included in the genetic study, along with four L.

praerosa, each from different river drainages, and one

each of the other taxa used in the previous study (Ap-

pendix 1).

Mitochondrial DNA sequences for a 1 kb portion of

the cytochrome oxidase subunit I gene (COT) were gen-

erated using published methods (Minton and Ly deacak

2003) for genomic isolation, PCR amplification, and ge-

netic an: alys sis that followed. Sequences were aligned i

eye (Hall, 1999) and phylogenetic hypotheses ¢ generated

by PAUP® 4.0b10 (Swofford, 2002) under maximum par-

simony with the following options: 50 replicates of heu-

ristic search with random addition, uninformative char-

acters were ignored, branches with minimum zero

length colle ipsed, and minimal length trees kept. This

gene fragment showed significant phylogenetic signal

and no base composition bias or sequence saturation in

a more inclusive pleurocerid dataset (Minton and Ly-

deard, 2003). Internal branch stability was assessed by

jackknife (Farris et al., 1996) and Bremer support (Bre-

mer, 1994).

RESULTS

Aligned sequences resulted in a data matrix of 890 char-

acters, of which 222 were parsimony-informative. Each

population of Leptoxis crassa anthonyi possessed a

unique haplotype, and all individuals from a population

shared the same haplotype. Maximum parsimony anal-

ysis yielded two trees (Figure 2, strict consensus). All

specimens of L. crassa anthony constituted a monophy-

letic group. Uncorrected p-distances were 1.46% be-

tween the Limestone Creek and Sequatchie River spec-

Pleurocera canaliculatum

Pleurocera walkeri

Pleurocera prasinatum

Elimia sp. 2

Elimia sp. 1

Elimia hydei

Lithasia

lo fluvialis

Leptoxis praerosa

Leptoxis c. anthonyi T (n=5)

Leptoxis c. anthonyi L (n=15)

Leptoxis c. anthonyi S (n=5)

71S, CL = 0.54) based on mitochondrial COL sequences.

50% above branches, Bremer values below. Com-

R. L. Minton and S. P. Savarese, Jr., 2005

imens, 2.02% between the Limestone Creek and Ten-

nessee River specimens, and 3.03% between the Se-

quatchie and Tennessee River specimens. The L. crassa

anthonyi clade was sister to a clade of L. praerosa spec-

imens. Most clades were well supported by jackknite and

Bremer values, as were the relationships within each

clade.

DISCUSSION

Our results further support the validity of Leptoxis cras-

sa anthonyi and its distinctiveness from L. praerosa, and

show each of the three populations of L. crassa anthonyi

to be genetically unique based on COT haplotypes. Un-

corrected pairwise genetic differences between the three

populations of L. crassa anthonyi were consistent with

published intraspecific differences seen in other pleu-

rocerids using COI sequences (Minton and Lydeard,

2003). The USFWS recovery plan calls for the presence

of no fewer than four viable populations of L. crassa

anthonyi before any change in listing status can occur.

For establishment of new “populations, introductions of

each haplotype to new areas within their current river

system would be preferred in order to avoid mixing of

unique evolutionary entities and possible elinninetion of

the current genetic diversity.

The USFWS has designated Leptoxis crassa anthonyi

from the Sequatchie and Tennessee Rviers as a single

population. Based on that assumption, their manage-

ment efforts might target one of the populations as a

source for augmenting the other. If translocations are to

be used, our fetal sis suggests that the Limestone Creek

population would be a better option for augmenting the

Sequatchie River population due to lower genetic dif-

ference. Additionally, if a single population is to serve as

a source of new introductions and augmentations, recent

survey work (J. Garner, pers. comm.) indicates that

Limestone Creek would be preferred because of its

large population size. Regardless, controlled experimen-

tal ‘populations should be established, either in the field

or laboratory, that could be monitored genetically and

for population ¢ growth and be compared with non-aug-

mented populations to determine whether fitness ne

been enhanced or diminished from the introduction of

unique haplotypes from other populations. This measure

would be especially prudent given that haplotypes are

being randomly introduced through human activity. Any

recovery plan that involves iencioosion of L. crassa an-

thonyi should use juveniles, as they are more easily iden-

tified by their strong keel. This will help ensure the

movement of L. crassa anthonyi and potentially prevent

the introduction of adult. non-endemic L. praerosa.

ACKNOWLEDGMENTS

This project was completed in the lab of C. Lydeard at

the University of Alabarna. We thank S. Ahlstedt and J.

Gamer for specimens. and two anonymous reviewers for

Page 13

their comments. Funding was provided by a USFWS

grant to C. Lydeard (1448-40181-97-G-033).

LITERATURE CITED

Bogan, A. E. and P. W. Parmalee, 1983, Tennessee’s Rare Wild-

life Volume II: The Mollusks. Tennessee Wildlife Re-

sources Agency, Nashville, 123 pp.

Bremer, K. 1994. Branch support and tree stability. Cladistics

10: 295-304.

Burch, J. B. and J. L. Tottenham. 1980. North American fresh-

water snails. Species list, ranges, and illustrations. Walk-

erana |: S]—215.

Dillon, R. T. and S. A. Ahlstedt. 1997. Verification of the spe-

cific status of the endangered Anthony’s riversnail, Athear-

nia anthonyi, using allozyme electrophoresis. Nautilus 10:

97-101.

Farris, J. S.. V. A. Albert, M. Kallersjo, D. Lipscomb and A.

G. Kluge. 1996. Parsimony jackknifing outperforms neigh-

bor-joining. Cladistics 12: 99-124.

Gamer, J. 1994. Survey of mollusks: Tennessee River mile

412.1. Aquatic Resources Center, Franklin, 2 pp:

Jenkinson, J. 1994. Freshwater mollusk survey at CSX railroad

bridge, near Bridgeport, Alabama, Tennessee River Mile

414.5. Tennessee Valley Authority, Chattanooga, 15 pp.

Haldeman, S. S. 1841. A monograph of the Limnaides and

other freshwater univalve snails of North America. No. 3,

[{Limnea]. J. Dobson, Philadelphia, 16 pp.

Hall, T. A. 1999. BioEdit: a user-friendly biological sequence

alignment editor and analysis program for Windows 95/

9S/NT. Nucleic Acids Symposium Series 41: 95-98.

Holznagel, W. E. and C. Lydeard. 2000. A molecular phylogeny

of North American Pleuroceridae ( (Gastropoda: Cerithioi-

dea) based on mitochondrial 16S rDNA sequences. Jour-

nal of Molluscan Studies 66; 233-257.

Minton, R. L. and C. Lydeard. 2003. Phylogeny, taxonomy,

genetics, and global heritage ranks of an imperiled, fresh-

water snail genus Lithasia (Pleuroceridae). Molecular

Ecology 12: 75-87.

Redfield, J. H. 1854. Descriptions of new species of shells.

Annual Report of the Lyceum of Natural History of New

York 6: 130-132

Say, T. 1821. Descriptions of the univalve shells of the United

States. Journal of the Academy of Natural Sciences of

Philadelphia 2: 149-179.

Swofford, D. 2002. PAUP®: Phylogenetics Analysis Using Par-

simony, version 4.0b10. Sinauer Associates, Sunderland,

Massachusetts.

Tryon, G. W. 1873. Land and freshwater shells of North Amer-

ica, Part IV. Strepomatidae. Smithsonian Miscellaneous

Collections 253: i-iv, 1-435.

Turgeon, D. D., J. F. Quinn, A. E. Bogan, E. V. Coan, F. G.

Hochberg, is G. Lyons, P. M. Mikkelsen, R. J. Neves, C.

F. E. Roper, }. Rosenberg, B. Roth, A. Schletema, F. G.

Thompson, “4 Vecchione and G. D. Williams. 1998.

Common and scientific names of aquatic invertebrates

from the United States and Canada: Mollusks (second edi-

tion). American Fisheries Society Special Publication 26,

Bethesda, 526 pp.

United States Fish and Wildlife Service. 1994. Endangered and

threatened wildlife and plants; determination of endan-

gered status for the royal snail and Anthony’ riversnail

Federal Register 59: 17 ‘994-17998.

United States Fish and Wildlife Service. 1997. Recovery plan

for Anthony’s riversnail. Atlanta, Georgia. 21 pp.

Page 14

Appendix 1. Systematic list of taxa and specimens used in this study. Complete locality information is available from the authors.

UAG = University of Alabama gastropod collection.

Taxon

Locality

Genus Elimia

E. hydei

Elimia sp. 1

Elimia sp. 2

Genus Io

Io fluvialis

Genus Leptoxis

L. crassa anthonyi

L. praerosa

Genus Lithasia

L. armigera

L. geniculata fuliginosa

L. lima

L. verrucosa

Genus Pleurocera

P. canaliculatum

P. prasinatum

P. walkeri

Black Warrior River, Tuscaloosa Co., AL

Green River, Hart Co., KY

Collins River, Warren Co., TN

Clinch River, Hancock Co., TN

Limestone Creek, Limestone Co., AL

Sequatchie River, Marion Co., TN

Tennessee River, Jackson Co., AL

Harpeth River, Davidson Co., TN

Shoal Creek, Lawrence Co., AL

Sequatchie River, Marion Co., TN

Tennessee River, Jackson Co., AL

Harpeth River, Cheatham Co., TN

Red River, Robertson Co., TN

Duck River, Maury Co., TN

Buffolo River, Humphreys Co., TN

Bear Creek, Colbert Co., AL

French Broad River, Knox Co., TN

White River, Woodruff Co., AR

Duck River, Maury Co., TN

Yellowleaf Creek, Shelby Co., AL

Shoal Creek, Lauderdale Co., AL

dCadaddddaada

UAG voucher

AG 584

AG 574

UAG 407

AG 585

UAG 581

AG 582

AG 583

AG 404

AG 560

UAG 588

AG 589

AG 555

AG 398

AG 403

AG 406

AG 570

AG 576

AG 577

AG 590

AG 591

AG 592

THE NAUTILUS, Vol. 119, No. 1

Genbank accession

AF435775

AF435759

AF435761

AF435776

AF435772

AF435773

AF435774

AF435779

AF435780

AF435781

AF435782

AF435739

AF 435754

AF435749

AF435747

AF435767

AF43577

THE NAUTILUS 119(1):15-26, 2005

Page 15

Fallen into oblivion—the systematic affinities of the enigmatic

Sulcospira Troschel, 1858 (Cerithioidea: Pachychilidae), a genus

of viviparous freshwater gastropods from Java

Frank Kohler!

Matthias Glaubrecht’

Museum of Natural History

Humboldt University

Institute of Systematic Zoology

Department of - o 1cozoology

InvalidenstraBe 43, D-10115 Berlin

GERMANY

' frank. [email protected]

> [email protected]

ABSTRACT

Sulcospira Troschel, 1858, is not only the taxonomically oldest

but also one of the most poorly known genera of Southeast

Asian Pachychilidae. It serves as an instructive case study as to

how the puzzling systematics of freshwater Cerithioidea has

hampered a deeper understanding of their phylogeny and evo-

lution. The genus has been established for the Javan freshwater

gastropod Melania sulcospira Mousson, 1849, on the grounds

of its round, multispiral operculum and an elongated main cusp

in the central radula teeth. Although of great systematic sig-

nificance, this taxon has been widely ignored by subsequent

authors. We here recapitulate the taxonomic history of the ge-

nus and describe and evaluate the morphological properties of

the type species S. sulcospira on basis of the limited existing

material, in order to facilitate a better understanding of pachy-

chilid systematics. In addition, in an attempt to clarify its sys-

tematic affinity, we compare the properties of another allege sly

related species from Java, S. martini (Schepmann, 198). We

show that these two viviparous species exhibit different pro-

toconch morphologies, which are indicative of reproductive

strategies distinct from other pachychilids. Finally, we outline

preliminary suggestions as to the systematics of Sulcospira

within the family Pachychilidae.

MATERIALS AND METHODS

MATERIALS

This study is based on the examination of material from

various museum collections worldwide (see repositories).

All of these samples comprise dry shells only, which as

a rule were empty. Only few shells contained fragmen-

tary soft parts; some of them were re-hydrated for ex-

aminations. However, these bodies gene rally did not fa-

cilitate morphological examinations except for the ex-

traction of small radula fragments. We have not tried to

extract DNA from dried tissues because earlier attempts

with comparable material of Brotia failed. In order to

acquire fresh material, collecting trips have been under-

taken in 2000 and 2002. We have searched rivers and

creeks in different sectors of there course (i.e. upstream,

midstream, downstream) for a period of altogether 7

days in West Java (along the roads between Jakarta and

Serang, Bogor and Sulabtme Sukabumi and Pelabuhan

Ratu, Bogor and Cipanas, Bogor and Cianjur, in the Bo-

tanical Garden Bogor) and for 3 days in East Java (be-

tween Taksimalaya, Cipatujah, and Pangadaran). During

these trips we were not able to find any mate ial of Sul-

cospira, though.

Because freshwater biotopes on Java are facing dra-

matic devastation by a multitude of causes related to the

dense population on this island such as pollution, flow

regulation, drainage, impoundment and a general deg-

radation of collecting areas by agriculture, industry and

settlements (own observations; see also Dudgeon, 2000,

for SE Asia in general), we believe that S. sulcospira has

become extinct in vast areas on Java. It remains unclear

whether and at which localities populations of this spe-

cies still exist.

Reposirory INSTITUTIONS

Voucher material is housed with the following museums:

Natural History Museum, London (BMNH), Museum

of Comparative Zoology, Cambridge, Mass. (MCZ), Mu-

séum cl Histoire Naturelle, € Geneve (MHNG), Muséum

National d'Histoire Naturelle, Paris (MNHN), Natural

History Museum Naturalis, Leiden (RMNH), Sencken-

bergmuseum, Frankfurt/Main (SMF), Zodlogisch Mu-

seum, Amsterdam (ZMA), Museum fiir Naturkunde,

Berlin (ZMB).

We were not able to locate material in the following

museum collections: Academy of Natural Sciences, Phil-

Page 16

THE NAUTILUS, Vol. 119, No. 1

adelphia (ANSP), United States National Museum,

Washington (UNSM), Zoologisches Institut und Muse-

um, Universitiit Hamburg (ZMH), Zoologische Staats-

sammlung, Miinchen (ZSM).

MORPHOLOGICAL EXAMINATIONS

Dimensions of all shells measured to 0.1 mm pre-

cision. The shell height (H) is the maximum dimension

parallel to the axis of ae breadth (B) the maximum

dimension perpendicular to H, including the aperture.

The length of the aperture (LA) is the greatest length

from fhe: junction of the outer lip with the penultimate

whorl to the anterior lip, the width (WA) the greatest

length perpendicular to LA. The height of the body

whorl (BW) is the distance from the ase of the shell to

the upper suture of the first whorl exactly above the

junction of the outer lip with the penultimate whorl.

Morphometrical parameters used in the analyses, beside

the shell dimensions, were: H/B, H/LA, H/BW, H/LA

and B/BW. These shell parameters were statistically an-

alyzed by performing t-tests, one-way ANOVA, and a

discriminate analysis.

Protoconchs removed from dried adults were cleaned

by soaking in 10% KOH solution, flushed in distilled

water, and sonicated to remove residual contaminations

prior to scanning electron microscopy. Radulae were tak-

en from dried shells or from historic preparations. Rad-

ulae from dried shells were enzymatically cleaned as de-

scribed by Holznagel (1998); an old radula embedded in

Canada balsam was cleaned with xylene followed by son-

ication. Radulae and juvenile shells were mounted on

aluminum specimen stubs using adhesive carbon tabs or

double-sided tape, respectiv ely. and coated with gold-

palladium for 120 s at 20 mA for examination under a

scanning electron microscope (LEO 1450 VP) at 10 keV.

NOMENCLATORIAL REMARKS

Some species-group names introduced by Troschel

(1857-1555) are open to discussion. Bouchet (pers.

comm,) argued that the usage of the names Bithyniae,

Lithoglyphi, Hydrobiae, Ancyloti, Thiarae, and Pachy-

chili by Troschel ( (op. cit.) contrasts with the rest of his

work (Troschel, 1856-1863), in which he stated the

ranks of the categories he used and formed names with

endings -idea, -ina, or -acea. Because Troschel stated

explicitly that he refrained from allocating these group-

ings at family rank given the somewhat ambiguous mor-

phological data he was faced with, it was suggested that

one should ignore these names (Bouchet, pers. comm.).

However, some of these names, such as Bithyniidae,

Thiaridae, or Hydrobiidae, have been usually published

with Troschel (1857-1858) as author.

stipulated, we prefer to refer to Troschel (1857-1858) as

Unless otherwise

author of these names not only because we regard them

as available and valid but also in order to maintain sta-

bility in their usage; for a statement to the contrary see

Bouchet and Rocroi (submitted).

Figure 1. Original drawing of the radula of “Sulcospira typ-

. (Melania sulcospira Mousson)” by Troschel (1858: pl. 9, fig.

. In the upper part of the figure a row of teeth is shown

ae a rachidian flanked on each side by a lateral tooth

and an inner and outer marginal tooth; a magnified represen-

tation of the rachidian is shown below. Characteristics that are

typical for Pachychilidae are, e.g., the enlarged main cusp of

the rachidian and lateral teeth, respectively, which is flanked

by three (or two) smaller cusps on each side that taper in size;

the presence and shape of the glabella (or ramp); and marginal

teeth possessing two cusps.

Nomenclatorial aspects raised in this paper refer to

the stipulations of the 4 edition of the International

Code of Zoological Nomenclature (“ICZN”) issued by

the Intern: eonal Commission of Zoological Nomencla-

ture (1999).

RESULTS

Sulcospira Troschel, 1858

Sulcospira Troschel, 1858: 117-118; Brot, 1874: 56; Thiele,

1929: 190; Morrison, 1954: 351.

Diagnosis: —Sulcospira ea a rather conical shell

et by spiral lirae; axial sculptural elements are

lacking. Protoconchs possess a smooth sculpture with a

fine granular texture or faint growth lines.

Type Species: 1849, by

mo ytypy.

Melania sulcospira Mousson,

Nomenclature and Systematics: The genus Sulcos-

pira was described by Troschel (1858) for the Javan spe-

cies M. sulcospira e xhibiting certain characteristics that

were held to be peculiar of his species, namely a round,

multispiral operculum with four regular whorls and the

radula with an enlarged main cusp of the rachidian (Fig-

ure 1, 17-18). Troschel (1858: 114) based his description

on material received from August Brot in Genéve, i.e.,

F. Kohler and M. Glaubrecht, 2005

Figure 2. Shell dimensions. B: breadth; BW: weight of the

body whorl; H: height; LA: length of the aperture; WA: width

of the aperture.

on material that is likely housed at MHNG today (see

Material Examined).

Sulcospira represents the oldest available generic

name established for representatives of the Southeast

Asian Pachychilidae and is here considered valid. How-

ever, the diagnosis of Troschel (1858) is not sufficient to

characterize the taxon unambiguously. Neither a round

and multispiral operculum w ith four whorls nor the pos-

session of a pronounced main cusp of the rachidian are

considered as diagnostic features of Sulcospira alone

(see discussion). Nonetheless, these features character-

ize Sulcospira as a member of the Pachychilidae (Glau-

brecht, 1996, 1999: Kohler and Glaubrecht, 2001, 2002,

2003).

In more recent literature Sulcospira has been widely

ignored. Thiele (1929) suggested subdividing Sulcospira

into two subgenera, Sulcospira and Tylomelania F. and

P. Sarasin, 1S98. According to Thiele, Sulcospira would

include Paracrostoma, Acrostoma Brot, 1870, and Bro-

tella Rovereto, 1899, as ee synonyms, consequently

comprising two species: S. sulcospira from Java and S.

Page 17

huegeli (Philippi, 1843) from South India. Tylomelania,

however, was considered to encompass a small number

of species restricted to Sulawesi.

This concept of Thiele (1929) led Subba Rao (1989:

107) to wrongly assume that M. huegeli would be the

type species of Sulcospira, which it is not (see Troschel’s

original designation).

Later authors had different taxonomic views. Morri-

son (1954) followed Abbott (1948) but not Thiele (1929)

and treated Acrostoma, Brotella, and Paracrostoma as

synonyms of Brotia, while considering Tylomelania as a

genus on its own. He also claimed that our understand-

ing of Sulcospira is not satisfactory. Based on some su-

perficial similarities with Brotia, but also with Tylome-

a and Balanocochlis (a thiarid), Morrison (1954) pro-

posed the allocation of Sulcospira “tentatively to the Me-

lanoides complex”. Since Melanoides is a thiarid, this

allocation is rejected here.

All a classification schemes were suggested in ab-

sence of phylogenetic analyses of morphological char-

acters. In addition to the type species, other taxa have

been assigned to the genus by previous authors, al-

though this has not hee m done consistently (Table 1).

The various opinions led also to different a dp

on the species circumscription and div ersity of this tax-

on. For example, Brot (1874) subsumed a number of

taxa under Sulcospira, of which we currently only con-

sider two to be actually pachychilids: Mc lania spadicea

Reeve, 1860, and M. hainanensis Brot, 1872. Yen (1939)

added two more taxa, M. ebenina Brot, 1883, and M.

biconica Brot, 1886. Boettger (1890), Oostingh (1932)

and Adam and Leloup (1838) treated M. te eeudnaria

von dem Busch, 1842, as member of Sulcospira but did

not mention the former taxa. In contrast, other authors

assigned those taxa to Brotia instead (Rensch, 1934;

Benthem: -Jutting, 1956, 1959; Knipper, 1958; Dudgeon,

1982, 1989: Kéhler and Glaubrecht, 2001, 2002). Even-

tually, Benthem-Jutting (1956) assumed that S. sulcos-

pira is the only representative of the genus.

In order to clarify the puzzling taxonomy and system-

atics, a revision of Sulcospira with an evaluation of its

anatomical characters is needed. A sound classification

has to be based on autapomorphic features, which is

lacking to date. Unfortunately, a comprehensive descrip-

tion of the morphology of Sulcospira suffers from the

Table 1. Comparison of previous views on the systematics and circumscription of Sulcospira

Authors

Morrison (1954);

Benthem-Jutting Kohler and Glaubrecht

Brot (1874) Thiele (1929) (1956) (2002)

Taxonomy at generic Melania (Sulcospira)

Sulcospira (Sulcospira),

Sulcospira Sulcospira

level Sulcospira (Tylomelania)

Included taxa M. sulcospira, M. spadicea,

M. hainanensis, and other

non-pachychilid taxa

sulcospira, S. huegeli,

T. neritiformis, T. carbo,

T. porcellanica

S. sulcospira S. sulcospira, 8. spadicea

Page 18

THE NAUTILUS, Vol. 119, No. 1

Table 2. Shell parameters [mm] of S. sulcospira and S. martini. Abbreviations: B: breadth; BW: weight of the body whorl; H:

height: LA: length of the aperture; m: median; N: number of whorls; No: number of shells; sd: st: sitar deviation; WA: width of

the aperture.

Lot No

S. sulcospira (total) m 26

Holotype M. sulcospira ]

S. sulcospira (ZMA) m 21

S. sulcospira (MNHN) m 4

S. martini (total) m 58

Syntypes M. spadicea (BMNH 19990497) m 3

Syntypes M. junghuhni (RMNH, ZMA) m 22

Sym pes var. flammulata, var. fasciata m 16

RMNH) sd

5 eee (ZMB 4.074) m 17

Oo) x

io)

aa ee bo

Olde WO mNWowns

NUMA ONUW Ui &

B LA WA BW N

10.1 9.5 4.9 14.3 4.7

1.3 1.0 0.6 1.9 0.5

12.0 9.7 5.0 15.2 4.0

10.2 9.5 4.9 14.4 4.6

1:2 0.9 0.5 0.6 0.6

9.9 9.3 4.9 14.1 4.8

1.8 L.5 0.9 2.9 0.4

12.0 11.0 5.9 17.7 6.0

1.5 1.3 0.8 2.1 1.0

11.6 10.2 5.2 16.1 6.0

0.2 0.9 0.6 0.9 0.5

12.4 11.6 6.0 18.4 6

1.8 1.4 0.9 2.4 0.7

12.3 11.7 6.1 17.9 6.6

1.2 11 0.7 1.7 0.6

11.2 11.0 5.7 16.9 4.9

1.3 1.0 0.5 hod 0.7

lack of well-preserved material of the type species. We

compile below the anatomical data based on the material

of S. sulcospira available.

Sulcospira sulcospira (Mousson, 1849)

Melania sulcospira Mousson, 1849a [1848]: 269; Mousson,

1S49b; 68, pl. 9, fig. 3; Martens, 1897; 245 (partim); Les-

chke, 1914: 251.

Sulcospira typica Troschel, 1858: 117, 118, pl. 9, fig. 6 [intro-

duced as replacement name for M. sulcospira Mousson,

1849].

Pachycheilus sulcospira |sic|—H. and A. Adams, 1858; 299.

Melania ( (Sulcospira) pee ae —Brot, 1870: 277: Brot, 1874:

56-57, pl. 6, fig. 11: Boettger, 1890; 245.

Sulcospira sulcospira ne 1954: 381; Kohler and Glau-

brecht, 2002: 149, fig. 3 L.

Diagnosis: Shell relatively small (Table 2), conical

with spiral lirae, and a subsutural depression; aperture

elongately ovate, abapically flared. Radula with squarish

rachidian exhibiting a straight upper and lower rim, the

cutting edge of all teeth with one very pronounced main

cusp of triangular shape.

Description: Shell (Figures 3-5): small, ovate to

conical, solid, spire with eroded apex and up to six flat-

tened whorls, separated by a narrow suture; sculpture

consisting of fine, reguli ar spiral lirae that are most prom-

inent at the base aad may almost lack on upper whorls,

and faint growth lines; whorls with a subsutural de spres-

sion; color from yellowish to olive or dark brown. Ap-

erture elongate ‘ly ovate, abapically flared, peristome

sharp; columella ‘slightly bent and thickened.

Protoconch (Figures 19-20): Relatively large, com-

pared to the adult: with he ight of about 1.2 mm com-

prising one and a half whorl: generally smooth. Apical

whorl inflated, dome-shaped, with a granular surface

sculpture, transition in sculpture visible on first whorl

from granular to faint growth lines. One sample of four

specimens housed at MNHN (ex coll. Staat) has a label

stating that * ‘about 140 embryos were obtained from the

large specimen”. However, the fate of these protoconchs

is unknown.

Operculum: With four whorls regularly increasing in

diameter and a sub-central nucleus.

Radula (Figures 16-17): Rachidian tooth squarish with

a straight upper and lower rim, cutting edge with one

he: avily enlarged main cusp of triangular shape, flanked

by two, simak smaller accessory cusps on each side; gla-

bella well developed, rather rectangular with a rounded

basal margin not exceeding the lower rim of the rachi-

dian tooth, lateral margins straight and not well defined.

Lateral teeth with one heavily enlarged main cusp, and

considerably smaller accessory cusps, one at the outer

side, two or three at the inner side. Inner and outer

marginal teeth with two cusps, the outer one being very

large, triangular in shape, and the inner one being point-

ed and small. Inner marginal teeth broader than outer

marginals. Outer lateral flange inconspicuous.

Unknown.

Type Material: Indonesia: Holotype ZMZ 522306, In-

donesia: Java, leg. Zollinger (Figure 3) {Mousson, 1849b

refers to “das einzige Exemplar dieser Art...” = the

only specimen of this species . . .].

Anatomy:

Type Locality: “Java”, Indonesia.

Other Material Examined: Indonesia: Java (MHNG;

MNHN: MNHN, ex coll. Staat; ZMA (2 lots); ZMB

200.101) (if not stated otherwise, a single lot from each

collection, mostly without reference number, was ex-

amined): museums without material: ANSP, BMNH,

MCZ, RMNH, SMF, USNM, ZMH, ZSM.

F. Kohler and M. Glaubrecht, 2005 Page 19

Figures 3-16. Shell morphology of Sulcospira sulcospira (3-5) and S. martini (6-16) (apertural and abapertural, respectively

3. S. sulcospira. Holotype (ZMZ 522306). 4-5. S. martini. Two shells from Java (ZMA). 6. Lectotype of M. spadicea (BMNH

19990497/A). 7-8. Two paralectotypes of M. spadicea (BMNH_ 19990497/B). 9. Lectotype of M. junghuhni (RMNH 71326). 10-

11. Two syntypes of M. junghuhni var. flammulata (RMNH 71327). 12-13. Two syntypes of M. junghuhni var. fusciata (RMINH

7132S). 14-16. Three shells from Java. Malangbong (ZMB 4.074). Scale bar = 10 mm

THE NAUTILUS, Vol. 119, No. 1

Figures 17-24. SEM images of the radula and juvenile shells removed from dry shells of S. sulcospira (17-20) and S. martini

(21-24). S. sulcospira: 17. Radula (ZMA). 18. Radula (ZMB 200.101). 19. Juvenile shell, apertural view (ZMA). S. martini: 20.

Juvenile shell, apical view (ZMA). 21. Juvenile shell, apertural view (ZMB 4.074). 22-23. Apical view. 24. Detail of the apical

portion. Scale bars = LOO zm.

F. Kohler and M. Glaubrecht, 2005

18 : —_— = a ]

}

167 o 50

° 2 |

4 |

14 oO

Co}

oo «Kod |

m 12 Ke 8

OR XK, & xPO |

oO ax x § e °

10 o ae oS |O'S. sulcospira |

aos |

| Oo a S. sulcospira holotype |

8 | 5 a =: |@ S. spadicea types —

| OS. martini types |

: XS martini (2B 4074) | |

10 15 20 25 30 35 40 45

Figure 25. Comparison of the shells of $. sulcospira and _S.

martini based on shell parameters height (H) and breadth (B).

Distribution: Indonesia: Java. More precise localities

were given by Boettger (1890: Bogor Botanical Garden)

and Martens (1997: Jakarta, Malangbong, Cipanas).

However, the hase of Boettger sould not be traced

(SMF? ), and Martens misidentified material of Melan-

oides tuberculata (Cipanas) and S. martini (Malangbong;

ZMB 4.076); no voucher material was found from Ja-

karta.

Nomenclature and Systematics: Melania sulcospira

is the type species of the genus Sulcospira by original

designation. When describing the new genus, Troschel

(1858), mentioned M. sulcospira as the * ‘typical species”,

and introduced the name S. typica for the same taxon

as a new, unnecessary substitute name meant to replace

an older available name (nomen novum). Thus, S. typica

is a junior svnonym of S. sulcospira (ICZN Art. 72.7.).

Sulcospira ee is invalidated as potential type name

(ICZN Art. 68.4.) making M. sulcospira the type by ab-

solute autonomy. Martens (1897) ued that M. spad-

icea is a synonym of this species, which was rejected by

Leschke (1914) and Benthem-Jutting (1956: 373), stat-

ing that Martens (1897) had misidentified specimens of

Melanoides tuberculata for M. spadicea.

Remarks: Compared to Sulcospira martini, S. sulcos-

pira is more conical in shape; the former lacks a con-

spicuous subsutural depression. Most conspicuously,

both species differ in their protoconch morphology.

Shells of B. testudinaria are more elongated in shape,

attain a larger size (between 25 to 40 mm in shell

height), lack a subsutural depression, and exhibit a dif-

ferent radular morphology: e.g., rachidian with inflated

and rounded upper corners, a smaller main cusp, mar-

ginal teeth possess two equally shaped cusps.

Given the scarcity of material and imprecise earlier

locality data for this material, we tried to restrict the type

locality from historical accounts. The Swiss malacologist

Albert Mousson (1805-1890) based his de scriptions on

material collected by the Swiss botanist Heinrich Zollin-

ger (1818-1859), who traveled in Indonesia between

1842 and 1848. However, as Zollinger collected not only

Page 2]

2.8

2.6

2.4

2.2)

N= 25 59

S. sulcospira S. martini

Figure 26. Comparison of S. sulcospira and S. martini based

on shell parameters ratio H/B. Box plot diagram showing the

median, the 25%- and 75%-percentile and largest non-ex-

tremes (less than 1.5 times of box height).

in West Java, but later also in East Java, and on some

other islands (Wanner, 1984), it turned out that a re-

striction of the type locality is not possible and “Java”

remains as the only known reference.

Sulcospira(? ) martini (Schepmann, 1898)

Melania spadicea Reeve, 1860: pl. 19, species 132 (not Melania

spadicea Philippi, 1549); Brot, 1S70: 277.

Melania (Sulcospira) spadicea.—Brot, 1874: 57-58, pl. 6, fig.

12,

Brotia spadicea.—Benthem-futting, 1956: 372-373, fig. 75

Sulcospira spadicea.—kohler and Glaubrecht, 2002: 14s. fig.

3G.

Melania junghuhni Schepman, 1896; 135-136, pl. 2, fig. 1.

(“Java”; lectotype and 41 paralectotypes RMNH 71326; 24

paralectotypes ZMA; two paralectotypes SMF 292406; var.

flammulata: 16 syntypes RMNH 71327; 3 syntypes MCZ

96926, var. fasciata: 18 syntypes RMNH_ 71528; 24 syn-

types ZMA; § syntypes MCZ 96598) (not M. junghuhni

Martin, 1879); Leschke, 1914; 251; Benthem-Jutting,

1929: S4.

Melania martini Schepmann, 1898: S4.

Brotia testudinaria.—Kohler and Glaubrecht, 2001; 301-304

(partim); Kohler and Glaubrecht, 2002: 140, 141, 150

(pe artim).

Diagnosis: Conical shell with convex to flattened

whorls sculptured with fine spiral ridges. Protoconch

with about 2.5 regular whorls; apical whorl not inflated,

no transition in shell structure is visible in the first two

whorls. Axial ribs may be present in the juvenile shell

from the second whorl on. Most conspicuously distin-

guished from all other pachychilids by its oe

morphology ( (Figures 27-34): distinguishable f rom. §.

sulcospira by its more elongated and I. irger shell (Figure

26); though adult shell not distinguish: ible from B. tes-

THE NAUTILUS, Vol. 119, No. 1

Figures 27-34. Comparison of protoconchs of different southeastern Asian Pachy chilidae (apertural and apical view, respectively).

27-28. Brotia costula. 29-30. Brotia hainanensis. 31-32. Tylomelania patriarchalis (with kind permission of Thomas von Rintelen).

33-34. Pseudopotamis semoni. Scale bars = 100 jum.

Desenptons Shell (Figures 6-16): Small to medium

sized (Table 2), ovate to conical, spire with eroded apex

and eight to ten flattened whorls; sculpture consisting of

fine, closely spaced regular spiral lirae, may lack ahniost

completely, and faint gr rrowth lines: color yellowish brown

to olive, brown spiral band or patches may be present.

Aperture elongated ovate, produced below, peristome

sharp.

Protoconch (Figures 21-24): Height of about 1.2 mm

comprising 2% whorls; apical wheel not inflated, corre-

sponding to the regular diameter of the subsequent

whorls; first two w honk smooth, only faint growth lines

visible, without transition in sculpture, our the second

whorl on smooth axial ribs may be present.

Operculum: Consisting of three whorls and a sub-basal

nucleus Brot (1874).

Anatomy and Radula: Unknown.

Type Material: Indonesia, Java: els and 41

paralectotypes of M. junghuhni, RMNH 71326, leg. Jun-

ghuhn (Figure 9), designated by Kohler and G laubrecht

(2002): 24 paralectotypes, ZMA; two paralectotypes,

SMF 292406; 16 syntypes of M. junghuhni var. flam-

mulata, RMNH 71327; three syntypes, MCZ 96926, 18

syntypes of M. junghuhni var. fasciata, RMNH 71328;

eight syntypes, MCZ 96898. Without locality: Lectotype

of M. spadicea, BMNH 19990497/A, Cuming collection,

anal by Kohler and Glaubrecht (2002): two para-

lectotypes, BMNH_ 19990497/2.

Type Locality: “Java”, Indonesia.

Other Material Examined: Without locality (ZMA;

MHNG): Indonesia: Java (MHNG), Malangbon (ZMB

4.074) (a single lot from each collection, mostly without

reference antaibee No material was found in the fol-

lowing museums: ANSP, MNHN, SMF. USNM, ZMH,

ZSM.

Nomenclature and Systematics: Melania spadicea

Reeve, 1860, is a primary homonym of M. spadicea Phi-

lippi, 1849 (ICZN Art. 53.3) and, as such, is permanently

invalid (ICZN Art. 57.2). The next available names are

M. junghuhni Schepmann, 1896, M. junghuhni var. flam-

F. Kohler and M. Glaubrecht, 2005

mulata Schepmann, 1896, and M. junghuhni var. fasciata

Schepmann, 1896. However, the first is a primary hom-

onym of Melania junghuhni Martin, 1879, a fossil spe-

cies from Java, as stated by Schepmann (1898). The oth-

er two are junior primary synonyms of Melania flam-

mulata von dem Busch in Philippi, 1843, and Melania

fasciata Menke, 1828, respectively.

Schepmann (1898) suggested Melania martini as a re-

placement name for M. junghuhni, which is the valid

name for this taxon. The two color morphs described by

Schepmann (1896), fasciata and flammulata, are not

considered here to represent extant evolutionary entities

and therefore are treated as junior synonyms of M. mar-

tini.

The classification of this species by former authors is

inconsistent. Benthem-Jutting (1956) considered it to be

a member of Brotia; but unaware of the fact that M.

spadicea Reeve, 1860, is not valid she assumed that this

name had priority over M. martini. Kohler and Glau-

brecht (2002) assumed that M. martini and M. spadicea

are distinct and treated the former as a synonym of B.

testudinaria and the latter as a species closely related to

S. sulcospira as was suggested earlier by Brot (1874).

Distribution: Java, as the only known locality. Ma-

langbon in Central Java, east of Bandung, is the only

known exact locality (ZMB 4.074, catalogued in 1859).

Analyses of Shell Morphometry: —Sulcospira martini

can be distinguished from S. sulcospira by its higher

shell and more slender shape (see analyses of she ll pa-

rameters below). However, to differentiate between

shells of “M. sulcospira” and “M. martini” is no easy task;

contradictory statements on their taxonomy abound in

earlier accounts (Brot, 1874; Benthem-Jutting, 1956;

kohler and Glaubrecht, 2002). In fact, shells exhibit a

very similar shape, sculpture, and coloration, However,

the two taxa can be distinguished by statistical analyses

of shell morphometry. We Taséd one-way ANOVA and t-

test for two independent groups of var iables to discrim-

inate specimens that were assigned beforehand either to

S. sulcospira or to S. martini according to their shell

oe The t-test showed that both taxa vary sig-

nificantly by the following parameters (P<5%): H, N,

H/B, H/LA, and H/BW: the one -way-ANOVA yielded

corresponding results.

The shells of the only lot from Java with precise lo-

cality data (ZMB 4.074) is identified here as S. martini

given its elongated shell (while the original label states

“M. sulcospira” instead). The statistical test has been

employed to explore whether these shells can signifi-

cantly be discriminated either from shells of S. sulcos-

pira or S. martini in regard to shell morphometry. Com-

parison of shells of the lot ZMB 4.074 with shells of S.

sulcospira (MNHN, ZMA, ZMB 200.101, ZMZ 522306)

by t-test reveals that both groups differ significantly in

the parameters H/LA, H/BW, and with a weak support

(P = 0.51) for H/B, whereas no significant differences

were found when comparing the lot ZMB 4.074 with

Page 23

Table 3. Results of the discriminate analysis of shell param-

eters.

Predicted group

membership S. sulcospira S. martini

S. sulcospira 95 (100.0%) 0 (0.0%)

S. martini 1 (1.7%) 59 (98.3%)

other shells of S. martini (BMNH 19990497, RMNH

71326-8).

A graphic comparison of the two taxa by means of

selected shell parameters is shown in Figures 95—26.

Shells assigned to each of the two taxa according to their

morphology were found to be correctly dlascified by a

diserantuate analysis of morphometric data with good

statistical support (Table 3). It is concluded that challe

of S. sulcospira are smaller and more conical in shape

than shells of S. martini (Table 1, Figure 26).

DISCUSSION

I. EVALUATION OF MORPHOLOGICAL CHARACTERS OF

SULCOSPIRA

The operculum and radula of S. sulcospira led Troschel

(1858) to describe a new genus for this species. How-

ever, among the Pachychilidae the operculum is known

to be relatively conservative in its general organization

(that is, to be. multispiral, rounded or ovate) bt quite

variable in relation to their number of whorls and in-

crease in diameter, even within a single genus (Kohler

and Glaubrecht, 2001, for Brotia; Kohler and Glau-

brecht, 2003, for Jagora; Glaubrecht and Rintelen, 2003,

for Pseudopotamis; Rintelen, 2003, for Tylomelania;

Kohler, 2003). Consequently, an operculum possessing

four regular whorls might be typical for S. sulcospira,

but only at the species les el. The possession of a round

to oval. multispiral operculum led Sarasin and Sarasin

(1898) to group several taxa within the so-called “pa-

laeomelanians”, as contrasted with the so-called “neo-

melanians”, which exhibit a paucispiral operculum. In

fact, this grouping coincides well with the modern con-

cept of the Pachychilidae and Thiaridae, respectively

(Glaubrecht, 1996, 1999: Kohler and Glaubrecht, 2001,

2002, 2003). Therefore, operculum morphology in S.

sulcospira corroborates the placement of the taxon with-

in the Pachychilidae. This, however, represents a ple-

siomorphic character state among the caer ae of

this family, and is not a suitable character to establish

generic distinction.

By and large, the same can be stated for the radula.

The onollascen radula is generally considered a conser-

vative character (Fretter ond Graham, 1994). The pat-

tern described and depicted by Troschel (1558) (Figure

1) is commonly found among ‘pachychilids (Kohler and

Glaubrecht, 2001, 2002. 2003: Glaubrecht and Rintelen,

ne Rintelen and Glaubrecht, 1999, 2003). This had

been already noticed by Troschel (1858), when allocating

taxa such as Pachychilus and Sulcospira (but also Me-

Iage 9

Page 24

THE NAUTILUS, Vol. 119, No. 1

lanopsis) under the “Pachychili”. However, we have

been unable to identify a single radular character pe-

culiar to Sulcospira, based on ine limited anatomical tea-

tures discussed here. An enlarged main cusp is also

found in other pachychilid species, such as B. pagodula

(Kohler and Glaubrecht, 2001). Moreover, it is evident

that radular characters may especially be prone to ad-

aptation, parallelism, and convergence and that intraspe-

cific variability and Sato d in general may be consid-

erable, as described, e.g., for littorinid gastropods (Pa-

dilla, 1998; Reid and Mak 1999; Reid, 2000). Thus, pos-

session of an elongated cusp alone is not considered a

characteristic suitable for the diagnosis of Sulcospira.

Gross anatomy of S. Seas and S. martini remains

unknown, due to the lack of ethanol-preserved speci-

mens. It has been shown for other pachychilids, though,

that particularly characters of the reproductive organs

(pallial oviduct, gonads, brooding structures) ) and the

protoconch may ‘bear essential systematic information

(Kohler and Glaubrecht, 2001, 2003).

Juvenile shells extracted from dried adults (Troschel,

1858: and own observations) deliver circumstantial evi-

dence that Sulcospira is viviparous. Furthermore, as

made evident from the following comparison, the pro-

toconchs of S. sulcospira are similar to those of some

species we hav e primarily assigned to the “Brotia-tes-

tudinaria-group” (Kéhler and Glaubrecht, 2001, e.g.,

Brotia testudinaria and B. hainanensis: Figures 29- 30).

In contrast, other taxa constituting the genus Brotia sen-

su stricto (denominated the “Brotia pagodula group” by

Kohler and Glaubrecht, 2001) have juveniles with an ir-

regularly wrinkled sculpture of the apical whorl of the

protoconch (Figures 25-26). Protoconchs of Tylomelania

and Pseudopotamis exhibit yet another fine morphology

(Figures 27-34). They attain a relative lar ve size and

exhibit a relatively small apical whorl with a Hier shell

as well as regularly increasing whorls (Rintelen and

Glaubrecht, 1999, 2003: Glaubrecht a Rintelen,

2003).

As discussed in some detail by Kohler and Glaubrecht

(2001), distinct protoconch morphologies of several pa-

chychilid genera are correlated with different reproduc-

tive strategies. For example, Brotia possesses a subhae-

mocoelic brood pouch while representatives of Tylome-

lania and Pseudopotamis are characterized by an eu-vi-

viparous mode of reproduction utilizing a modified

oviduct as brood pouch (= uterine brood pouch; Rin-

telen and Glaubrecht, 1999; Glaubrecht and Rintelen,

2003). Females of the latter two taxa retain a small num-

ber of embryos in the uterus that are nourished by se-

cretions produce ‘d by the albumen gland.

The protoconchs of Sule ospira corre spond to those of

the so-called “Brotia ae group” (Kéhler and

Glaubrecht, 2001), which indicates to us ra S. suleos-

pira possesses a subhaemocoelic brood pouch as well.

Il. COMPARISON OF S. SULCOSPIRA AND S. MARTINI

Sulcospira martini, which has tentatively been allocated

the genus by Kohler and Glaubrecht (2002), can be

distinguished by its shell, but much more conspicuously

by its “different ‘protoconch morphology, which is unique

among the Pachychilidae.

Because the protoconch morphology is related to the

mode of reproduction, it is assumed that S. martini ex-

hibits reproductive features (e.g. incubatory structure,

reproductive strategy) that may be distinct from those

known from any other pachychilid taxon.

IIL. SysTeMATIC CONCLUSIONS

Shell, operculum, and radula of S. sulcospira (and also

of S. martini) are typically pachychilid; a basally well

rounded and flared aperture, a round to oval, multispiral

operculum, and a rachidian tooth with an enlarged main

cusp flanked by up to three accessory cusps that taper

in size are diagnostic characters of this family. However,

these characters are symplesiomorphic and, thus, unin-

formative at the generic level. Soft body morphology,

which could bear eee information, is not known. Still,

some systematic conclusions can be drawn based on the

evaluation of protoconch morphology, which is consid-

ered to be more or less constant at the generic level:

First, species of Brotia (Figures 27-28), Tylomelania

(Figures 31-32), ), and Pse Lilacs (Figures 33-34) are

not congeneric with S. S. sulcospira ( see Figures 19-20)

since they exhibit each a distinct protoconch, which is

testimony to a different reproductive strategy as dis-

cussed above. The same holds true for Jagora (Kohler

and Glaubrecht, 2003). All these pachychilid genera

have been shown to represent independent monophy-

letic lineages characterized by morphological features,

such as See: reproductive morphologies. Second, the

protoconch of S. sulcospira is very similar to that known

from spe cies of the “Brotia-te studinaria- -group * denom-

inated by Kéhler and Glaubrecht (2001) (Figures 29, 30

for B. hainanensis). Hence, Sulcospira Troschel, 1858,

being available and valid, might be an appropriate ge-

neric name for this species group under the precondition

that it can be shown that a protoconch with a smooth

and dome-shaped apical whorl is a character possessed

by the members of this group and derived by shared

ancestry. However, the alternative explanation that a

similar protoconch re presents an ancestral state that is

present in two different lineages has to be ruled out.

Otherwise, inferring generic relationship in absence of a

phylogenetic evaluation could lead to the erection of

pe araphy letic taxa when the characters considered are

plesiomorphic. Before we cannot show by phylogenetic

analyses oF morphologic il or molecular data that S. sul-

cospira and species of the “Brotia testudinaria group”

indeed belong to the same taxon, we therefore refrain

from a respective taxonomic suggestion.

Third, we conclude that S. martini can be recognized

as a species and that it does not represent a synonym of

S. sulcospira. Furthermore, it is clear that the morphol-

ogy of the protoconch of S. martini does resemble

ther of the known pachychilid genera very closely (Fig-

ures 27-34) including that of S. sulcospira (Figures 19-

F. Kohler and M. Glaubrecht, 2005

20). Just based on this single feature, it could be de-

duced that S. martini might bea represents itive of a yet

undescribed genus. However, as has been stated for S.

sulcospira, a sound decision on its systematics should

rely on a more comprehensive data set.

For the time being, we suggest to maintain Sulcospira

as a monotypic genus endemic to Java. Furthermore, we

refrain from a taxonomic decision on the generic rela-

tionship of “Sulcospira™ martini. It likely is not a mem-

ber of one of the described pachychilid genera, but its

true relationships remain unknown.

Irrespective of our anticipation that both species dealt

with in this paper may have already become extinct in

large parts of their original distribution area, we still

hope t that suitable material will turn up, eventually al-

lowing to find an answer to the remaining questions and

to solve another of the many puzzling aspects of pachy-

chilid phylogeny and syste inatics that long hampered a

deeper understanding of the evolution of this intriguing

and instructive case study among the lymnic C erithioi-

dea.

ACKNOWLEDGMENTS

We are most grateful to the curators Philippe Bouchet

(MNHN), Yves Finet (MNHG), Edmund Gitte snberger

and Jeroen Goud (RMNH), Ronald Janssen (SMF), Tru-

di Meier (ZMZ), Robert Moolenbeek (ZMA) and Kathie

Way (BMNH) for making material of their collections

available to us. Thomas von Rintelen kindly provided

SEM images of a juvenile shell of Tylomelania patriar-

chalis. We thank Philippe Bouchet and an anonymous

reviewer for most helpful comments on the manuscript,

in particular as related to nomenclatorial aspects. Their

comments helped much to improve the quality of this

paper. The work of F. K. was funded through a post-

graduate scholarship of the Konrad-Adenauer-Stiftung,

Sankt Augustin (Germany). This paper is part of a pro-

ject on Southeast Asian pachy chilids supported through

grant GL 297/4 to M.G. by the Deutsche For schungs-

gemeinschaft.

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THE NAUTILUS 119(1):27-42, 2005

Page 27

Six new species of Paryphantopsis (Gastropoda: Pulmonata:

Charopidae) from the Papuan Peninsula of New Guinea

John Slapcinsky

Florida Museum of Natural History

University of Florida

Gainesville, FL 32611 USA

[email protected]

ABSTRACT

Six new species of Paryphantopsis, a genus of charopid snails

endemic to New Guinea, are described from the geologically

complex eastern terminus of the Papuan Peninsula. All descrip-

tions are based on material collected in 2002-2003 and sa

information on shell, genital, and radular anatomy. Of the

previously described species, genital anatomy was known i

three and radular morphology for only one species. Examina-

tion of these under-utilized character sets has uncovered tax-

onomically useful interspecific variation. The six new species

appear to have distributions limited to particular mountain

groups on the extreme eastern terminus of the Papuan Pen-

insula. Species in close proximity or sympatry share unique

shell, genital and radular characters suggesting local speciation

and poor dispersal ability. Diversity in Paryphantopsis has been

underestimated and it is likely that there are many species yet

to be discovered with narrow geographic and ecological ranges

in the under-explored mountains of New Guinea.

INTRODUCTION

Paryphantopsis, a genus of charopid snails endemic to

New Guinea, are found at moderate and high altitudes

(600-4000 m) and are distributed from Western Papua

(Irian Jaya) to the Louisiade Archipelago. The genus

contains fourteen previously described species: P. arcu-

ata Jutting, 1964, P. dualoensis Solem, 1970, P. elegans

(Fulton, 1902). P. filosa Jutting, 1964, P. fultoni (Coen,

1922), P. globosa (Hedley, 1890), P. lame lligera (Thiele,

1925), P. latior Jutting, 1964, P. lowisiadarum (Méllen-

dorff, 1899), P a eine Jutting, 1964, P. pygmaca

(Bavay, 1908), P. sculpturata Jutting, 1964, P. similis

(Thiele. 1928), ss P. striata (Fulton, 1902). Solem

1970) reviewed the genus, redescribing all species ex-

cept those then recently described or reviewed by Jut-

ting (1964). Most A a le species are known

from small samples of shells, often only from their type

localities, and data on genital morphology are limited to

three species, and on radular morphology to only one

species (Solem, 1970, Wiktor, 2003). Paryphantopsis has

not been reported previously east of longitude 148° E

on the relatively poorly sampled eastern part of New

Guinea, which is known as the Papuan Peninsula. The

eastern end of the eee includes the Cloudy Moun-

tains to the south, and the disjunct terminus of the

Owen Stanley Range to the north, separated from the

main Owen Stanley uplands by extensive lowlands west

of Mount Suckling. The geologically complex Papuan

Peninsula is formed largely by the East Papua Compos-

ite Terrane (EPCT), a tectonic province compose dof at

least 4 separate geological units with differing ages, or-

igins, and histone s. These units appear to have assem-

led northeast of modern New Guinea during the Pa-

leocene, 62-57 Myr ago, and fused to the main body of

the island in the Late “Oligocene to Early Miocene, 25—

22 Myr ago (Pigram and Davies, 1987). Because of its

likely initial offshore amalgamation, the EPCT may have

develope ed a distinct and large ly endemic biota. This is

the first in a series of papers de scribing the results of

ten weeks of field surveys that took place during April-

May, 2002 and January-March, 2003. These surveys ex-

plored the extreme eastern terminus of the Papuan Pen-

insula where two geological units lie in close proximity,

the Cloudy Mountains of the Port More ssby Terrane and

eastern terminus of the Owen Stanle ny Range of the Kutu

Terrane (Figure 1).

MATERIALS AND METHODS

Specimens were hand-collected or sifted from samples

of leaf-litter. Live collected animals were drowned over-

night and then preserved in 75% ethanol. Gross anatom-

ical dissections were made under 75% ethanol using a

dissecting microscope. Radulae were isolated from dis-

sected buceal masses using a saturated KOH solution.

Scanning electron micrographs of radulae were made us-

ing a field emission SEM. Drawings of the genital anat-

omy were made with the assistance of a camera lucida,

and measurements were taken using an ocular microm-

eter. Shell measurements were made as follows. Whorl

count (W) was measured from the suture of the first

whorl to the body whorl and fractions of a whorl were

determined with the aid of a cardboard circle divided

into 10 equal parts of 36° (Figure 2, line 1-2.9). Spire

Page 28

THE NAUTILUS, Vol. 119, No. 1

= 9°

F Mount

Suckling

S 10° MKK «2550080

Mount

Simpson

0-100m

100-1000m

1000-2000m

2000-3000m

Owen Stanley Range

a 11°

E 149°

E250"

E i5i?

Figure 1. Distribution of Paryphantopsis on the eastern terminus of the Papuan Peninsula, Papua New Guinea between 9° S,

149° E and 11° S, 151° E. A = P. abstrusa, K = P. koragae. L = P. lebasii, M = P. matawanensis, U = P. ubwamensis, Y = P.

yawii, O = other sites sampled.

diameter (SD) was the length of a straight line passing

from the apertural edge of the suture through the mid-

dle of the apex to the opposite suture (Figure 2, line A-

B). Diameter (D) was the greatest width of the shell

perpendicular to the shell axis (Figure 3, line C—D).

Height (H) was the greatest distance between the apex

and the base of the aperture measured parallel to the

shell axis (Figure 3, line D-E). Spire height (SH) was

measured from the top of the body whorl to the apex of

the shell (Figure 3, line F—G). Aperture width (AW) was

the greatest distance from the columellar edge to the

outer edge of the aperture (Figure 3, line E-H). Aper-

ture height (AH) was measured from the suture to the

base of the aperture, parallel to the shell axis (Figure 3,

line H-I). The lengths of radular teeth were measured

from the top of the mesocone to the posterior edge of

the basal plate. The widths of radular teeth were mea-

sured as the greatest width of the cusps, not the basal

plate. The following abbreviations are used in figures of

genital anatomy: AT = atrium, DI = diverticulum, EP

= epiphallus, OV = free oviduct, PE = penis, PG =

prostate gland, PP = penial pilasters, PR = penial re-

tractor muscle, SD = spermathecal duct, SP = sper-

matheca, VA = vagina, VD = vas deferens, VP = vergic

J. Slapcinsky, 2005

Dace. OC

Page 29

Periostracal

Extensions

Periostracal

Processes

H E

Figures 2-3. Diagram of shell measurements, 2. Whorl count (line 1-2.9), spire width (line A-B). 3. Diameter (line C—D),

height (line D-E), spire height (line F-G), aperture width (line E-H), aperture height (line HI).

papillae. Specimens are deposited in the following insti-

tutions: Bernice P. Bishop Museum, Honolulu (BPBM),

Florida Museum of Natural History, Gainesville (UF),

Papua New Guinea National Museum, Port Moresby

(PNGNM), Wroclaw University Museum of Natural

History (MNHW).

SYSTEMATICS

Family Charopidae Hutton, 1884

Genus Paryphantopsis Thiele, 1928 (Type species:

Flammulina (Paryphantopsis) lamelligera Thiele, 1928,

by original designation.)

Description: Moderate to large-sized charopid snails

with loosely coiled shells of approximately three whorls.

Shell shape varies among species from globose to de-

pressed with an elevated to flat spire and a rounded to

keeled margin. The umbilicus, usually covered by an ex-

pansion of the peristome, is sometimes perforate. Nu-

clear whorls (protoconch) are sculptured with spiral rows

of small pits that become weaker and less regular on

later whorls. Postnuclear whorls (teleoconch) are usually

sculptured with growth lines accentuated with short per-

iostracal extensions punctuated with occasional longer

extensions at regular intervals. These longer extensions

can bear additional processes at the shell margin (Figure

2). A few species do not have longer periostracal exten-

sions and some species do not bear any extensions. Body

color is usually yellow in life. The epiphallus is apically

inflated. often with an apical diverticulum. The penis is

textured with convoluted pilasters of varying complexity.

Central teeth of the radula are tricuspid and of similar

size to the tricuspid and slightly assymetrical lateral

teeth. The many lateral teeth become shorter and less

symmetrical, grading in shape with the marginal teeth.

Marginal teeth are usually assymetric, their endocones

longer than their ectocones. Endocones and ectocones

and less often mesocones can bear accessory cusps.

Paryphantopsis abstrusa new species

(Figures 4-10, Table 1)

Description: The adult shell is small for the genus,

4.4-4.8 mm (mean = 4.6) in diameter and 3.5-3.8 mm

(mean = 3.6) in height, with 2.9-3.0 (mean = 3.0) rap-

idly expanding whorls (Figures 4-6, Table 1). The spire

is elevated 0.3 mm. Postnuclear whorls descend regu-

larly and the shell height/diameter ratio is ().76—0.81

(mean = 0.79). The shell has 1.3 evenly rounded nuclear

whorls, sculptured with 12 spiral rows of small pits. The

postnuclear whorls bear weak and irregular malleations,

most readily visible at the base, and indistinct and irreg-

ular striae on the apical surface. The postnuclear whorls

are also sculptured with weak growth lines. Approxi-

mately every fourth growth line is accentuated by a per-

iostracal extension. The periostracal extensions bear

small (0.15 mm) trigonal processes at the periphery.

These processes occur on every periostracal extension

for the first two whorls, then become less regular and

finally absent from the final % whorl. The fragile pro-

cesses are often partially worn from the earlier whorls

of adult shells. The nuclear whorls are white, the post-

nuclear whorls brown. The umbilicus is closed by a re-

flection of the peristome. The aperture is large, ovate to

nearly circular, with an aperture-diameter to aperture-

height ratio of 0.54-0.67 (mean = 0.63).

The body color is uniform bright yellow-white in life,

fading to cream in specimens preserved in ethanol. The

vas deferens narrows rapidly from the prostate gland and

remains narrow to the swollen head of the epiphallus

(Figure 7). A long coiled diverticulum inserts laterally

on the epiphallus soon after the junction with the vas

deferens. The epiphallus narrows slightly after the di-

verticulum and widens towards the junction with the pe-

nis. The penis is half the length and three times the

width of the epiphallus, robust, widest centrally, and nar-

rowing slightly at each end. Apically there are several

small convoluted pilasters and one much larger convo-

luted pilaster that extends to near the base (Figure §).

The penial retractor muscle is robust, originating from

Page 30 THE NAUTILUS, Vol. 119, No. 1

Figures 4-10. Paryphantopsis abstrusa. 4-6. Photographs of shell, Holotype UF 308235, diameter 4.6 mm. 7-8. Camera lucida

drawing of genitalia, UF 299677, maximum width 7.2 mm. 9-10. Scanning electron micrograph of radula, UF 299677, field width

of central and lateral teeth 41 xm, marginal teeth 42 xm

he is

J. Slapcinsky, 2005

Page 3]

Table 1. Measurements in mm of undamaged adult shells of six species of Paryphantopsis, N = count, H

= height, D = diameter,

SH = spire height, SD = spire diameter, AH = aperture height, AD = aperture diameter, W = number of whorls.

Species N H D SH sD AH AD W

P. abstrusa 3 mean+SD 36+02 46+02 32+ 0.0; QI=O0 26202 28=01 30+ 01

range 3.5-3.8 44-AS 0.3-0.3 2.0-2.] 2.4-2.7 2.7-2.9 2.9-3.0

P. koragae 3 mean+ SD 29+ 0.1 42+00 02+0.0 16+ 0.1 2.1 EO 2.6 =O] 2.8 + 0.1

range 2.8-3.0 4.14.2 0.1-0.2 1.5-1.7 2.0-2.1 2.5-2.7 2.7-2.8

P. lebasii 41 mean+ SD 36+ 0.2 6.8 + 0.5 0.1 + 0.1 25+ 0.2 2.8 + 0.2 43+ 0.4 28 + 0.1

range 3.34.0 6.1-7.5 0.0-0.2 2.3-2.8 2.4-3.0 3.9-4.9 2.7-2.9

P. matawanensis 5 mean+SD 45+05 70+04 O12+01 26+03 36+05 452+04 29+02

range $.0—-4.9 6.6-7.3 0.0-0.1 2.2-2.8 3.24.1 4.14.8 2.7-3.0

P. ubwamensis 3 mean+SD 50202 71+07 O38 01 30+03 38+03 43+05 30201

range 4,8-5.1 6.4-7.7 0.2-0.4 2.7-3.3 3.64.1 3.94.9 2.9-3.]

P. yawii 55 mean+SD 312+03 59+04 02+01 22+03 232403 37404 272011

range 2.6-3.6 5.0-6.5 0.1-0.3 1.9-2.6 2.0—2..7 3.24.1 2.6-2.8

the diaphragm and inserting at approximately mid-point

on the epiphallus. The spermathecal duct is robust, nar-

rowing abruptly at mid-point and remaining narrow until

joining the relatively small, ovate spe srmatheca. The free

oviduct joins the moderate length vagina above the atri-

um.

The central teeth of the radula (second row from left)

are tricuspid, S-9 wm wide and 11-12 wm long, roughly

the same shape as, but smaller than, the first lateral

teeth, which are 9-10 zm wide and 12-13 wm long (Fig-

ure 9). The mesocones of both the central and first lat-

eral teeth are tall, slender, and blade-shaped, joining the

rectangular basal plates close to, but not on, their pos-

terior edge. The mesocones of the ce sntral teeth barely

project beyond the anterior edge of the basal plates,

those of the lateral teeth project well beyond the edge.

The ectocones are trigonal and short only one third of

the height of the mesocones, joining the posterior edge

of the basal plates. The lateral teeth are asymme strical,

their endocones are slightly taller then their ectocones.

The marginal teeth are dorsoventrally compressed and

tricuspid to multicuspid, S-9 jzm wide and 11-12 ym

long (Figure 10). The endocones of the marginal teeth

the height of the mesocones and only slightly

taller than the ectocones. The mesocones of the margin-

al teeth often bear small cusps near the mid-point.

Type Material: Holotype: UF 308235, J. Slapcinsky,

16 April 2002; oe UF 299667 (2 specimens), UF

303558 (4 specimens), type locality, J. Slapcinsky, 16

April 2002.

are half

Type Locality: Papua New Guinea, Milne Bay Proy-

ince, Cloudy Mountains, Ubwam Mountain, headwaters

of the Watuti River, 10° 29.8’ S, 150° 14.02’ E, 670 m

altitude.

Habitat: Observed crawling near the base of trees

with smooth bark, in native forest, during wet weather

at 670 meters altitude.

Etymology: From the Latin abstrusa, a feminine ad-

jective meaning hidden, concealed and reserved, allud-

ing to the difficulty finding this species and to its subtle

alieniost concealed pe ieiosteacal processes.

Remarks: = Paryphantopsis abstrusa difters from most

other Paryphantopsis smaller than 5 mm in diameter by

having periostracé il extensions with processes at their

margins. Paryphantopsis similis is the only other small

species with periostracal extensions. It is more de-

pressed, with a height/diameter ratio of 0.70 compared

to a height/diameter ratio of 0.79 in P. abstrusa.

Paryphantopsis koragae new species

(Figures 11-17, Table 1)

Description: The adult shell is small for the genus,

4.14.2 mm (mean = 4.2) in diameter and 2.$—3.0 mm

(mean = 2.9) in height, with 2.7-2.8 (mean = 2.8) rap-

idly expanding eihoels (Figures 11-13, Table 1). The

spire is slightly elevated, 0.1-0.2 mm Rese = (0.2),

postnuclear whorls descend slowly and regularly. Shell

height/diameter ratio is 0.67—0.71 (mean = 0.69). There

are 1.1 evenly rounded nuclear whorls, sculptured with

12 spiral rows of small pits that are not continued on

the postnuclear whorls. The postnuclear whorls have

regular growth wrinkles with irregular, short, periostracal

extensions that are slightly we ae basally. None of the

periostracal extensions extend further than the others.

The nuclear whorls are white, the postnuclear whorls are

dark brown. The umbilicus is closed by a reflection of

the peristome. The aperture is large, flattened apically

and slightly angled at the periphery and base of the col-

umella. The aperture-diame to aperture-height ratio

is 0.78—0.80 (mean = 0.79).

The body color is uniform yellow in life, fading to

cream in ae preserved in ethanol. The vas def-

erens narrows rapidly after the prostate gland and re-

mains narrow until entering the swollen, ovate head of

the epiphallus (Figure 14). The interior of the head of

the epiphallus bears two strong pilasters that enter the

short broad diverticulum, which is roughly one quarter

of the length of the epiphallus. The e piphi illus is two to

three times longer than the penis and one third narrower

Page 32 THE NAUTILUS, Vol. 119, No. 1

Figures 11-17. Paryphantopsis koragae. 11-13. Photographs of shell, Holotype UF 305237, diameter 4.2 mm, 14-15. Camera

lucida drawing of genitalia, UF 303586, maximum width 6.3 mm. 16-17. Scanning electron micrograph of radula, UF 303586,

field width of central and lateral teeth 55 2m, marginal teeth 41 jzm

J. Slapeinsky, 2005

at the junction with the penis. The very short penial

retractor muscle originates on the diaphragm and inserts

near the mid-point of the epiphallus. The interior of the

penis bears several smooth, low, regular Me and

one much larger convoluted pilaster (Figure 15). The

spermatheca is oblong-ovate, its duct is apicall y narrow

and widens abruptly at the mid- point. The free oviduct

is relatively robust, joining the long vagina well above

the atrium.

The central teeth of the radula (center row) are tri-

cuspid, 9-10 am wide and 12-13 jm long, slightly

smaller than the first lateral teeth, which are 10-11 xm

wide, 13-14 pm long (

both central and lateral teeth barely project beyond the

basal plate. The ectocones of the central and lateral

teeth are about one half the height of the mesocones.

The lateral teeth are tricuspid and very slightly assy-

metric with the endocone of each lateral sligl itly taller

than the ectocone. The marginal teeth are domox entrally

compressed and tricuspid or weakly and irregularly mul-

ticuspid, 5-13 zm wide and 8-10 jm long (Figure 17).

The endocones of the marginal teeth are nearly the

height of the mesocones aad can have very weak sec-

ondary cusps; the ectocones are shorter and unicuspid

to irregularly multicuspid.

Type — Holotype: UF 308237, J. Slapcinsky,

20 February 2003; Paratypes: Papua New Guinea, Milne

Bay Province, Mount Matawan (Mount Simpson): UF

303586 (2 specimens), UF 303587 (2 specimens), type

locality; UF 303584 (1 specimen), plateau ENE of sum-

mit, 10°2.1' S, 149°34.6’ E, 2567 m altitude, J. Slapcin-

sky, 17 February 2003; UF303585 (1 specimen), Bunisi

V: illage, 10°1.1' S, 149°36.2’ E, 1450 m altitude, J. Slap-

cinsky, 16 February 2003.

Type Locality: Papua New Guinea, Milne Bay Prov-

ince, Mount Matawan (Mount Simpson): NE of summit,

10°L.7’ S, 149°34.7' E, 2100 m altitude.

Habitat: Active on plants and leaf litter usually near

the ground. Observed aestivating in suspended leaf litter

within 1.5 m of the ground in tr opical hardwood forest

and cloud forest from 1450 m to 2600 m altitude.

Etymology: This matronym honors Ms. Helen Kor-

age, Councelor for the Village of Ikara, who facilitated

our access to the Mount Matawan area.

Remarks: = Paryphantopsis koragae differs from other

species of Paryphantopsis in being sculptured with

growth lines accentuated with very short periostracal ex-

tensions. All other small species (= 5 mm diameter),

including P. filosa, P. pygmaea, P. arcuata, P. sculpturata,

P. similis, P. “platycephala, and P. abstrusa have occasion-

al longer periostracal extensions. The anatomy of only

one Pega species of similar size has ‘been fig-

ured previously: P. filosa ee Karkar Island near Ma-

dang (Wiktor, 2003. fig. 9). This species differs from P.

koragae in not having a bistieraet Cas on the epiphallus.

Din orp 28

age 33

Figure 16). The mesocones of

Paryphantopsis lebasii va species

(Figures 18-24, Table 1

Description: The adult shell is slightly larger than av-

erage for the genus, 6.1-7.5 mm (mean = 6.5) in di-

ameter and 3.3-4.0 mm (mean = 3.6) in height, with

2.7-2.9 (mean = 2.8) 1 rapidly expanding whorls (F Figures

18-20, Table 1). The spire is flat to slightly elevated, 0.0—

0.2 mm (mean = 0.1). Postnuclear whorls descend slow-

ly and regularly and shell height/diameter ratio is 0.45—

0.61 (mean = 0.53). There are 1.3 nuclear whorls, with

weak, peripheral and supraperipheral angles, and sculp-

tured with about 12 spiral rows of small pits. These pits

become larger and less regular on the postnuclear

whorls, where they are visib sle through the periostracum

as weak malleations. The sculpture of spiral rows of ob-

long pits is clearer where the periostracum is removed.

The shell periphery is weakly keeled. Apical surface of

the whorls is broadly rounded or with a very weak su-

praperipheral angle. Regular growth w rinkles accentu-

ated with short periostracal extensions are present on

the postnuclear whorls. On the apical surface, these ex-

tensions are folded along their length towards the ap-

erture and are weakly appressed to the shell. Approxi-

mately every fifth periostracal extension protrudes about

0.5 mm beyond the shell margin forming large rectan-

gular processes. These processes ov erlap "each: other on

the penultimate whorl, but not on the body whorl. They

are approximately equally spaced, rectangular, distally

rounded, and of approximately equal length. Periostracal

extensions of the growth lines are shorter, erect, and less

prominent basally. Nuclear whorls are white; postnuclear

whorls pale yellow brown. The umbilicus is perforate or,

less often, closed, covered to varying degrees by a re-

flection of the peristome. The aperture is large, de-

pressed-ovate with an aperture-diameter to aperture-

height ratio of 0.51-0.75 (mean = 0.65).

The body color is uniform bright creamy-yellow in

life, fading to creamy-white in specimens preserved in

ethanol. The vas deferens narrows to the junction with

the ovate head of the epiphallus (Figure 21). The epi-

phallus is approximately one quarter the diameter of the

penis and does not bear a diverticulum. The penial re-

tractor muscle is a little less than half the length of the

epiphallus, originating from the diaphragm and inserting

at the base of the epiphallus. The robust penis is a little

shorter than the epiphallus, with three strong pilasters

that run its entire length (Figure 22). The atrium is

short, expanding slig] itly towerd the junction with the

penis and vagina. The spermathec cal aut is massive at

the base, tapering rapidly at mid point, the remainder is

relatively narrow until its junction with the spherical

spermatheca. The free oviduct is slightly coiled and nar-

row, joining the very short vagina just above the atrium.

The central teeth of the radula (fifth row from left)

are symmetrically tricuspid, 89 jzm wide and 13-14 jm

long, and are similar in shape and length to the slightly

wader (Q- " wm), and slightly asymme strical lateral teeth

(Figure 23). The bluntly Sonics ‘and erect mesocones of

Page 34 THE NAUTILUS, Vol. 119, No. 1

Figures 18-24. Paryphantopsis Iebasii. 18-20. Photographs of shell, Holotype UF 308233, diameter 6.5 mm, 21-22. Camera

lucida drawing of genitalia, UF 299671, maximum width 6.7 mm. 23-24. Scanning electron micrograph of radula, UF 299671,

field width of central and lateral teeth 67 zm, marginal teeth 38 jum

J. Slapcinsky, 2005

the central and lateral rows join their basal plates cen-

trally and barely project beyond the anterior of their bas-

al plates. The ectocones of both the central and lateral

rows are trigonal and short, about half the height of the

mesocones; they join the posterior edge of their basal

plates at a low buttress. The e ndocones of the lateral

teeth are slightly larger but otherwise of similar shape

to their ectocones. The marginal teeth are dorsoventrally

compressed, multicuspid, about 9-LO jum wide and 10—

11 wm long (Figure 24). The endocones are nearly the

same height as the mesocones while the ectocones are

much shorter, about one half to one third the height of

the mesocones and divided into three cusps.

Type Material: Holotype: UF 308233, J. Slapeinsky,

9 April 2002; Paratypes: Papua New Guinea, Milne Bay

Province: UF 299676 (1 specimen); UF 299699 (5 spec-

imens), Cloudy Mountains, Ubwam Mountain, heacwa-

ter of the Watuti River, 10°29.8’ S$, 150°14.0' E, 675

meters altitude, J. Slapcinsky, 16 April 2002; UF 299674

(S specimens); UF 303593 (4 specimens), Pini Range,

Duabo Mission Station, 10°25’ 05” S, 150°1S’ 24 EB, 325

meters altitude, J. Slapcinsky, 9 April 2002; BPBM (2

specimens); MNHW 97S (2 specimens); PNGNM 004-

105 (2 specimens); UF 299671 (19 specimens) UF

303591 (1 specimen), 30 April 2002; UF 303590 (7 spec-

imens), 2 March ae Pini Range, E of “a tbo Mission

Station, 10°25.0’ S$, 150°18.6" E, 325 meters altitude, J.

Slapcinsky; UF prea 1 specimen), 30 April 2002: UF

303592 (1 specimen), | May 2002, Pini Range, aban-

doned logging road W of f Duabo Mission Station, 10°

24.9’ S, 150° 18.3" E, 325 meters altitude, J. Slapcinsky.

Type Locality: Papua New Guinea, Milne Bay Prov-

ince, Pini Range, Duabo Mission Station, 10°25’ 04.7"

S. 150°18’ 24.4” E, 325 meters altitude.

Habitat: Found on logs, mossy rocks and wet ground,

usually near streams in disturbed and undisturbed

broadleaf forest in hilly terrain from 325 to 700 meters

altitude. Observed active during the day.

Etymology: This patronym honors Mr. Biga Lebasi,

our host ad guide at Duabo Mission Station, the type

locality.

Remarks: Only four other Paryphantopsis species, P.

elegans, P. fultoni, P. yawii, and P. lamelligera have weak

to strong peripheral keels. Paryphantopsis lebasii is un-

like all species except P. yawii and P. fultoni, in having

large, rectangular periostracal extensions oriented par-

allel to the keeled shell mar gin. These peripheral exten-

sions are all of equal length, unlike in P. fultoni, and do

not overlap on the body whorl, unlike in P. yawii.

Paryphantopsis matawanensis new species

(Figures 25-31, Table 1)

Description: The adult shell is larger than average

size for the genus, 6.6-7.3 mm (mean = 7.0) in diameter

and 4.0-4.9 mm (mean = 4.5) in height, with 2.7-3.0

(mean = 2.9) rapidly expanding whorls (Figures 25-27

Table 1). The spire is flat or very slightly elevated, 0. 0.

0.1 mm (mean = 0.1). Postnuclear eos descend reg-

ularly and shell height/diameter ratio is 0.61—0.69 (mean

= (0.64). There are 1.3 rounded nuclear whorls, sculp-

tured with 6 spiral rows of small pits that grade into

weak, spiral striae on the penultimate whorl; shell sculp-

ture is obscured by periostracum on the body whorl. The

shell is wider and slightly angular below the mid- point.

The postnuclear whorls have regul: ir growth wrinkles ac-

centuated with low periostracal extensions that alternate

with several much longer extensions approximately every

10 growth-lines. The grouping of several longer exten-

sions appears like a single very thick pe sriostracal exten-

sion to the naked eye. The nuclear whorls are white, the

postnuclear whorls are dark brown to red-brown. A re-

flection of the peristome closes the umbilicus. The ap-

erture is large, depressed-ovate, with an aperture-di-

ameter to aperture-height ratio of 0.54-0.67 (mean =

0.63).

In life the body color is bright-yellow with lateral

patches of dark purple- brown, the yellow fades to cream

in specimens preserved in ethanol. The vas deferens nar-

rows toward the junction with the inflated spherical head

of the epiphallus. Immediately after, and perpendicular

to the head of the epiphallus, there is a finger-shaped

diverticulum that is roughly one quarter the length, and

slightly narrower than the diameter of the epiphallus

(Figure 28). The remainder of the epiphallus is some-

what twisted and approximate ly the same length as the

penis. The penial retractor muscle is short, originating

from the diaphragm and inserting on the basal ee of

the epiphallus. The epiphallus is roughly half the di-

ameter of the penis. The penis expands for its apical

third and then tapers basally to its junction with the sim-

ilarly sized atrium. The penis apex bears seve ral regular

pilasters oriented perpendicular to the length of fhe pe-

nis (Figure 29). At the pe nis mid- -point there i is one very

large and convoluted pilaster that extends basally. The

Ww: all of the base of the pe nis is thin, bearing regular small

pustules. The atrium is short and narrow ‘expanding

slightly towards the junction with the penis and the long

vagina. The base of the spermathecal duct is relatively

narrow, about the same diameter as the base of the penis

and free oviduct; it triples in size to its mid-point then

narrows abruptly for the remaining third before joining

the spherical spermatheca. The free oviduct is narrow,

joining the long vagina well above the atrium.

The central teeth of the radula (center row) are tri-

cuspid, 11-12 zm wide and 18-19 jum long, roughly the

same size and shape as the first lateral tee sth ( Figure 30).

The mesocones of both the central and first late al teeth

are tall and sharp, tapering apically and narrowing ba-

sally. Mesocones are atti ie d to their basal plates along

their entire le ngth, except for their apical quarter that

extend beyond the anterior margin of the basal plates.

The ectocones of the central teeth and the symmetric

ectocones and endocones of the lateral teeth are trigo-

nal, about half the height of the mesocones. The mar-

Page 36 THE NAUTILUS, Vol. 119, No. 1

Figures 25-31. 9 Paryphantopsis matawanensis. 25-27. Photographs of shell, Holotype UF 308236, diameter 7.3 mm. 28-29.

Camera lucida drawing of genitalia, UF 303581, maximum width 11.7 mm, 30-31. Scanning electron micrograph of radula, UF

303581, field width of central and lateral teeth 63 zm, marginal teeth 59 jm

J. Slapceinsky, 2005

ginal teeth are dorsoventrally compressed and irregularly

multicuspid, 11-12 pm wide and 10-12 pm long (Figure

31). The ectocones of the marginal teeth are slightly

shorter than their endocones, which are slightly shorter

than their mesocones. Both the ectocones and endo-

cones are irregularly multicuspid, the mesocones are

broadly trigonal to bro: idly rounded,

Type Material: UF 308236, J. Slapcinsky, 19 Febru-

ary 2003; Paratypes: Papua New Guinea, Milne Bay

Province, Mount Matawan (Mount Simpson): UF

303581 (9 tag UF 303582 (3 specimens), type

locality; UF 306529 (1 specimen), NE of summit,

10°2.1' S, 149° 34.4" r ae m altitude, J. Slapcinsky, 18

February 2003; UF 303583 (1 specimen), NE of sum-

mit, LO°L.7' S, aga 7 E, 2100 m altitude, ]. Slapcin-

sky, 20 February 2003.

Type Locality: Papua New Guinea, Milne Bay Prov-

ince, Mount Matawan ( Mount Simpson), ), ridge top E of

summit, 10°2.5’ S, 149°34.6’ E, 2700 m altitude.

Habitat: All specimens were collected in cloud forest

from 2100 io 700 m altitude. Individuals were observed

from 1-2 m above ground, crawling on a variety of cloud

forest vegetation in wet or foggy w eather In drier weath-

er specimens were observ ed in leaf litter that was sus-

pended in trees, especially in the crowns of Pandanus

sp.

Etymology: Named for the type locality and known

range of dhis species: Mount Matawan ( (Mount Simpson).

Remarks: Of the other larger (>6 mm diameter) spe-

cies of Paryphantopsis, P. P. globosa, and P. louisiadarum

do not have periostracal extensions on the growth lines,

unlike P. matawanensis. Of the species with periostracal

extensions, P. latior, P. lamelligera, P. fultoni, and P. stri-

ata do not have a repeating pattern of approximately LO

short periostracal extensions followed by several longer

extensions. P. matawanensis further differs from P. To

melligera and P. fultoni in not having peripheral pro-

cesses on the periostracal extensions. The genital anat-

omy of P. matawanensis differs from P. lamelligera in

having a diverticulum.

Paryphantopsis ubwame - new species

(Figures 32-38, Table 1

Description: The adult shell is large for the genus.

6.47.7 mm (mean = 7.1) in diameter and 4.8-5.1 mm

mean = 3.0) in height, with 2.9-3.1 (mean = 3.0), rap-

idly expanding whorls (Figures 32-34, Table 1). The

spire is elevated, 0.2-0.4 mm (mean = 0.3), the post-

nuclear whorls descend relatively ails espe cially near

the aperture. The shell height/diame ter ratio is 0.66—

0.75 (mean = 0.70). The 1.2 nuclear whorls are evenly

rounded and sculptured with 12 spiral rows of small pits.

These pits do not continue on the postnuclear whorls,

which are sculptured only with weak srowth lines that

do not bear periostracal extensions. The whorls are in-

Page 37

flated, the sutures deeply impressed, and the periphery

evenly rounded. The nuclear whorls are white, the post-

nuclear whorls brown, with irregular lighter patches.

The umbilicus is perforate, narrowed by a reflection of

the peristome. The aperture is large, ovate, with an ap-

erture-diameter to aperture- height ratio of 0.79-0.93

(mean = 0.85).

The body color is uniform yellow in life, fading to

cream in specimens preserved in ethanol. The vas def-

erens is wide at the prostate gland, narrowing slags!

and remaining narrow until the junction with the inflated

ovate tip of the epiphallus (Figure 35). The epiphallus

bears a long (approximately a quarter of the length of

the epiphallus), ), finger-she ped diverticulum just after the

junction with the vas deferens. The epiphallus is about

one third the diameter of the apex of the penis. The

penis is broad apically, narrowing abruptly to half its api-

cal diameter slightly before mid-point and remaining the

same diameter to the junction with the atrium, The pe-

nis is sculptured with several slightly convoluted pilasters

that extend in air are from near basally to near apically,

and one much larger and more convoluted pilaster near

the penis mid-point (Figure 36). The penial retractor

muscle originates from the diaphragm and inserts at the

mid-point of the epiphallus. The spermathecal duct is

basally robust and narrows at mid-point, remaining nar-

row until the junction with the spherical spermatheca.

The free oviduct is narrow joining the moderate length

vagina above the atrium.

The central teeth of the radula (center row) are tri-

cuspid, 10-11 pm wide and 15-16 jum long, roughly the

same width and shape as, but a little shorter than, the

first lateral teeth, which are 15-19 wm long (Figure 37).

The mesocones of both the central and lateral teeth are

long, slender and blade shaped, projecting slightly be-

yond the basal plate. The ectocones and endocones of

the lateral teeth are symmetrical and half the height of

the mesocones. The marginal teeth are dorsoventrally

compressed and irregularly tricuspid-multicuspid, 12-15

um wide and 13-15 pm long (Figure 38). The endo-

cones are tall, large to very large and sometimes irreg-

ularly multicuspid. The ectocones are unicuspid.

type Material: Holotype: UF 303589, J. Slapcinsky,

22 April 2002; Paratypes: Papua New Guinea, Milne Bz ly

Price: Cloudy Mountains, Ubwam Mountain: UF

299666 (1 specimen), type locality; UF 299668 (2 spec-

imens), headwaters of the Watuti River, 10°29.8' S,

150°14.02' E, 670 m altitude, J. Slapcinsky, 16 April

2002.

Type Locality: Papua New Guinea, Milne Bz wy Proy-

ince, Cloudy Mountains, Ubwam Mountain, suuminit,

10°30.4' S, 150°13.5' E, 1000 im altitude.

Habitat: This species was found crawling on a moss-

covered rotting log in cloud forest at 1000 meters. Dead

shells were also found at 670 meters in leaf litter at the

base of a tree with smooth bark.

Page 38 THE NAUTILUS, Vol. 119, No. 1

OT alae i emma Mle es iM ae.

7.7 mm. 35-36.

Camera lucida drawing of genitalia, UF 299666, maximum width LO. mm. 37-38. Scanning electron micrograph of radula, UF

Figures 32-38. Paryphantopsis ubwamensis. 32-34. Photographs of shell, Holotype UF 303559, diameter

299666, field width of central and lateral teeth 66 jm, marginal teeth 45 pom.

J. Slapeinsky, 2005

Etymology: This species is named for the type local-

ity, Ubwam Mountain, and indirectly for one of our

guides who is named after the mountain.

Type Material: Holotype: Papua New Guinea, Milne

Bay Province, Cloudy Mountains, Ubwam Mountain,

summit, 10°30.4" S, 150°13.5' E, 1000 m altitude, J.

Slapcinsky, 22 April 2002 (UF 303589); Paratypes: Papua

New Guinea, Milne Bay Province, Cloudy Mountains,

Ubwam Mountain: type locality (UF 299666, 1 speci-

men); headwaters of the Watuti River, 10°29.8’ S,

150°14.02’ E, 670 m altitude, J. Slapcinsky, 16 April

2002 (UF 299668, 2 2 specimens).

Paryphantopsis yawii new species

(Figures 39-45, Table 1)

Description: The adult shell is average in size for the

genus, 5.0-6.5 mm (mean = 5.9) diameter, and 2.6-3.6

mm (mean = 3.1) height with 2.6-2.5 (mean = 2.7)

rapidly expanding whorls ( Figures 39-41, Table 1). The

spire is elevated, 0.1-0.3 mm (mean = 0.2). Postnuclear

whorls descend slowly and regularly. Shell height/diam-

eter ratio is 0.47-0.55 (mean = 0.53). The 1.2 2 nuclear

whorls bear rounded peripheral | supraperipheral

ridges: sculptured with about 15 spiral rows of small pits.

These pits become larger and less regular on postnuclear

whorls but usually are obscured by the periostracum.

However, pits are visible in areas where the periostra-

cum is removed, and within the aperture. The shell has

a peripheral keel and blunt supraperipheral ridge; the

surface between is flattened. Postnuclear whorls are

sculptured with regular growth wrinkles that are accen-

tuated with periostrac ‘al extensions. On the apical surface

of the shell, periostracal extensions are folded along their

length toward the aperture and appressed to the shell

except at the keeled margin where they form rectangular

processes that extend about 0.5 mm beyond the shell

margin. The large, distally rounded processes over rlap,

forming a continuous periostracal fringe of uniform

length at the shell periphery. Pariostracal extensions on

the crowth lines extend basally but are short and erect.

Nuclear whorls are white, postnuclear whorls yellow

brown. The umbilicus is closed by a reflection of the

peristome. The aperture is large, de »pressed-ovate, with

an aperture-diameter/aperture-height ratio 0.53-0.66

(mean = 0.63).

The body color is uniform bright creamy-yellow in

life. fading to creamy-white in specimens preserved in

ethanol. The vas deferens narrows toward the junction

with the slightly inflated head of the epiphallus (Figure

42). The epiphall us is approximately three times longer

and one quarter of the diameter of the penis and ioe:

not bear a diverticulum. The penial retractor muscle is

long, about two thirds the length of the epiphallus, orig-

inating from the diaphragm and inserting on the cal

third of the epiphallus. The penis is shit and robust

with poorly defined pilasters in the apex (Figure 43).

The atrium is short and narrow, expanding slightly to-

Page 39

wards the junction with the penis and the short vagina.

The base of the spermathecal duct is massive, tapering

slowly but remaining broad for more than one third of

its length; the remainder is relatively narrow until the

junction with the spherical spermatheca. The free ovi-

duct is slightly coiled and narrow, joining the short va-

gina just above the atrium.

The central teeth of the radula (fifth row from left)

are tricuspid, 9-10 «zm wide and 14-15 wm long, rough-

ly the same size and shape as the first lateral teeth (Fig-

ure 44). The mesocones of both the central and first

lateral teeth are short, conical and erect, joining the rect-

angular basal plates nearly centrally and barely project-

ing beyond the plates’ anterior margin. The ectocones

are triagonal and short, only one third of the height of

the mesocones, joining the posterior edge of the basal

plates. The lateral teeth are asy mmeticak their endo-

cones are slightly taller then their ectocones. The mar-

ginal teeth are dorsoventrally compressed, tricuspid to

multicuspid, 8-10 wm wide and 10-12 wm long (Figure

45). The endocones of the marginal teeth are near ly the

same height as the mesocones while the ectocones are

much shorten only one half to one third of their height

and often divided into three cusps.

Type Material: Holotype: UF 308235, J. Slapcinsky.

6 April 2002; Paratypes: Papua New Guinea, Milne Bay

Province: UF 299675 (2 specimens), UF 303594 (2 spec-

imens), Wowow Mountain, W of Naura, 10°16.9’ S,

150°9.9' E, 635 m altitude, J. Slapcinsky, 8 May 2002;

UF 299673 (1 specime n), UF 303595 (1 specimen), 4

April 2002, UF 303598 (1 ‘specimen), 27 February 2003,

waterfall on Upalai Creek, 3 km WNW of Watunoa,

10°19.6' S, 150°34.6' E, 60 m altitude, J. Slapcinsky; UF

299670 (3 specimens), UF 299672 (6 (6 specimens), 6 April

2002, UF 303596 (13 specimens), 7 March 2003, head-

water of Goilayoli River at road crossing, 30 km ENE

of Alotau, 10°18.7' S$, 150°37.3' E, 275 m altitude, J.

Slapcinsky; BPBM (4 specimens), MNHW 977 ( (4 spec-

imens), PNGNM 004-104 (4 specimens), UF 303597 (45

specimens), small waterfall on Kinahidamadamana River

near Budo Village, 10°17.1' S, 150°26.7' E, 125 m alti-

tude, J. Slapcinsky, 4 March 2003.

Type Locality: Papua New Guinea, Milne Bay Proy-

ince, headwater of Goilayoli River at road crossing, 30

km ENE of Alotau, 10°1S.7’ S, 150°37.3’ E, 275 m al-

titude.

Habitat: This species was active during the day in na-

tive forest on rocks and logs with moss and algae, and

on moist soil near streams. It was encountered in hilly

terrain at relatively low altitudes for the genus, ranging

from 60 to 635 meters.

Etymology: This patronym honors Mr. Benjamin Yawi

of Budo Village, Milne Bay Province, Papua New

Guinea. He and his family located, arranged permission

to visit. and helped to collect at m: iny of the sites where

this species was found.

Page 40 THE NAUTILUS, Vol. 119, No. 1

Figures 39-45. Paryphantopsis yawii. 39-41. Photographs of shell, Holotype UF 308238, diameter 6.5 mm. 42-43. Camera

lucida drawing of genitalia, UF 299672, maximum width 6.4 mm. 44-45. Scanning electron micrograph of radula, UF 299672,

field width of central and lateral teeth 63 2m, marginal teeth 48 jum.

J. Slapeinsky, 2005

Page 4]

Remarks: Peripheral keels are unusual among known

Paryphantopsis species and are found only in P. elegans,

P. fultoni, and to a lesser extent in P. le basii and P. la-

melligera. Paryphantopsis yawii is unlike all other spe-

cies, except for P. lebasii and P. fultoni, in having large,

rectangular periostracal extensions oriented parallel to

the keeled shell margin. These peripheral extensions are

all of equal length, unlike in P fultoni and they overlap,

forming a continuous periostracal fringe, unlike in P. le-

basii.

DISCUSSION AND CONCLUSIONS

The family Charopidae was previously considered to be

a minor component of the terrestrial molluscan fauna of

New Guinea, with relatively few species and genera, and

to lack the spectacular radiations exhibited by this and

the related Endodontidae in the oceanic islands of the

Pacific (Solem, 1983: 305). Ongoing surveys indicate that

this is not the case; inadequate * sampling, rather than low

diversity, is the cause of the perceived low number of

charopid species in New Guinea. Likewise, reports of

low generic diversity of charopids in New Guinea are

more likely the result of insufficient sampling and may

reflect the paucity of anatomic material available to de-

fine generic units (Solem, 1970: 241). Despite the short

duration and limited geographic scope of the current

survey, six new species of Paryphantopsis are reported

here. increasing the known diversity of the genus by al-

most 50%. In addition to Paryphantopsis, species be-

longing to several other charopid genera were also col-

lected: these will be treated in later publications.

On the eastern terminus of the Papuan Peninsula,

Paryphantopsis species that occur in close proximity or

sympatry share unique shell, genital, and radular char-

acters. For example, both species from the Mount Ma-

tawan area, P- matawanensis and P. koragae, have un-

usually short penial retractor muscles, long vaginas and

angled apertural margins. Species in the uplands of the

Cloudy Mountains, P ubwamensis and P. abstrusa, are

unusually tightly coiled and globose. The two lowland

species, P. lebasii and P 1 yawii, share distally rounded

rectangular periostracal processes and the unusual origin

of fer mesocones from the center of their basal plates.

These unusual characters shared by different species in

close proximity or sympatry suggest that Paryphantopsis

species have speciated locally on a fine geographic scale;

speciation in these cases is presumably Peilitited by

their poor dispersal ability. Because much of New

Guinea remains under-explored, the true diversity of the

islands’ Paryphantopsis, other charopids, and land snails

in general is almost certainly greatly underestimated.

The geographic distibacon: of the six Paryphantopsis

species appears to be limited to particular mountain

ranges on different terrains, despite the proximity of

these mountains to each other (Figure 1). Three species,

P. matawanensis, P. koragae, and P yawwii are restricted

to the Owen Stanley Range, part of the Kutu Terrane,

while three others, P. abstrusa, P. lebasii, and P. ubwa-

mensis, are restricted to the Cloudy Mountains of the

Port Moresby Terrane. These distinct suites of endemic

species are consistent with the terrane-accretion hypoth-

esis (Davis et al. 1997) proposed for the formation of

the East Papua Composite Terrane and suggest that the

low vagility and high diversity of charopids and other

land cial may viralee them ideal to test hypotheses of

terrain acc1 retional | ustory.

ACKNOWLEDGMENTS

I thank the landowners of Alotau, Budo, Bunisi, Gadow-

alai, Ikara, and Naura for permission to work on -

land and for field assistance; J. Anamiato, I. Bigilale,

Kraus, F. Malesa, B. Uruwa, and B. Yawi for cia

field assistance; B. Lebasi for hosting my stay at Duabo

Mission Station; B. Yawi for help accessing land through-

out the Alotau area; G. Kula and D. Mitchell of Conser-

vation International for pr oviding logistical support and

advice; PNG National Museum a Art Gallery for pro-

viding in-country collaborative assistance; PN es Depart-

ment of Environment and Conservation, PNG National

Research Institute, and Milne Bay Provincial Govern-

ment for permission to work in Milne Bay Province: and

]. Worsfold for sharing bibliographic information. Field

work for this pesenecly Was supported by National Sci-

ence Foundation grant DEB 0103794 and the University

of Florida Foundation, McGinty Endowment. kK. Em-

berton, F. Kraus, G. Paulay, F. G. Thompson, and two

anonymous reviewers suggested i improvements to earlier

drafts of this manuscript.

LITERATURE CITED

Bavay, A. 1908. Mollusques terrestres et fluviatiles. Nova

nar Zoology 5: 269-292, pl. 14.

Coen, G. 1922. Descrizione di nuovo specie di molluschi

del Tate Civico di Genova. Annali del Museo Civico di

Storia Naturale, Genova 9(3): 359-363.

Fulton, H. C. 1902. Descriptions of new species of land Mol-

lusca from New Guinea. Annals and Magazine of Natural

apes) 7(9): 182-184.

Hedley, C. 1890. Description of a new Rhytida from New

Cuned, Annual Report of British New Guinea, 1555—

1889: 94.

Jutting, W. S. S. v. B. 1964. Non-marine Mollusca of West New

Guinea. Part 3, Pulmonata, I. Nova Guinea, Zoology 26:

1-74, pls. 1-2.

Mollendorff, O. 1899. Neue arten aus der Strubell’schen sa-

mmlung. Nachrichtsblatt der Deutschen Malakozoolo-

gischen Gesellschaft 31(5): S9-92.

Pigram, C. J. and H. L. Davies. 1987. Terranes and the accre-

tion history of the New Guinea orogen. BMR Journal of

Australian Geology and Geophysics LO: 193-211.

Solem, A. G. 1970. The Endodontid land snail genera Pilsbry-

charopa and Paryphantopsis (Mollusca: Pulmonata). The

Veliger 12: 239-264.

Solem, A. G. 1983. Endodontoid land snails from Pacific Is-

lands (Mollusca: Pulmonata: Sigmurethra). Part Il. Fam-

ilies Punctidae and Charopidae, Zoogeography. Field Mu-

seum of Natural History, Chicago ix + 336 p

Page 42

Thiele, J. 1928. Mollusken vom Bismark-Archipel, von Neu-

Guinea und Nachbar-Inseln. Zoologische Jahrbiicher 55:

119-146.

Wiktor, A. 2003. Terrestrial gastropods (Mollusca) of province

THE NAUTILUS, Vol. 119, No. 1

Madang in Papua-New Guinea. Part III. Pulmonata:

Rathousiidae, Ellobiidae, Succineidae, Agriolimacidae,

Endodontidae (partim), Ariophantidae, Euconulidae, Sub-

ulinidae, Streptaxidae. Folia Malacologica 11(1/2); 1-21.

THE NAUTILUS 119(1):48-49, 2005 Page 43

First record of Akera Miiller, 1776, from the eastern Pacific,

with the description of a new species

Angel Valdés

Natural History Museum of Los

Angeles County

900 Exposition Boulevard

Kelvin Barwick

City of San Diego

EMTS Laboratory

2392 Kincaid Road

Los Angeles, CA 90007 USA

[email protected]

San Diego, CA 92101 USA

[email protected]

ABSTRACT

The description of the new species Akera julicae is based on a

complete specimen collected from southern California and

three empty shells from Costa Rica. These specimens consti-

tute the first record of Akera in the eastern Pacific. Akera ju-

lieae is distinguishable from the widespread Indo-Pacific spe-

cies Akera soluta by its radular and jaw morphology; the rach-

idian teeth of A. soluta are more solid sr larger cusps and

flanking denticles, the mid-lateral teeth of A. soluta are den-

ticulated, whereas in A. julieae they are ace and the out-

ermost lateral teeth of A. julieae are proportionally more elon-

gate and straighter than those of A. soluta; the jaws of A. soluta

are well developed and composed of a number of rows of

strong rodlets, whereas in A. julieae the jaws are more rudi-

mentary with only five rows of { fragile-looking rodlets. There

are no consistent differences between these two species in giz-

zard plate and adult shell morphology, but the protoconch of

A. soluta is slightly larger and more elongate. The western At-

lantic species Akera bayeri is distinguishable from A. julicae by

having a more elevated shell spire and stronger radular teeth

with denticles on all lateral teeth.

INTRODUCTION

The opisthobranch family Akeridae Mazzarelli, 1891,

includes opisthobranchs with an external, cylindrical

shell into which the animal cannot retract completely. It

contains the single genus Akera Miiller, 1776, and only

a few Recent valid species distributed throughout trop-

ical and temperate regions. The type species, Akera bul-

lata Miiller, 1776, has been reported from the northeast

Atlantic, from the Baltic shores of Denmark and Norway

to the British Isles, continuing on to the Atlantic and

Mediterranean costs of continental France and Spain

(Thompson, 1976), Italy (Rinaldi, 1988), and the Canary

Islands (Ortea et al., 2001).

In the Indo-Pacific, the widespread species Akera so-

luta (Gmelin, 1791) has been reported from South Af-

rica and Mozambique to Australia, Marshall Islands, and

the Philippines (Gosliner, 1987). Both Akera bicincta

(Quoy and Gaimard, 1833), from Australia, and Akera

constricta Kuroda 1947, from Japan, have been regarded

as junior synonyms of A. soluta, see Wells and Bryce

(1993) and Hamatani (2000) respectively. This expands

the known range. for A. soluta. Other nominal Indo-Pa-

cific species, Akera tumida (A. Adams in Sowerby, 1850),

Akera tasmanica Beddome, 1852, and Akera aperta

Hedley, 1899, were all described from the southwestern

Pacific, and are poorly known. They are likely synonyms

of A. soluta, but because they were described based sole-

ly on shell morphology, their identities are unclear and

in need of revision. In the western Atlantic, Olsson and

McGinty (1951) reported for the first time a species of

Akera from Florida under the name “Akera thompsoni.”

Because the animal was figured but not described, this

species name is a nomen nudum in accordance with Ar-

ticle 13.1 of the International Code of Zoological No-

menclature (ICZN, 1999). The first available name for

the western Atlantic Akera is Akera bayeri Ev. Marcus

and Er. Marcus, 1967, which has been reported from

the southwestern Caribbean Sea and Brazil (Ev. Marcus,

1970).

There are no Recent species of Akera known from

the eastern Pacific. Akera maga Vokes, 1939, the only

known species from this area, was described from the

lower to middle Eocene, Domengine Formation, Fresno

County, California (Vokes, 1939). ). Squires (2001) report-

ed this species from the Llajas Formation (also lower to

middle Eocene), Ventura County, California. In the

present paper we describe the first occurrence of Recent

Akera in the eastern Pacific based on a complete spec-

imen collected in the Channel Islands, California and

three shells from Guanacaste, Costa Rica.

MATERIALS AND METHODS

The specimen from California was collected as part of

the Southern California Bight 1998 Regional Marine

Monitoring Survey (Bight, 1998) conducted in the sum-

mer of 1998. Infaunal samples were taken with a 0.1 m°

Van Veen grab sampler and screened through a 1 mm

mesh. The sample was then placed in a relaxant solution

of Epsom salts (magnesium sulfate heptahydrate—

Page 44

THE NAUTILUS, Vol. 119, No. |

Table 1. Comparative material examined in this study. The specimen marked with an asterisk (*) was collected alive and included

soft parts.

Species Locality

Date Depth Number

Akera soluta Kii, Japan

Uala Reef, Quezon, Philippines

New Zealand

Hardwick Bay, South Australia

Phuket, Thailand

Akera bullata Finmark, Norway

May 1959

— LACM 153414

3-15 m LACM 073035

LACM 153415

LACM 157942

LACM 153416

LACM 153417

LACM 046386

LACM 153415

MegSO,-7H,.O) and freshwater for a minimum of 30 min-

utes. The sample was then fixed in 10% buffered for-

malin and preserved in 70% ethanol. The preserved

whole animal was photographed using a digital camera

mounted on a dissecting scope (Wild Epimarkroskop®

M450).

The Costa Rican shells were collected during the

Searcher 401 Expedition of the Natural History Muse-

um of Los Angeles C ounty.

Several additional specimens and shells belonging to

other species of Akera were examined for comparison

purposes. These specimens are listed in Table 1

The shell is very fragile and could not be dissected

intact from the specimens examined, which rendered

the description below incomplete. Once the shell was

removed, the internal organs were dissected and drawn

using a Nikon SMZ LOOO microscope equipped with a

drawing tube. The radula, jaws, and gizzard plates were

didceoted and photographed using a Hitachi S-3000N

Scanning Electron Microscope (SEM). The apical region

of the shell was separated from the rest of the shell vie

mounted for SEM photography.

SYSTEMATICS

Akeridae Mazzarelli, 1891

Akera Miiller, 1776

Type Species:

typy.

Akera bullata Miiller, 1776, by mono-

Diagnosis: Shell external, fragile, translucent, cylin-

drical to slightly bulloid. Spire flattened to elevated with

a pi artially e »mbedde d protoconch. Ape rture equé al to the

length of the spire or Slightly shorter. Thin periostracum

forming a raised flange at the keel. Animal unable to

retract fully into the shell, but can stretch over twice its

length; with a

tending laterally, meeting mid-dorsally over the shell.

yosterior pallial tentacle. Parapodia ex-

Radula with rachidian tooth having a median cusp and

smaller denticles on either side. Rachidian tooth flanked

by 21-52 lateral teeth. Gizzard with a number of irreg-

ular plates arranged in three tiers.

Akera julicae new species

(Figures 1-4)

Akera sp.—Behrens, 2004: 1S, pl. LD.

Holotype: LACM 3033, from type locality.

Paratypes: 3 shells, southeastern corner of Bahia

Jobo, off sand beach west of Bahia de Salinas, Guana-

caste Province, Costa Rica (11°02'22” N, 85°45'16" W),

14 Feb. 1972, 1.5-10.7 m depth (LACM 3034).

Type Locality: Southwestern corner of Santa Catalina

Island, California, USA (33°18'24" N, 118°22’05” W), 24

Jul. 1998, m depth, in gray colored silt and clay

(LACM 303:

External sarees The body is oval, 10 mm long

in the preserved holotype. The cephi alic shield is trian-

gular, comprising about 4 of the body length in the pre-

served specimen (Figures 1A, 2A). The parapodia are

narrow and . not reach the midline of the body (Fig-

ures LA-C, ). The gill is unipinnate, with 11] simple

lamellae ae 2B). The color of the living animals is

unknown; the preserved holotype is grayish hike:

Shell Morphology: The shell is bullomorph, fragile,

well-calcified (Figure 1). The protoconch is smooth, aad

only the outer whorl is visible externally (Figure 3C). Its

maximum diameter is 200 jm. The whorls are se parated

by a deep, channelled suture, which is shallower on the

apical whorls. The periphery of the whorls, near the

apex, is angulated and forms a conspicuous keel, which

divides the whorls into two parts (Figure 1D, 3C). The

inner part has a characteristic pattern of strong, Gue

wrinkles. The whorls are attached to the preceding

whorl just below the periphery of this whorl. The apical

region is flattened and the external whorls overlap the

most internal. The whorl sides are clearly curved, con-

vex. The aperture is broad below and narrow above,

where it extends into a deep sinus along the suture of

the upper lip. There is a thin callus in the columella.

The sculpture consists of numerous, low and thin spiral

ribs and axial growth lines. The shells are covered with

a thin brownish periostracum.

Anatomy: The buccal bulb is oval; it connects poste-

riorly to the long esophagus and the salivary glands (Fig-

A. Valdés and K. Banwick, 2005

Figure 1.

B. Ventral view. Scale bar as in A. C. Lateral view. Scale bar

ure 2C). Two strong retractor muscles attach laterally to

the buccal bulb. The radular formula is 19*23.1.23 in

the holotype. The rachidian teeth are broad, with a tri-

angular based. basally concave, and a pointed central

Akera julieae new species, photographs of the preserved holotype (LACM 3033). A. Dorsal view. Scale bar = 5 mm

5 mm. D. Api al view of the shell. Scale bar 5 mm

cusp (Figure 4A). There are 3-5 denticles on each sid

of the cusp varying in shape and size. The two innermost

teeth on each row have a long and narrow ba

ni ]

conspicuously wider cusp bearing de noacires O

Page 46

THE NAUTILUS, Vol. 119, No. 1

Figure 2.

Scale bar = 1 mm. B. Detail of the gill. Scale bar =

of the reproductive system. Scale bar = 0.5 mm. E. Penis and prosate. Scale bar =

Akera julieae new species, drawings of the preserved holotype (LACM 3033)

_ A. Dorsal view of the complete body.

1 mm. C. General view of the anatomy. Scale bar = 1 mm. D. General view

0.5 mm. Abbreviations: ag, albumen gland;

am, ampulla; bb, buccal bulb: be, sais copulatrix; cs, cephalic shield; dg, digestive gland; es, esophagus; esg, external seminal

groove; fco, female copulatory organ;

ga, genital atrium; gl, gill: @z, gizzi ard:

i, intestine; mg, mucous gland; pn, penis; pr, prostate;

ps, penial sheath; rm, retractor nants sh, shell; sg, salivary gland: sr, seminal receptacle.

and outer sides. The rest of the lateral teeth are hook-

shaped, with a long and narrow cusp and lack denticles

(Figure 4B). In the outermost teeth the base is shorter

and the cusp proportionally longer than in the mid-lat-

erals (Figure 4C)

simple, elongate rodlets | (Figure 3B), The esophagus

opens into a large muscular gizzard, which contains sev-

eral gizzard plates. The gizz ard plates vary in shape and

size; they are irregular with angular edges ( (Figure 3A).

The reproductive system is monoaulic (Figure 2D).

The ampulla is long and convoluted; it opens into the

genital atrium at the same point where the albumen and

mucous glands open. The bursa copulatrix is oval; it con-

nects with the genital atrium through a wide and curved

duct. The genital atrium is long and is connected to a

complex female copulatory organ near the opening.

From the gonopore an open seminal groove runs in an-

terior direction to the protrusible cephalic penis and the

prostate. The penis is long and externally enclosed in a

). The jaws are compose d of 6 rows of

sheath (Figure 2E). The prostate is short and simple,

and connects proximally to the penis.

Etymology: Dedicated to Julie Barwick, the daughter

of the junior author.

DISCUSSION

Akera julicae has been included in Akera because of the

presence of a fragile, cylindrical external shell, a flat-

tened spire, and a parti lly embedded protoconch. The

radula of this species has a single broad, triangular rach-

idian tooth, with a median cusp and smaller denticles on

either side, as well as several hamate lateral teeth. The

gizzard contains a number of irregular gizzard plates ar-

ranged in three tiers. All these characteristics are diag-

nostic of the genus Akera (see above).

This is the first record of Akera in the eastern Pacific.

Behrens (2004) cited this species as Akera sp., based on

A. Valdés and K. Barwick, 2005

Page 47

Figure 3. Akera juliae new species and A. soluta (Gmelin, 1791). A-C. Akera julieae, scanning electron micrographs of the

22)

preserved holotype (LACM 3033). A. Gizzard plate. Scale bar =

300 pm. B. Jaw. Scale bar =

50 wm. C. Protoconch. Scale bar

= 200 am. D-E. Akera soluta (Gmelin, 1791), scanning electron micrographs of a specimen from Japan (LACM 153414). D.

Gizzard plate. Scale bar = 300 jm. E. Jaw. Scale bar = 50 wm. F. Protoconch. Scale bar = 200 jxm.

the material here examined and information, photo-

graphs. and descriptions provided by the junior author.

The description of the new species is mainly based on

the holotype, collected from California, but the three

empty shells collected from Costa Rica seem to belong

to the same species. However, this needs to be verified

when complete specimens from Costa Rica become

available.

Akera julieae differs from other described species of

the genus. Several shells and one specimen of the Indo-

Pacific Akera soluta were examined for comparison (see

Table 1) confirming the presence of several external and

internal differences between these two species. For in-

stance, the rachidian radular teeth of A. soluta are more

solid with larger cusps and flanking denticles (Figure

4D). The three innermost lateral teeth have wide cusps

Page 48

THE NAUTILUS, Vol. 119, No. 1

Figure 4.

of the holotype (LACM 303:

\. soluta, whereas only the two inner-

most teeth of A. fall ae have similar characteristics. The

mid-lateral teeth in A. soluta also have denticles (Figure

4), whereas they are smooth in A. julieae. The outer-

with denticles in A.

most lateral teeth of A. julicae are proportionally more

elongate and straighter than those ie \. soluta (Figure

4F). More importantly, the jaws of A. soluta are well

developed and composed of a number ae rows of strong

rodlets (Figure 3E), whereas in A. julicae the jaws are

Akera juliae new - cies and A. soluta (Gmelin, 1791).

A-C, Akera julicac, scanning electron micrographs of the radula

A. Rachidian and innermost lateral teeth. Scale bar = 50 pum, B. Mid-lateral teeth. Scale bar = 50

p.m. C. Outermost lateral realy Scale bar = 50 pm. D-F, Akera soluta, scanning electron micrographs of the radula of a specimen

from Japan (LACM 153414). D. Rachidian and innermost lateral teeth. Scale bar = 50 ym. E. Mid-lateral teeth. Scale bar = 50

pm. F, Outermost lateral teeth. Scale bar = 50 jm.

more rudimentary with only 5 rows of fragile-looking

rodlets. There are no consistent differences between the

gizzard plates of A. julicae and A. soluta (Figures 3A,

3D). Externally, A. soluta has a slightly larger and more

elongate seen The adult she lls of the “se two spe-

cies are indistinguish: able due to the morphological Var-

iability in A. soluta.

Akera oe ri is the only other species of Akera known

from the Americas. The external morphology and anat-

A. Valdés and K. Barwick, 2005

nae 40

Page 49

omy of this species was described in detail by Marcus

and Marcus (1967) and Marcus (1970). Akera bayeri dif-

fers from A. julieae in two important regards. The shell

of A. bayeri has a more elevated spire and the radular

teeth are stronger with denticles on all lateral teeth,

whereas in A. julieae the lateral teeth are smooth.

LITERATURE CITED

Behrens, D. W. 2004. Pacific coast nudibranchs, Supplement

I1—New species to the Pacific coast and new information

on the oldies. Proceedings of the California Academy of

Sciences 55: 11-54.

Gosliner, T. 1987. Nudibranchs of Southern Africa. Sea Chal-

lengers, Monterey, 136 pp.

Hamatani, I. 2000. Family Akeridae. In: Okutani, T. (ed.), Ma-

rine Mollusks in Japan. Toaki University Press, Tokyo, xlvii

+ 1173 pp.

ICZN (International Commission of Zoological Nomenclature).

1999. International Code of Zoological Nomenclature, 4"

Edition. International Trust on Zoological Nomenclature,

London, 306 pp.

Marcus, Er. and Ev. Marcus. 1967. Opistobranchs from the

southwestern Caribbean Sea. Bulletin of Marine Science

17: 597-628.

Marcus, Ey. 1970, Opisthobranchs from northern Brazil. Bul-

letin of Marine Science 20: 922-995.

Olsson, T. and L. McGinty. 1951. “Akera thompsoni.” The

Nautilus 65: pl. 3, figs. 7-7b.

Ortea, J. A.. L. Moro, J. J. Bacallado and R. Herrera. 2001.

Catalogo actualizado de los Moluscos Opisthobranquios

de las Islas Canarias. Revista de la Academia Canaria de

Ciencias 12: 105-134, pls. 1-4.

Rinaldi, E. 1988. Primi rinvenimenti di Akera bullata O. F.

Mueller, 1776 sulla costa Romagnola. Bollettino Malacol-

ogico 24: 25-26.

Squires, R. L. 2001. Additions to the Eocene megafossil fauna

of the Las Llajas Formation, Simi Valley, Southern Cali-

fornia. Contributions in Science 489: 1-40.

Thompson, T. E, 1976. Biology of Opisthobranch Molluscs,

Volume 1. The Ray Society, London, 207 pp.

Vokes, H. E. 1939. Molluscan faunas of the Domengine and

Arroyo Hondo formations of the California Eocene. An-

nals of the New York Academy of Sciences 35: 1-246, pls

]-22.

Wells, F. E. and C. W. Bryce. 1993. Sea Slugs of Western

Australia. Western Australian Museum, 184 pp.

THE NAUTILUS 119(1):50-54, 2005

Page 50

Description of Calliotropis pulvinaris new species (Gastropoda:

Trochidae: Eucyclinae: Calliotropini) from West Madagascar

Claude Vilvens

Rue de Hermalle, 113

B-4680 Oupeye

BELGIUM

[email protected]

ABSTRACT

Calliotropis pulvinaris new species is described from West

Madagascar and compared with similar iy cies in the trochid

subfamily Eucyclinae, particularly with C. patula (Martens,

1904), C. concavospira (Schepman, 1908), C. blacki. Marshall,

1979, and C. vaillanti (Fischer, 1582). The new species can be

separated from these by a rather depressed spire, a rounded

periphery, tumid whorls be saving four spiral cords of which nod-

ules decrease in size and increase in number from adapical

cord to abapical cord, and five spiral cords on the base.

INTRODUCTION

The malacofauna of this area remains poorly known, de-

spite earlier surveys (1971-1973) by ORSTOM (Office

de la Recherche Scientifique et Technique Outre-Mer,

now IRD: Institut de Recherche pour le Développe-

ment) on the continental slope of Madagascar (Crosnier

and Jouannic, 1973). Independently of the inherent eco-

nomic interest, this endeavor yielded abundant zZ00logi-

cal material, more particularly mollusks now de sposited

at the MNHN (Muséum national d'Histoire naturelle,

Paris).

Commercial fishing boats have trawled for deep-water

shrimp off Madagascar. The commercial dredging off

West Madagascar from these last years brought various

specimens of trochid species, some of them described

in the past (Watson, 1886; Martens and Thiele, 1904;

Thiele, 1925; Barnard, 1963), others recently named as

new — (Vilvens, 2001 and 2002).

Guido T. Poppe entrusted me with trochid shells cal:

lected in deep water. These shells, originally labeled as

C calliotropis patula (Martens, 1904), are conspecific with

material trawled by French expeditions and deposited at

MNHN in the 1970s. Closer examination and compari-

son with the type of the supposed species leads me to

Two years ago,

conclude that all these shells belong to an unnamed spe-

cies that is described here as new.

Text abbreviations used are: IRSNB: Institut royal des

Sciences naturelles de Belgique, Bruxelles, Belgium:

MNIIN: Muséum national d'Histoire naturelle, Paris,

France; NMNZ: Museum of New Zealand Te Papa Ton-

garewa, Wellington, New Zealand; ZMA: Zodlogisch

Museum, Amsterdam, The Netherlands; ZMB: Zoolo-

gisches Museum of Berlin, Germany; Pl, P2, P3, P4:

primary cords (P1 is the most adapical); stn: station; dd:

no live specimens present in sample.

I follow below the classification of Hickman and Mc-

Lean (Hickman and McLean, 1990) at the suprageneric

level.

SYSTEMATICS

Family Trochidae Rafinesque, 1$15

Subfamily Eucyclinae Koken, 1897

Tribe Calliotropini Hickman and McLean, 1990

Genus Calliotropis Seguenza, 1903

Type Species: Trochus ottoi Philippi, 1544, Pliocene—

Pleistocene, Italy; by original designation.

Calliotropis pulvinaris new species

(Figures 1-6)

Description: Shell rather long for genus (height up to

18.3 mm, width up to 29.0 mm), rather depressed, rath-

er thin, cyrtoconoidal; spire Yr ather low, he ight 0.6—

0.7Xwidth, he ight 2.3-4.5Xaperture height; smb?

deep and large. Protoconch about 300 jm wide, with

about 1 whorl, partially damaged on available specimens,

remaining part smooth. Teleoconch of up to seven con-

vex whorls, bearing four spiral granular cords and pro-

socline threads; nodule .s from cords produced by inter-

sections with axial folds on four first whorls; additional

axial threads not connecting nodules on last whorls. Su-

ture visible, impressed, not canaliculated. First teleo-

conch whorl convex, sculptured by about LS—20 proso-

cline smooth riblets, interspace be ‘tween riblets twice as

wide as riblets; primary spiral cords P2 and P3 appearing

almost immediately, evenly spaced, similar in size me |

shape, bearing rounded nodules produced by intersec-

tion with axial "ible ts. On second whorl, P2 and P3 stron-

ger, P1 appearing at end of whorl, close to P2. On third

whorl, nodules of Pl and P2 becoming sharp, with weak-

C. Vilvens, 2005 Page 5

Figures 1-8. Calliotropis species. 1-6. Calliotropis pulvinaris new species. 1-3. Holotype MNHN, northwestern Madagascai

18.129.0 mm. 4. Paratype IRSNB, West Madagascar, 1$.1*26.6 mm. 5-6. Paratype, collection C. Vilvens, West Madagascar,

18.3%25.4 mm. 7-8. C. concavospira Schepman 1908), syntype ZMA Indonesia, 6.0%8.7 mm

Page 52 THE NAUTILUS, Vol. 119, No. 1

Figures 9-16. Calliotropis species. 9-12. Calliotropis patula (Martens, 1904). 9-10. Syntype ZMB 55919, between Zanzibar and

Brawa, 16.2%25.1 mm. 11-12. Syntype ZMB 109933, Somaly, 19.0% 27.2 mm. 13-14. C. blacki Marshall, 1979, holotype NMNZ

M226932, Raoul Island, Kermadec group, 11.3% 13.0 mm. 15-16. C. vaillanti (Fischer, 1582), MNHN, Azores Islands, 10.6% 12.2

C. Vilvens, 2005

Table 1. Calliotropis pulvinaris. Shells measurements in min for all types cited and type material (n=8

aperature height; TW: number of teleoconch whorls.

. H: height; W: width; HA:

Measurement

TW H W HA L/W H/HA

Range 5.75-7.00 15.4-18.3 21.8-29.0 4.00-6.90 0.60—-0.70 2,304.50

Mean 6.69 17.11 94.7 5.71 0.69 3.10

Standard deviation 0.41 0.97 9.13 0.93 0.03 0.70

ly adapically oriented tip; P4 partially covered by suc-

ceeding whorl, with nodules smaller and more numerous

than ones on other cords. From fourth whorl on nodules

on cords decrease in size and sharpness, and increase in

number from Pl to P4; cords evenly distributed on

whorl: axial threads in area between cords remain thin,

distance between threads similar to threads width. On

last whorl, P4 ae aia no secondary spiral cords; pe-

riphery rounded.

Aperture almost circular, forming angle at junction of

inner and outer lips; this angle almost aaade d and outer

lip slightly flared in fully mature specimens; inner lip

flanged i in a curved are projecting weakly over umbilicus:

parietal lip forming thin, transparent | glaze. Columella

slightly curved, ata tooth, abapical part prosocline.

Base moderately convex, with 5 granular spiral cords,

innermost one stronger than others and bordering um-

bilicus; interspace between cords twice as wide as cords,

smaller for two innermost cords on young adult speci-

mens: very fine axial lamellate threads between cords,

poorly visible. Umbilicus wide, funnel shaped, diameter

about 35% of shell diameter, with very fine crowded axial

lamellae and no spiral cord within. Color of protoconch

and teleoconch pinkish white, with no maculations; first

two whorls somewhat brownish.

Type Locality: West Madagascar, 22°17.0' S,

43°02.2" E, 640-660 m, Indian Ocean.

Type Material: Holotype MNHN unnumbered (dd),

Chalutages Vauwban, stn. CH 112, coll. A. Crosnier, from

type locality, 18.1X29.0 mm; Paratype 1 MNHN un-

numbered (dd), 12°50'S, 48°09'E, 580-585 m, north-

western Madagascar, coll. A. Crosnier; Paratypes 2

MNHWN (dd), 1 IRSNB (IGnr 30 185) (dd), 1 ZMB

(Moll. 108.519) (dd), 1 G. T

. Poppe collection (dd), 1

C.Vilvens collection (dd), all from West Madagascar, off

Mahajanga (formerly Majunga), commercial trawlers

said to be from 800 m.

Etymology: Of a cushion (Latin); with reference to

the soft and oval form of the shell, without angulations.

Remarks: Calliotropis pulvinaris new species is simi-

lar to C. patula (Martens, 1904) (Figures 9-12) from

East Africa (off Somalia and Zanzibar, 977-1019 m), but

this species differs from the new species by having a

subangulated periphery, less tumid whorls, four spiral

cords on base (instead of 5) and also by the nodules of

Pl and P2 that become bigger than those of P3 much

later (4° whorl).

The new species weakly resembles C. concavospira

(Sche pman, 1908) (Figures 7—S) from Indonesia (also

deep water from $35 to $83 m), but this smaller species

has only three cords on the whorls and these whorls are

more angulate dd.

C alliotropis pulvinaris new species may ne pe com-

pared to C. blacki Marshall, 1979 (Figures 13 ), from

Kermadec Islands, but this Indo-Pacific species is ein

er for a similar number of whorls, has a more elevated

spire and only four spiral cords on the base.

The new — is also superficially similar to C. vail-

lanti (Fischer, 1882) (Figures 15-16) and C. ambigua

(Dautzenberg and pie »r, 1896), both from eastern At-

lantic, and to C. actinophora (Dall, 1890) from western

Atlantic, but these three spe cies have a more elevated

spire, only 3 spiral cords on the whorls and only four

spiral cords on the base.

ACKNOWLEDGMENTS

I would like to express my warm thanks to P. Bouchet

(Muséum national d'Histoire n: ture lle, Paris) for access

to the malacological resources of the MNHN, and V.

Héros (MNHN) f or her help in my search for scientific

papers. Also, I am especially grateful to J. L. Van Goe-

them (Institut royal des Sciences naturelles de Belgique)

for his constant he Ip, particularly with loan of er I

also would like to thank F. Koehler (Museum fiir Na-

turkiinde, formerly Zoologisches Museum, Berlin), B. A.

Marshall (Museum of New Zealand Te Papa Tongarewa,

Wellington) and R. Moolenbeek (ZoGlogisch Museum,

Amsterdam) for the loan of types from their institution.

Finally, 1 highly appreciate the judicious advice of R.

Houart.

LITERATURE CITED

Barnard, K. H. 1963. Contributions of South African marine

mollusca. Part IV. Gastropoda: Prosobranchiata: Rhipidog-

lossa, Docoglossa. Tectibranchiata. Polyplacophora. Solen-

ogastres. Scaphopoda. Annals of the South African Mu-

seum 47 (2): 201-360.

Crosnier, A. and C. Jouannic. 1973. Note dinformations sur

les prospections de la pente continentale malgache effec-

tuées par le N.O. Vauban (Bathymétrie—Sédimentolo-

gie—Péche au chalut). Documents scientifiques du Cen-

tre — YM de Nosy Bé 42.

Hickman, S. and J. H. McLean. 1990. Systematic revision

and ane neric classification of trochacean gasteropods

Page 54 THE NAUTILUS, Vol. 119, No. 1

Natural History Museum of Los Angeles County Science Vilvens, C. 2001. Description of a new species of Calliostoma

Series vit 169 pp. (Gastropoda: Trochidae: Calliostomatinae) from Madagas-

Martens, E. von and J. Thiele. 1904. Die beschalten Gastro- car. Novapex 2: 175-178.

poden der Deutschen Tiefsee-Expedition, 1895-1899. A. Vilvens, C. 2002. Description of Lischkeia mahajangaensis n.

Systematisch-geographischer Teil. Wissenschaftliche Fr- sp. (Gastropoda: Trochidae: Eucyclinae: Calliotropini)

gebnisse der deutschen Tiefsee-Expedition auf dem from East Madagascar. Novapex 3: 127-131.

Dampfer “Valdivia” 1595-1899, 7 (A): 1-146. Watson, R. B. 1886. Report on the Scaphopoda and Gas-

Thiele, J. 1925. Gastropoda der Deutschen Tiefsee-Expedition tropoda collected by HMS Challenger during the

IL Teil. Wissenschaftliche Ergebnisse der deutschen Tief- years 1873-1876. Report on es Heiae results ef the

see-Expedition auf dem Dampfer “Valdivia” 1898-1899, voyage of HMS Challenger, 1876. Zoology 15: 1-

17(2): 35-282. 680.

Notice

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The Curatorship, established originally in accordance with the wishes of the late Dr. R. Tucker Abbott, Founding Director of

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mea

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ett desl

THE NAUTILUS

Volume 119, Number 2

July 20, 2005

ISSN 0028-1344

A quarterly devoted

to malacology.

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The Bailey-Matthews Shell Museum

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ee 7 IN el a

CONTENTS

Volume 119, Number 2

July 20, 2005

ISSN 0028-1344

Guido Pastorino

Alvar Carranza

Walter Norbis

A revision of the genus Trophon Montfort, 1810

(Gastropoda: Muricidae) from southern South America................... 55

Latitudinal trends in shell characters of the neogastropod

Olivancillaria urceus (Gastropoda: Olividae) in the

temperate southwestern Atlantic Ocean... 0... cc eee 83

Errata .............0...

“ ,

‘ i

nad

ste

ane

‘

THE NAUTILUS 119(2):55-82, 2005

Page 55

A revision of the genus Trophon Montfort, 1810 (Gastropoda:

Muricidae) from southern South America

Guido Pastorino

Museo Argentino de Ciencias

Naturales

Angel Gallardo 470, 3° piso lab. 57

. 1 405D]R Buenos Aires

ARGENTINA

[email protected]

ABSTRACT

The genus Trophon from southern South America is revised

and restricted to ten valid species from among the 36 nominal

species still currently used. In addition, a new species, Trophon

parodizi from Patagonian waters, is described. Adult specimens

of the new taxon are illustrated, described and compared with

other living species of the same genus and similar geographic

distribution. Redescription and re-illustration of types, based

on material from several institutions around - world are pro-

vided for Trophon geversianus (Pallas, 1774), T. plicatus (Light-

foot, 1786), T. patagonicus (dOrbigny, 18: a T. acanthodes

Watson, 1882, T. pelseneeri Smith, 1915, T. amettei Carcelles,

1946, T. clenchi (Carcelles, 1953), T. wilhelmensis Ramirez-

Bohme, 1981, and T. bahamondei McLean and Andrade, 1982.

In addition, “Trophon” malvinarum Strebel, 1908, assigned to

genera incerta, is also redescribed and illustrated.

INTRODUCTION

Among many groups of marine gastropods from South

American waters in need of a modem comprehensive

revision, the high diversity and abundance exhibited by

the muricid genus Tro} hon Montfort, 1810, renders it

particularly interesting. Ear ly collections include a large

and varied array of specimens obtained by 19" century

expeditions from shallow waters of a vast area including

more than 5,000 km of coast in Argentina. A taxonomic

revision of these gastropods sevesled that a large num-

ber of names should be placed in synonymy. The tax-

onomy and nomenclature thus cle wified is sure to im-

aes ‘the usefulness of this genus and its species as tools

for biogeographic and evolutionary interpretations, with-

out forgetting that clear specific delimitation is crucial

to other uses such as the commercial exploitation of the

type species (Trophon geversianus) in southern Chile.

The subfamily Trophoninae is one of the most con-

spicuous groups of marine gastropods living presently

around the southern tip of South America. The southern

origin of the group seems to be beyond doubt, as dis-

cussed by Griffin and Pastorino (2005), when. revising

the numerous extinct species appearing in the fossil re-

cord since the late Oligocene.

This article Constitutes a review of all living species of

Trophon from both coasts of southern South America.

The study involves only those taxa living in environments

associated with the continental shelf. Accordingly, T. mu-

crone Houart, 1991, from 1500-1575 m off Brazil and

the subantarctic T. veronicae Pastorino, 1999, are not

considered herein. These two deep-water species seem

to belong in a different group according to data available

on the radula, protoconch, and penis of T veronicae.

Such anatomical data remain unknown for T) mucrone.

In addition, T. ohlini Strebel, 1904, with a distinct pro-

toconch and radula, different from those of other Pata-

gonian species and resembling the boreal Boreotrophon

truncatus, will be the subject “of a future paper. Finally,

T. triacanthus Castellanos et al., 1987, recently described

under Trophon, is also considered as belonging to a dif-

ferent genus—possibly Apyxistus—according to several

differences in protoconch and shell morphology.

Houart (2003) recently described three new species

under the genus Trophon sensu lato from enten

more than 1000 m in de »pth. No radular, anatomical, o

protoconch information is included in the diescristions.

However, enough differences can be observed in the

shells that, as mentioned by Houart himself, a new genus

may be granted for these species. A similar situation is

true for * ‘Trophon” malvinarum:; however, as there are

no accurate illustrations or recent descriptions of this

species I decided to include it in this work as belonging

to an indeterminate genus.

For each of the species considered herein, adult spec-

imens, operculum, gross anatomy, radula, protoconch,

and ultrastructure of the shell are described whenever

enough material was available. This work is part of a

complete revision of the genus including all species from

South America and Antarctica.

MATERIALS AND METHODS

All the material examined is housed in the collections of

the Museo Argentino de Ciencias Naturales “Bernardino

Page 56

Rivadavia”, Buenos Aires (MACN-In); Museo de La Pla-

ta, La Plata (MLP); National Museum of Natural His-

tory, Smithsonian Institution, Washington, D.C.

(USNM) and American Museum of Natural History,

New York (AMNH). Part of the type material is housed

in the following museums: The Natural History Muse-

um, London, (BMNH). Zoologisches Institut und Zool-

ogisches Museum der Universitit Hamburg, (ZMH);

Swedish Museum of Natural History, Stockholm,

(NHRM): Museum national d'Histoire naturelle, Paris

(MNHN): Museo Nacional de Historia Natural, Santia-

go, Chile (MNHNS): Museo Nacional de Historia Nat-

ural, Montevideo, Uruguay (MNHNM); Academy of

Natural Sciences of Philadelphia (ANSP) and Los An-

geles County Museum of Natural History (LACM). Fi-

nally, several historical specimens from the Auckland In-

stitute and Museum, New Zealand (AK) were examined

for comparative purposes. Material from the USNM

originates from the United States Antarctic Program

(USAP) and was mostly collected by two ships: R/V

Hero and R/V ELTANIN. Material collected by the Uru-

guayan ship R/V ALDEBARAN is also included.

Dissections were performed on ethanol-preserved

specimens for study of gross anatomy, with emphasis on

the morphology of the anterior alimentary system, and

the pallial portions of the male and female reproductiv e

systems. Radulae were prepared according to the meth-

od described by Solem (1972) and observed using a

LEO 440 scanning electron microscope (SEM) at the

USNM and a Philips XL 30 at MACN. Radular termi-

nology follows Kool (1993, fig. 6B). Shell ultrastructure

data were procured oat freshly fractured colabral sec-

tions taken from the central portion of the lip on the

last whorl of two individuals per taxon, whenever suffi-

cient material was available.

Photographs were taken using a digital scanning cam-

era. Several images were scanned from black and white

35 mm negatives using a slide scanner. All images were

digitally processed.

For the convenience of the reader in the Additional

Material Examined sections, “D” means that the speci-

mens were collected dead and “A” means alive.

SYSTEMATICS

Class Gastropoda Cuvier, 1791

Subclass Ortogastropoda Ponder and Lindberg, 1996

Superorder Caenogastropoda Cox, 1959

Order Sorbeoconcha Ponder and Lindberg, 1996

Infraorder Neogastropoda Wenz, 1938

Family Muricidae da Costa, 1776

Subfamily T [rophoninae Cossmann, 1903

Genus Trophon Montfort, 1S10

Murex magellanicus Gmelin, 1791

=Buccinum geversianus P all is, 1774) by original des-

ignation. Poly) plex Pe srry, LS11 (type species Polyple x bul-

bosa Perry, 1811] as ica geversianum Pallas, 1774)

and Muricidea Swainson, 1840 (type species Murex ma-

gzellanicus Chemnitz, 1780 (nomen nudum) =Murex ma-

el Species:

THE NAUTILUS, Vol. 119, No. 2

gellanicus Gmelin, 1791 =Buccinum geversianum Pallas,

1774) are synonyms.

Description: Shells variable in size, ranging from

about 1 to more than 10 centimeters high, fusiform, su-

bquadrate to elongate. Protoconch paucispiral, with 2 to

24) asymmetrical, ‘smooth, slightly globose, and regularly

convex whorls. Spire of about 4 whorls, equal or slightly

shorter than aperture height, never higher. Subsutural

ramp generally present, variously developed and slightly

‘nclined posteriorly, defining a conspicuous keel. Axial

sculpture variable, from weak growth lines to strong la-

mellae—in some instances even growing over the sub-

sutural ramp; axial sculpture generally betes developed

than spiral sculpture. eerie along keel sometimes

growing into fairly strong, always open, spines, which

may even curve backwar asi Spiral ornamentation. vari-

ably developed; sometimes missing, but generally con-

sisting of slightly rounded and equally dev eloped cords,

der than interspaces and sometimes accompanied by

weaker secondary intercalated ones. Spiral ornamenta-

tion usually restricted to surface abapical to keel, while

missing along subsutural ramp and in some cases only

present in the earliest whorls. Aperture subovoid; outer

lip sharp, sometimes slightly reflected, but always

smooth throughout. Siphonal canal always present and

open, although variably developed in length, sometimes

curved. Umbilicus variable, usually open, wide, although

in some species totally absent or represented bya hare

Shell ultrastructure arranged invariably in 2 layers: in-

nermost layer of crossed lanellar aragonite, outer layer

of amorphous calcite. Relative thickness of both layers

variable according to species or—within a given spe-

cies—latitude at which the population lives.

Radulae rachiglossate, rachidian teeth with 3 median

cusps, the central one the larger and the lateral ones

with a denticle, sometimes obsolete but always present,

attached to the interior margin (never free). Rachidian

base always sinuous and with the base offset under the

proximal tooth. Marginal cusps always single, never bifid.

Lateral teeth atonive: thin, with the attachment area also

thin. Operculum circular or suboval tear- shaped, attach-

ment area with horseshoe shaped scars.

Accessory salivary glands always developed, tubular,

single or coiled and miflled. Esophagus with a loop run-

ning along the left side of the gland of Leiblein; esoph-

ageal glands externally invisible.

~Egg-capsules always erect, never lenticular, usually

with nurse eggs.

Trophon geversianus (Pallas, 1774)

(Figures 1-21)

“Purpurschnecken” Knorr, 1769: 47, Pl . 30, fig. 2

“Buccin feuilleté” Knorr, 1770: 53, pl. 30, fig. 2.

Buccinum geversianum Pallas, 1774: 33, pl. 3, figs. 1, 2.

Buccinum foliaceum multifariam frondosum Chemnitz, 1780:

130, pl. 139, fig. 1297 {non-binominal, rejected by Opin-

ion 184 (ICZN, 1944) ) [Lectotype of Murex mage lanicus

Gmelin, 1791 (Beu, 1978)].

G. Pastorino, 2005 Page 57

Figures 1-16. Trophon geversianus (Pallas, 1774). 1-3. MACN-In 36036, Punta Cavendish, Puerto Deseado, Santa Cruz province

in 5 m. 4-5. MACN-In 36042, Cueva del Indio, Puerto Deseado. 6-9. MACN-In 36041, Sierra Grande, Rio Negro province, in

tide pools. 10-11. MACN-In 36037, Punta Penas, San Julian, Santa Cruz Province in 2 m. 12. MACN-In 36043, Bahia Almanza

Puerto Harberton, Tierra del Fuego, in 3 m. 13-14. MACN-In 36038, both specimens from Playa La Mina, San Julian, Santa Cruz

Province intertidal. Scale bar for all shells = 1 cm. 15-16. Two views of protoconch, arrow head the transition to teleoconch. Scale

bar = 500 wm

THE NAUTILUS, Vol. 119, No. 2

Figures 17-21.

Buccinum fimbriatum Martyn, 1754: fig. 6

Murex magellanicus Gmelin, 1791: 3548 partim (var. B excl.):

dOrbigny a 451: Ais 1828: 127, pl. 26, fig. 90;

Hanley, 1556 oe ig. 9O

Neptunea . foliacea eas 758. 116

?Murex ventricosus ve 1S10: 178

Polyplex bulbosa Perry, 1811: pl. 9, fig. 5

Murex foliatus Schumacher, 1$17: 215, sensu Vokes 1971

Fusus magellanicus Lamarck. Gray, 1839: 115

?Murex varians dOrbigny, 1839: pl. 42, figs. 4-7; d’Orbigny,

$41:452

T. geversianus Pallas—Montfort, 1810: 483, fig.; H. and A

Adams, 1853: 77, pl. 8, fig. 3 c.; Tapparone-Canefri, 1874

5; Kobelt, 1878: 205, pl. 72, fig. 1-3; pl. 73, fig. 1; Sow-

erby I, 1880: pl. 404, figs. 7,8; Tryon, 1880: 144, pl. 32,

7-340, 343-347; pl. 70, figs. 433; Watson, 1886:

64: Rochebrune and Mabille, 1889: H.53; Strebel, 1904:

73, pl 4, figs. 11-23; pl. 5, figs. 24-42: pl. 6, figs. 43-52

Lamy, 1906: 3: Ihering, 1907: 404; Melvill and Standen,

907: 106; Strebel, 1908: 37, pl. 6, figs. 94 a, b.; Carcelles

946: 60, figs. 1-5; 1946: 69, figs. 6Ga,b; TZa,b,c,d; 8; Powell,

951: 151, fig. L, 81; N, 107; Castellanos, 1970: 76, pl. 5

ig. 2: Dell, 1971: 210; Harasewych, 1984: 13, figs. 1-3

25: Vokes, 1991: 7, fig.; 1992: 3, figs le, d: 3c, d; Kool

993: 47, figs. 9-14, 30-31: Castellanos and Landoni

1993: 3, pl. 1 figs. 1-15, 1S—2]

figs. 33

Trophon geversianus (Pallas, 1774). 17. Radula, frontal view. Scale bar = 100 jm. 18. Lateral view of the same

radula. Scale bar = 30 jzm. 19. Ultrastructure of the shell. Scale bar = 100 wm. 20. Penis, critical-point dried. Scale bar = 800

pm. 21. two views of the operculum. Scale bar = 1 cm.

Fusus intermedius Hupé in Gay, 1854: 166, pl. 4, fig. 6, non

Cristofori and Jan, 1832 nomen nudum: nec A. J. Mich-

elotti, 1846 nomen nudum: nec G. Michelotti, 1547:

Rochebrune and Mabille. 1SS9: 11.53

F. geversianus Pallas —Hupé in Gay, 1$54: 167; Gould, 161:

pl. 16, fig. 277 a, b

Trophon geversianus var. calva Kobelt, 1S78: 305, pl. 75; fig. 1

T. geversianus var. lirata Kobelt, 1878: 305, pl. 76, fig. 1, 2.

philippianus Dunker in Kobelt, 1878: 277, pl. 72, figs. 4, 5;

Melvill and Standen, 1907: eo Powell, 1951. 152.

philippinarum Dunker. Sowerby Il, LSS0: pl. 405, fig. 21.

°T. varians (VOrb,)—Carcelles, oe 131, figs. 1, 2, 3, 6, 7;

Castellanos, 1970: 75, pl. 5, fig. 3, 4; Vokes, 1992: 3 fig. 4c.

plicatus (Lightfoot)—Calvo, 1987: 135, fig. 99

Trophon sp Vokes 1991: 9, figs. 1-13

Description: Shell large (up to 100 mm) and extreme-

ly variable, fusiform, subqui idrate profile, chalky, whitish;

protoconch of 2 whorls, smooth, cylindrical, slightly olo-

bose, slightly asymmetrical; te leoconch of 4 shouldered

whorls, spire less than % of total shell he ight. S Spire angle

about 50°; suture impressed; subsutural shelf straight,

aperture ovoid, interior glossy pinkish; anterior siphonal

canal moderately long (half the height of aperture); um-

bilicus closed or deep. some specimens with a pseu-

G. Pastorino, 2005

Page 59

doumbilical chink; outer lip rounded, with reflected edg-

es; inner lip curved, adpressed.

Axial ornamentation of irregular, low lamellose varices

on first whorls, becoming S—10 well-defined lamellae on

last ones. Lamellae growing across entire whorl, attached

to the shell, sometimes curving adavxially, Lamellae end-

ing in shallow peripheral spine, in some specimens grow-

ing adapically.

Spiral ornamentation of about 15 cords beginning at

periphery of whorls. Smooth specimens common in in-

tertidal pools : and ‘ayelia } yanks. Coloration varying from

creamy white to dark brown. Growth lines regularly

space od, present throughout shell. Geographic variation

conspicuous from north to south and from intertidal to

infralittoral specimens, expressed as a series of smooth

to profusely ornamented specimens according to area of

collection.

Shell ultrastructure composed of two layers; inner-

most layer (50% shell thickness) of * colabrally aligned

crossed lamellar aragonite, outer layer (50% shell thick.

ness) with amorphous calcite.

Operculum oval, brownish, with terminal nucleus. Ex-

ternal surface covered by concentric, irregular, growth

lines. Inner surface attachment area with 3 horseshoe-

shaped scars, thick glazed rim present in all specimens.

Anatomical and radial: ar characters as described by

Harasewych (1984) and Kool (1993).

Type Material: The type material upon which Pallas

(1774) based Buccinum geversianum could not be lo-

cated. It could not be found at the Zoological Institute

of the Russian Academy of Sciences (ZIL) St. Peters-

burg, where most of the material studied by that author

is housed (B. Sirenko, pers. comm.).

Additional Material Examined: 53°39’ S, 70°55.5' W,

5 A, R/V Hero Cruise 702, Sta. 466, 25 April 1970, 20

m (USNM 901605); 53°39’ S, 70°55.5' W, 6 A, R/V

Hero Cruise 702, Sta. 473, 26 ue 1970, 15-18 m

(USNM 901609); 53°30'48” S, 70°50'33" W, 4 A, R/V

Hero Cruise 692, Sta. 69-11, 24 ner 1969, intertidal,

(USNM 901604); 53°24.8' S, 69°39.2' W, 1 D, R/V

Hero Cruise 702, Sta. 481, 27 April 1970, 1S m, (USNM

901605): 53°17’ S, 68°13’ W, 1 A, R/V Hero Cruise 712,

he 71-2-6, 21 April 1971, 1 m, (USNM 886739);

3°51'32” S, 70°25'52"” W, 1 A, R/V Hero Cruise 692,

ae 69-22, 13 May 1969, 2-3 m (USNM_ 886187);

53°37.9' S, 70°14’ W, 1 D, R/V Hero Cruise 702, Sta.

486, 28 April 1970, 292-296 m (USNM 901602); 54°59"

S, 64°53’ W, 1 D, R/V ELTanin Cruise 11, Sta. 970, =

February 1964, 586-641 m (USNM 870515); 53°48.7'

70°24.1’ W, 1 D, R/V Hero Cruise702, Sta. 489, 29 heal

1970, 13-18 m (USNM 901601); 54°27’ S, 66°12' W, 2

D, R/V ELTaNIN Cruise 6, Sta. 453, 21 January 1963, 31

m (USNM 901600); 53°26’ S, 68°35’ W, 6 A, R/V EL-

TANIN Cruise 21, Sta. 297, 12 January 1966, 0 m (USNM

901607): 53°35’ S, 70°50’ W, 1 D, 1 A, R/V ELTANIN

Cruise 21, Sta. 292, § January 1966, 0 m (USNM

901606); 53°17’ S, 68°13’ W, 3 D, B/V ELTANIN Cruise

712, Sta. 71-2-6, 21 April 1971. 0-1 m (USNM 901608):

Puerto Basil Hall, Isla de los Estados, 10 A, 21 May

1971, 0-1 m (MLP 4243): Buen Suceso Bay, Tierra del

Fuego, 4 A, 23 October 1941 (MLP 27218): San Julian,

Punta Penas, 6 A, 1 March 1924 (MLP 526 partim):

Puerto Lobos, Chubut, 12 A, 2 February 1938 (MLP

2021); Puerto Golondrina, Ushuaia, 9 A, January 1962

(MLP 27201); Puerto Piramides, Chubut, 1 A (MLP

4715); San Julian, 8 A (MLP 1583); Puerto Hoppner,

Isla de los Estados, 7 A (MACN-In 22547); Punta Cav-

endish, Puerto Deseado, 7 A, 6 February 1961 (MACN-

In 26171): Punta Cavendish, Puerto Deseado. Santa

Cruz province, 5 m (MACN-In 36036); Chubut, 6 A

(MACN-In 4097); Tierra del Fuego, 4 A (MACN-In

5777-1); Cueva del Indio, Puerto Deseado (MACN-In

36042); Sierra Grande, Rio Negro province, in tide pools

(MACN-In 36041); Punta — es Julian, Santa Cruz

province, 2 m (MACN-In 36037 » Playa La Mina, San

Julian, Santa Cruz province eee (MACN-In

36038); Bahia Almanza, Puerto Harberton, Tierra del

Fuego, 3 m (MACN-In 36043); San Antonio Oeste, Rio

Negro, intertidal, 3 A (MACN 35387); 54°47'36" S,

64°22'35"” W, 1 A, R/V ELTANIN Cruise 712, Sta. 71-2-

44, 24 May 1971, intertidal (USNM 881127); 54°46'12”

S, 64°24'42” W, 1 A, R/V ELTANIN Cruise 712, Sta. 71-

2-46, 21 May 1971, intertidal (USNM 881131); 54°4536"

S, 64°02'36" W, 1 A, R/V ELTANIN Cruise 7151, Sta. S69,

23 October 1971, intertidal (USNM 881132); 54°45'45”

S, 64°09'55" W, 1 A, R/V Hero Cruise 712, Sta. 71-2-

40, 21 May 1971, intertidal (USNM 881130); 54°47'48"

S, 65°16’ W, 1 A, R/V Hero Cruise 712, Sta. 71-2-8, 23

April 1971, intertidal (USNM 881126); 54°48’, 65°14" W,

1 A, R/V Hero Cruise 712, Sta. 71-2-14, 25 April 1971,

intertidal (USNM_ 881129); Punta Arenas, Cabeza de

Mar, Chile (ANSP 101444); Ushuaia ee peas

Malvinas Is. (ANSP 277535 and 277538): Cape Fair-

weather, Santa Cruz province, Argentina (ANSP 78080);

Straits of Magellan (ANSP 36241 and ANSP 366497);

Puerto San Julian ( (ANSP 312324); Santa Cruz River

(ANSP 101445); Puerto Gallegos (ANSP 312319); Golfo

San Jorge (ANSP 178645): Puerto Madryn, Chubut

(ANSP 170471); Puerto Parry, Isla de los Estados, 54°46’

S, 64°23’ W (ANSP 402810); mouth of Santa Cruz River

(ANSP. 88536); Punta Arenas (ANSP $8549); Puerto

Madryn (ANSP 170474); Punta Arenas, Chile (ANSP

199711).

Distribution: Trophon geversianus has the widest

geographic range of all species of Trophon, i.e., from

Buenos Aires province to Burdwood Bank in the south-

western Atlantic, Tierra del Fuego and Malvinas Is., and

the Magellan Strait in Chile. Literature records from

around Antarctica are almost certainly wrong assign-

ments, probably mistaking it for Trophon nucelliformis

Oliver and Picken, 1984, T. mac quariensis Powell, 1957,

or T. albolabratus Smith, 1875.

Remarks: = Troplion geversianus is the best-known spe-

cies of the entire genus. The great morphological vari-

ation can be appreciated from the large number of

names proposed for the different morphological variants

Page 60

in this species. Zaixso (1973) and Penchaszadeh (1976)

both studied the egg capsules of this species but only

the latter confirmed the existence of short-lived nurse

eggs in the capsules.

Trophon varians is a dubious species described by

dOrbigny from material he collected in northern Pata-

gonia (“au sud du Rio Negro”). This species has no la-

mellae, weak spiral cords, ad its shell is unusually thick.

Houart (1998) illustrated a paralectotype (as syntype)

housed in Paris and Aguirre (1993) designated and fig-

ured the lectotype (BMNH 1854.12.4. 539) from 13 syn-

types from the BMNH collection. According to

dOrbigny’s illustration (Plate 42, figs. 4, 5) and the lec-

totype and the paralectotype housed in Paris, it appears

that its distinctive characters are the thickness of the

shell and absence of lamellae. However, despite this dif-

ference between T. geversianus and these primary types

of T. varians, the rest of the paralectotypes are very sim-

ilar to other thin-shelled specimens of Trophon gever-

sianus usually found exposed during low tides in noite

em Patagonia, Golfo San Matias and around the Valdés

Peririsula: The anatomy and radula of the latter are

identical to those of T. geversianus. The specimens with

heavy shells are characteristic of the mouth of Rio Ne-

gro. Nothing is known about the anatomy and radula of

Hess heavy specimens. Perhaps these characters may

prove that it is only a local variation of Trophon gever-

sianus, as it was suggested originally by d’ Orbigny him-

self (1841: 452). D’Orbigny (1841: 452; 1839, plate 42

figs. 6-7) also described and illustrated the egg- -capsules

Fone what he supposed were z. varians. The illustration

agrees better with capsules of Urosalpinx haneti (Petit,

1856).

Trophon plicatus (Lightfoot, 1786)

(Figures 22-42)

Le Sabot Magellanique Favanne, 1780: 3

men nudum.

Buccinum laciniatum Martyn, 1784: fig. 42, nomen nudum.

Murex plicatus Lightfoot, 1786: 104.

M. lamellosus Mz oa Gmelin, 1791: 3536; Wood and Han-

ley, 1856: 133, pl. 27, fig. 100

Polyplex gracilis Pe srry, ISL1: pl. 9, fig. 4

Fusus laciniaius Martyn.—Reeve, 1847: spec. 14, figs. a, b, c.;

Hupé in Gay, 1854: 168; Gould, 1861: pl. 16, fig. 278.

Trophon laciniatus Martini—H. and A. oe 1853: 77, pl.

8, figs. 3 a, b.; Kobelt, 1878: 280, pl. 72, figs. 6, 7; Tryon,

1880: 143, pl. 31, figs. 330-332. Rochen and Mabille.

1889: H.53:; Stre shel, 1904: 199, pl. 3, figs. 1-8; Lamy,

1906: 3; Strebel, 1908: 37; Caste distes 1970: 74, pl. 5,

fic. |

?Trophon antarcticus Philippi, 1868: 225 (sensu Tryon 1880),

T. lacineatus Martyn. Sowerby IL, 1880: pl. 404, fig. 13.

T. (Stramonitrophon) laciniatus (Martyn).—Powell, 1951: 156,

fig. L, 86.

T. (Stramonitrophon) lamellosa (Ginelin)—Dell; 1971: 212.

T. plicatus (Lightfoot, 1786).—Rehder, 1967; 20; Cernohorsky,

1977: 117, fig. 1s Vokes, 1991: 7, fig.; 1991b: 9, figs. 14—

16; 1992: 3 bec 2. b, e, d.; Castellanos and Landoni, 1993:

5, figs. 16, 15, 22

42, pl. 79, fig. LT. no-

THE NAUTILUS, Vol. 119, No. 2

Description: Shell of medium to large size (to 50

mm), smooth, fusiform, thin, somewhat "challey: proto-

conch smooth of 2% whorls; teleoconch of 6 shouldered

whorls, spire less than % of total shell height. Spire angle

about 50°; suture impressed; subsutural ramp straight,

aperture small, ovoidal, interior pale brownish; anterior

siphonal canal long (more than half height of the aper-

ture), narrow, curved, open; umbilicus absent; outer lip

rounded with reflected edges; inner lip curved, adpres-

sed. Axial ornamentation of irregular, low lamellose var-

ices in the first whorls, that become 8-10 real lamellae

in the last ones. Lamellae growing over the entire whorl,

but attached to the shell, sometimes curving adaxially.

Lamellae ending in a peripheral spine, in some speci-

mens growing adapically. Spiral ornamentation poorly

developed to almost smooth, sometimes consisting of 6

weak cords on the base of the last whorl and the back

of the siphonal channel. Regular, very weak growth lines

present throughout shell.

Shell sitvastractune composed of two layers; inner-

most layer (25% of shell thickness), composed of cola-

brally aligned crossed lamellar aragonite, outer layer

thick (75% of shell thickness) with amorphous calcite

(Figure 42).

Operculum oval, with terminal nucleus. External sur-

face covered by concentric, irregular, growth lines. Inner

surface attachment area with two or three horseshoe-

shaped scars, glazed rim present in all specimens (Figure

36).

Animal of medium size relative to shell. Mantle large,

mantle roof thin, Cephalic tentacles broad in basal fae

flat, blunt, with rounded large black eyes in the middle;

mantle edge thickened, smooth; pallial organs arranged

as in other rachiglossans; brown osphradium less than

half of ctenidium length, slightly asymmetrical, with 50—

55 leaflets per side; ctenidium as wide as osphradium,

containing triangular leaflets. Hypobranchial gland

brownish and inconspicuous, rectum to the right of hy-

pobranquial gland.

Proboscis pleuroembolic, long, broad. Radular ribbon

extending beyond rear of buccal mass, long (0.69 * ap-

erture height vs. 0.78 * aperture height in Trophon gev-

ersianus). Esophagus loops beneath buccal mass, where

esphagus receives embedded ducts of salivary glands an-

terior to a small Leiblein valve. Esophagus curving dis-

tinctly and running along entire side of gland of Lei-

blein. Esophageal ¢ olands ( “Glandule framboise’ *) whitish

in color, posterior to nerve ring slightly marked on the

external side of the esophagus. Large s saliv ary glands en-

veloping Leiblein valve and accessory salivary glands. Ac-

cessory salivary glands distinctly large, sometimes dark-

colored, tubular, compact, coiled, embedded in saliv. ary

glands. Gland of Leiblein conspicuous, brown, ending in

a medium size blind duct and very small ampulla.

Radula rachiglossi in with rachidian teeth wide (~ 150

um), central cusp thin, large; lateral cusps wider and

slightly shorter than central cusp, with inner edge

straight; with sharp straight denticle in upper third of

inte al edge of lateral cusp, external edge with 6 very

G. Pastorino, 2005 ase 6]

Figures 22-36. Trophon plic atus (Lightfoot, 1786). 22-24. MACN-In 36033, Bahia Ensenada, Ushuaia, Tierra del Fuego, Ar-

gentina. 25-26. MACN-In 9032-16, Comodoro Rivadavia, Chubut province, Argentina. 27. BMNH 19990354, 45°55.219' S

73°39.522' W, intertidal. Islet NE shore of Isla Huemules, Golfo Elefantes, Chile. 28-29. AK 133035, Station WS7S8 Discovery

15°07" S, 65°W. 30-31. MACN-In 36034, Punta Pefias, San Julian, Santa Cruz province, Argentina. 32. Same lot as Figure 6. 33.

MLP 27202. Bahia Golondrina, Ushuaia, Tierra del Fuego. 34-35. Two protoconchs, scale bars 100 jum. 34. USNM $70535

52°30’ S, 67°14’ W, in $2 m. 35. USNM 901620, 53°35’ S, 69°45’ W, 1 D, R/V Hero Cruise 692, Sta. 404, 37-46 m. 36. MLP

979

27232. operculum. external (left) and internal (right) views, scale bar = 1 cm. Scale bar for all shells = 1 cm

eee)

Page 62 THE NAUTILUS, Vol. 119, No.

Figures 37-42. Trophon plicatus (Lightfoot, 1786). 37. MLP 27232, Dorsal view of radular ribbon. Scale bar = 50 jm. 38.

MLP 526, San Julian, Punta Penas, | March 1924, rachidian teeth. Scale bar = 30 jzm. 39. Variations of rachidian teeth, Rocamora,

Ushuaia, 1 A, 8-10 m (MACN-In 36053). Scale bar = 30 wm. 40. MLP 272

27202, Bahia Golondrina, Ushuaia, Tierra del Fuego,

Argentina, detail of the rachidian teeth, Scale bar = 20 wm. 41. Same specimen as 40, lateral view of rachidian teeth, scale bar =

30 pm. 42. USNM 870535, 52°30’ S, 67°14’ W, in 82 m, shell ultrastructure, fracture surface commarginal. Scale bar = 30 wm.

G. Pastorino, 2005

well defined denticles, always present. Base of rachidian

tooth sinuous, sliding bene rath base of next tooth. Mar-

ginal area with single conspicuous cusp. Lateral teeth

with single, long and narrow cusp, slightly attached basal

plate. Cusps of rachidian teeth pyramid-like in lateral

view (Figures 38-40)

Male and female organs as in Trophon geversianus

(see Harasewych, 1984).

Type Locality: Islas Malvinas.

Type Material: Probably lost. According to Dance

(1966) part of the materi: il from the Portland ¢ Jatalogue

is in London (BMNH), however this is not the case of

T. plicatus.

Additional Material Examined: 52°30’ S, 67°14’ W,

2 D, RV ELranin Cruise 11, Sta. 980, 14 February

1964, $2 m (USNM 870535); 53°35’ S, 69°45’ W, 1D,

R/V Hero Cruise 692, Sta. 404, 37-46 m (USNM

901620); 52°56’ S, 75°00’ W, 1 D, R/V ELTANIN Cruise

Ll, Sta. 958, 5 February 1964, 92-101 m (USNM

$70423):; 53°06' S, 67°04’ W. 3D, B/V Hero Cruise 702,

Sta. 450, 5 March 1970, 86 m (USNM_ 901622):

53°39'24" S$. 70°55'30" W, 1 D, R/V Hero Cruise 702,

Sta. 467, 25 April 1970, 24 m (USNM 901623); 52°35’

S, 65°0S’ W, 1 D, R/V ELTanin Cruise 11, Sta. 976, 13

February 1974, 128 m (USNM 870525): 46°04’ S, 83°55’

W, 1 A, R/V ELTANIN Cruise 25, Sta. 326 9 October

1966, 298 m (USNM 901621): 53°48.7' S, 70°24.1' W,

1 D, R/V Hero Cruise 702, Sta. 489, 29 April 1970, 13—

IS m (USNM 901624); 53°32’ S, 64°57’ W, 2 A, R/V

ELTANIN Cruise 11, Sta. 974, 12 February 1964, 119-

124 m (USNM 881960): 53°39’ S, 70°55.5' W. 1 A. R/V

Hero Cruise 702, Sta. 466, 20 m (USNM 901754): Ba-

hia Golondrina, Ushuaia, 1 A, (MLP 2 ae Rocamora,

Ushuaia, 1 A, S-10 m (MACN-In 36053); Bahia Ensen-

ada, Ushuaia, Tierra del Fuego, en 36033):

45°07’ S, 65°W, 1 A, py eee WS7SS, 13 De-

cember 1931, $2-88 m (AK 133035) illustrated o figs,

28-29: Comodoro Rivadavia, 17 D ie N-In 9032-16):

54°34’ S, 64°00'18” W, 1 D, 1 A, B/V Hero Cruise 715,

Sta. 870, 24 October 1971, 84 m (USNM 881128): Bahia

Buen Suceso, | A, 23 October 1941, (MLP27230): Cabo

Colnett, Isla de los Estados, 1 A, R/V Hero Cruise 712,

Sta. 853, 20 October 1971, 91 m (USNM 869720): 2 D,

7S m (USNM 96176); Orange Harbor (USNM 5676);

Paso Richmond, Tierra del Fuego 55 m, (MACN-In

24940): 55°7’ S, 66°33’ W, 82 m (MACN-In 23938): Ti-

erra del Fuego (MACN-In 5240-2); Punta Penas, San

h ilidn, Santa Cruz eet Argentina, 4 A (MLP

27232); Punta Penas, San Julian, Santa Cruz, Argentina,

| A (MLP 27212); Puerto San Julian, Santa Cruz prov-

ince, Argentina (49°15’ S, 67°39’ W), 4 A, 2-3 m

(MACN-In 36034): 45°55. 219' S, 73°39. 599! W intertid-

al, Islet NE shore of Isla Huemules, Golfo Elefantes,

Chile (BMNH 19990384).

Distribution: This is a typical species from the Ma-

gellanic province. It has been recorded from Peninsula

Page 63

Valdés to Tierra del Fuego in Argentina and Chile to

49°S in the north (Reid and Osorio, 2000).

Remarks: The authorship of the name according to

Dance (1962) and Rehder (1967) should be oF to

J. Lightfoot, the anonymous compiler of the “Catalogue”

where the name was used for the first time, not to So-

lander.

There is an interesting range of conchological varia-

tion in Trophon plicatus, from almost smooth specimens

to highly lamellate. However, the profile is always slen-

der. Trophon bahamondei McLean and Andrade, 1982,

is a morphologically similar species, with peripheral

spines and shallow lamellae. On the other hand, F ba-

hamondei has no spiral ornamentation and Trophon pli-

catus has 6 weak cords on the base of the last as and

the back of the siphonal canal.

The gross anatomy is that customary for most Pata-

gonian Trophoninae. However, a distinctive anatomical

feature is the morphology of the accessory salivary

glands, which are compact, tubular, somewhat coiled and

large, and completely embedded in the salivary glands.

Most Pat: wonian Tega? (e.g. T. bahomanda as

well as T. geversianus), have small, kidney-shaped, ac-

cessory saliv ary glands.

Radular features of 7. plicatus are clearly different

from T. bahamondei (see Figures 26-27). The latter has

a distinctive small cusp on the outer margin of the lateral

cusp on the rachidian teeth. In addition, the rachidian

base is thinner and wider.

Trophon antarcticus Philippi, 1868, is probably a syn-

onym according to Tryon (1880); however, the type ma-

terial is missing and the description is rather obscure.

Powell (1951) described the subgenus Stramonitro-

phon to include only T. plicatus {(as T. laciniatus Martyn,

nomen nudum rejected by Opinion 456 (ICZN, 1957)]

a species with Stramonita-like radula, i.e., rachidian

teeth with three cusps, where the marginal side of the

lateral cusps bears several en This radula illus-

trated by Powell (1951: 194, fig. L, 86) was dissected

from the spe cime n illustrated in 98 29. The radular

morphology of T. plicatus is different from that of T.

geversianus. However, Coronium coronatum (Penna-

Neme and Leme, 1978) and T. acanthodes Watson,

1882, bear the same denticles on the lateral cusps of the

rachidian teeth. This appears to be a common feature in

several Trophoninae from the southwestern Atlantic.

The shell of Trophon iarae Houart, 1998, shows some

similarity with some specimens of T. plicatus. It was

based on only two specimens collected . fishing boats

apparently from Brazil and off U Jruguay. The anatomy

and soft parts of T. iarae remain unknown. The radula

apparently has been illustrated by Calvo, 1987 (although

according to Houart, 1998: 127, there is no certainty

about the ide tity of the specimen from where this r: ad-

ula was taken). In any event, this illustration does not

allow for a detailed comparison with other species. Fur-

ther comments about the validity of this species or its

Page 64

THE NAUTILUS, Vol. 119, No. 2

affinity with other species of Trophon mostly depends

on anatomical data which are not available.

Houart (1998: 127) mentioned the specimen of T. pli-

catus illustrated by Cernohorsky (1977) from Lively Is.

(Malvinas Is.) as belonging to his new species, T. iarae.

However, Cernohorsky’s specimen could be easily in-

cluded in the range of geographic distribution and mor-

phological variation of T. plicatus. He also illustrated two

specimens of T. patagonicus as T. plic atus (Figures 5-10

in Houart, 1998).

Trophon patagonicus (VOrbigny, 1839)

(Figures 43-65)

Murex patagonicus dOrbigny, 1839: pl. 62, figs. 2-3; 1941: 452

non Fusus patagonicus Sowerby, 1846 (=Trophon).

Trophon necocheanus thering, 1907: 404, pl. 16, fig. 106.

lie laciniatus ( Martyn). Carcelles, 1946: 70-72, figs. 6

a, b, 7 a-d, 8.

Trophon plicatus (Lightfoot, 1786),—Rios, 1985: 88, : 31, fig.

386; 1994: 114, pl. 37, fig. 453; Houart, 1998: 130, figs.

8-10 non Lightfoot, 1786.

Description: Shell large (up to 72 mm), biconic,

heavy, chalky or glossy we sometimes brownish grey,

opaque; protoconch smooth of 2% (~2.0 X 1.5 mm)

asymmetrical whorls; transition to teleoconch well de-

fined; teleoconch of 7 shouldered whorls, spire less than

¥% of total shell height. Spire angle about 70°; suture

impressed: aperture oval, interior ‘glossy white; anterior

Siphonal Canal rather long tor ve genus (ha reigh of

iphonal | rather long for tl (half height of

aperture), narrow, open; pse sidouenbiens deep and

widely open; outer lip reflexed. Axial ornamentation of

irregular strong, sharp lamellae covering entire whorl

surface, numbering 9-12 in the last whorl Continuum

of morphologies present from completeh sly smooth shells

with no axial ornamentation (Figures 53-56), to some

incipient lamellae along periphe ral keel (Figures 45—50),

to strongly developed lamellae (Figures 43-45). Spiral

ornamentation missing, except on first teleoconch whorl

which bears 3—4 invariably present spiral cords (Figures

62-63). Irregular growth lines present throughout “shell

surface. Some uncommon specimens have about three

weak greenish or dark spiral lines on the last three or

four whorls.

Shell ultrastructure arranged nieen’y of two layers;

innermost layer (25% shell thickness)

crossed lamellar aragonite, outer layer very thick (75%

shell thickness) of amorphous calcite (Figure 61).

Operculum oval, with terminal ep sus. External sur-

face covered by concentric, irregular, growth lines. Inner

surface attachment area with two or three horseshoe-

shape scars.

Rachiglossan radula with rachidian teeth distinct, cen-

tral cusp thin, large; lateral cusps slightly shorter than

central cusp, sharp straight denticle in the upper third

of the internal edge of the lateral cusp, external edge of

lateral cusps smooth. Base of rachidian tooth sinuous,

sliding beneath base of next tooth. Marginal area with

single conspicuous cusp. Lateral teeth with single, long

compose dof

and narrow cusp, attached to basal plate (Figures 64—

65).

Penchaszadeh (1976) described the egg capsules of T.

patagonicus {identified as T. laciniatus (Martyn) and T.

varians (dOrbigny)].

Type Material: Four syntypes are housed in the Nat-

ural History Museum, London under the number

1854.12.4.538, from “Baie de San Blas, Patagonie”. One

of them is herein illustrated (Figures 43, 44),

Additional Material Examined: Puerto Quequén,

Buenos Aires province, 3 A (MLP 26309); Miramar,

Buenos Aires province, | A, 1 D (MLP 417); Necochea,

Buenos Aires province, 3 A, 30 m (AMNH 173640); Ne-

cochea, | A, 30 m (AMNH 181220); Puerto Quequén

(ANSP 236034 and 236032): 18 miles off Puerto Que-

quen, 4 D (MACN-In 20441); 37°20’ S, 56°50’ W, 4 D,

55 m (MACN-In 15104); Mar del Plata, Buenos Aires

province, Argentina (MACN-In 10289; 12902; 2 D, 45

m, 9361-51: 7 D, 11374; 11587; 115S87-1; 4 D, 10249:

10320: 2 D, 10749: 11118; 3 D, 12066; 3 A, 25775; 9361-

53: 5 D, 10290; 10190; 10248; 12216): 36°25’ S, 54°38"

W, 54 m, 1 D wet 93426): 34°40’ S, 52°18’ W, 1

A, 100 m (MACN-In 23491); Fondos de Querandf,

Buenos Aires province, 5 A (MACN-In 14334); 25 miles

off Puerto Quequén, 22 D (MACN-In 21138); Fondos

de Querandf, | D (MACN-In 25774); Necochea, 2 D,

40-50 m (USNM 710024); Mar del Plata, 4 D, 1 D

(USNM 568240: USNM 346826): off Necochea, 1 A, 30

m (USNM 876123, Bledsoe collection); Mar del Plata,

1 D (USNM 363768); Uruguay, 1 D (USNM 346786);

1 D, 36°30" S, 54°44" W, 26 m (MACN-In 24259).

Distribution: This species is common on the mussel

banks off southern Uruguay and Buenos Aires province

in depths of 25-40 m.

Remarks: = Trophon patagonicus is a variable species,

endemic to the littoral of Buenos Aires province and the

Uruguayan coast. The species has had quite a confusing

taxonomic history. Those specimens with well-dev eloped

lamellae have been usually identified as T. plicatus, and

those with smooth shells and no omamentation as T.

geversianus or T. varians. Specimens collected on the

same location were identified as two different species

according to the presence or absence of lamellae. How-

ever, some specimens (Figures 50-52) are clearly inter-

mediate forms, and no another anatomical feature seems

to separate them. There is no reason to consider them

as two different species. The whole lot of specimens

a d by Carcelles (1946) shows clearly that he con-

fused dOrbigny’ s species with T. plicatus. Such a mis-

take was alos made by Rios (1985, 1994) and several

other authors. Trophon plicatus is a typical magellanic

species with a thinner and more cylindrical profile, me-

dium size (never reaching more than 50 mm. high).

Specimens of T. plicatus here identified were never col-

lected at latitudes north of 45°S. Its protoconch is always

smooth and with fewer whorls, and the lamellae along

the keel usually develop into strong triangular projec-

G. Pastorino, 2005 Page 65

Figures 43-59. Trophon patagonicus (d¢ Irbigny, 1539). 43-44. Syntype BMNH 1554.12.4.538, Bahia San Blas, Argentina. 45-

46. MACN-In 36040, Puerto Quequén, Buenos Aires. 47-49. MLP 417, Miramar, Buenos Aires, in 54 m. 50-52. MACN-In 12066

Mar del Plata, Buenos Aires. 53-56. MACN-In 36031, Puerto Quequén. 57-59. MACN-In 21047, “Carmen de Patagones?”

Buenos Aires. Scale bar for all figures = 1 cm

SEE

Page 66

THE NAUTILUS, Vol. 119, No.

Figures 60-65. Trophon patagonicus (WOrbigny, 1839). 60. Penis, critical-point dried. Lateral and frontal view. Scale bar = L000

wm. 61. Shell ultrastructure, fracture surface commarginal. Scale bar = 300 zm. 62-63. MACN-In 11374, protoconch, three views,

Mar del Plata, Buenos Aires province. Scale bar for all figures = 600 pm. 64. Dorsal view of radular ribbon. Scale bar = 50 jm.

65. Lateral view of rachidian teeth. Scale bar = 50 jm

tions approximately parallel to the coiling axis. The ap-

erture is larger and subcircular in T. patagonicus, very

different from the smaller and almost circular one on T.

plicatus. Trophon geversianus usually has strong spiral

ornamentation on the entire shell surface, a feature also

observed in T. varians

Ihering (1907) described TL necocheanus based on

(Quaternary?) specimens collected at Necochea, Buenos

\ires province, Carmen de Patagones, Buenos Aires

province, and Sierra Laziar, Santa Cruz province (“For-

mation araucanienne’). Most of the type material is lost,

but the remaining types fall within the range of variation

of T. patagonicus. One of the specimens, from a lot of

three, with uncertain locality (MACN-In 21047, rela-

beled “Carmen de Patagones?”) is here illustrated (Fig-

ures 57, 59). This specimen was acquired through an

exchange with Museu Paulista, S40 Paulo, Brazil (where

Ihering worked for most of his professional life) and is

G. Pastorino, 2005

Page 67

part of the original type series. All of the specimens fit

well in the original description of T. patagonicus and

there is no doubt that is the same species described by

d Orbigny.

Sowerby I (2! of name) described in 1846 Fusus pa-

tagonicus from the Tertiary of San Julian (Santa Cruz

province, Argentina), a species that clearly belongs in

Trophon. Therefore, as the two species are quite distinct,

dOrbigny’s name has priority over Sowerbys (Griffin

and Pastorino, 2005).

Trophon acanthodes Watson, 1882

(Figures 66-78)

Trophon acanthodes Watson, 1882: 386; 1886: 166, pl. 10, fig.

6; Cemohorsky, 1977: 112, fig. 9 (holotype); Pain, 1980:

S, fig.; Rios, L985: SS, pl. 31, fig. 358; Castellanos, 1986:

22, fig: C Castellanos and Landen 1993: 8, pl. 2, fig. 28

Fusus haenhones (Watson).—Carcelles, 1947: 12, pl. 2 figs. 1,

2; pl. oi figs. 3, 4 (not figs. 5, 6 which is Corontin coron-

atum).

Pagodula acanthodes (Watson).—Kaicher, 1980: fig. 2599.

“Fusinus” acanthodes (Watson) —Calvo, 1987: 153, fig. 127.

“Trophon™ acanthodes Watson.—Rios, 1994: 37, pl. 37, fig. 486.

Description: Shell large, about 125 mm high, thick,

slender in profile, ch alky white: protoconch worn in all

specimens; teleoconch of 7 shouldered whorls: spire less

than % of total shell height. Spire angle about 45°; suture

impressed, subsutural shelf oblique, somewhat convex,

aperture small, subcircular, interior glossy white; ante-

rior siphonal canal very long (longer than aperture

height) deep, straight or gueved always open; outer lip

rather reflected, rounded: inner lip adpressed. Axial or-

namentation of 12-15 regularly spaced, axial lamellae,

slightly raised, almost attached along the entire whorl,

producing open, conspicuous, regularly spaced spines at

periphery; spines becoming more closely packed on last

whorl. Spiral ornamentation of 4-5 weak rounded

threads on the lower part of the first whorls, becoming

more than 20 in last one. Growth lines present through-

out shell, producing wrinkly surface by intersection with

spirals.

Operculum elliptical, nucleus terminal, older speci-

mens tear-shaped. External surface covered by concen-

tric, irregular, growth lines. Inner surface with a con-

spicuous margins al rim; attachment area cov ering almost

the whole operculum but the rim; oreeshGe- shaped

scars present.

Rachidian teeth wide (~130 ym), central cusp large:

lateral cusps wider and shorter than central cusp, inner

edge with an almost obsolete denticle (particularly in

adult specimens), external edge with 5-6 denticles de-

creasing in size towards the edge. Base of rachidian

tooth curved, somewhat sinuous, sliding beneath base of

next tooth. Marginal area with single oe Lateral teeth

with single, long cusp, slightly atached | yasal plate.

Shell ultrastructure composed of two layers: inner-

most layer (55% of total thickness of shell) composed of

crossed lamellar aragonite, outer layer (45%) of amor-

phous calcite.

Animal as in other Trophon species but with some size

differences. Tentacles long and well defined, joined at

base; eyes deeply marked. Salivary and accessory salivary

glands as in T. plicatus. Osphradium a bit more than one

third of penile length, asymmetrical, with about 112

leaflets, ctenidium w ith about 250 leaflets twice as large

as osphradium. Typical pleuroembolic proboscis, shorter

than in T. geversianus, valve of Leiblein also smaller:

gland of Leiblein large with a long blind duct. Radula

long; digestive tract with the usual loop on the left side

of the gland of Leiblein, before the duct to the gland.

Male and female organs similar to T. geversianus.

Type Material: BMNH_ 1887.2.9.568, holotype

50°8'30" S, 74°41’ W, 229 m (illustrated by Cernohorsky,

1977, fig. 9).

Additional Material Examined: 52°53’ S, 74°05’ W,

3 D, B/V ELTANIN Cruise 23, Sta. 1605, 1 April 1966,

522-544 m (USNM 901756): off Mar del Plata, 5 A,

January 1962 (MLP 26283); 52°41’ S, 74°35’ W, 1 D,

R/V ELTANIN Cruise 21, Sta. a 6 January 1966, 188—

247 m (USNM 870115); 51°56’ S, 56°39’ W, 1 D, R/V

ELTANIN Cruise 7, Sta. 557, . March 1963, 855-866 m

(USNM 870345): 52°52’ S, 75°18’ W, 1 D, R/V ELTANIN

Cruise 21, Sta. 288, 119-329 m (USNM 901758): 52°51’

S, 74°13’ W, L D, B/V Etrantin Cruise 21, Sta. 291, 523-

539 m (USNM 901757); 52°53’ S, 74°05’ W, 2 D, R/V

ELTANIN Cruise 23, Sta. 1605, 1 April 1966, 522-544 m

(USNM 897615): 40°15" S, 57°40' W, 1 A (MACN

18425): 40°03’ S, 57°00’ W. 1 D, 50 fathoms (91.5 m)

(MACN 15699): 30 miles off Mar del Plata (ANSP

262989 and 236028); off Mar del Plata, 1 A, 2 D (MACN

17671 and 16449): 39°26’ S, 56°40’ W, 1 A, 1 D, 90 m

(MACN 17040); 39°02’ S, hai W, 1 A, 1 D; East of

Punta Médanos (39°—39°30' S), 4 D, 50 fathoms (91.5

m) (MACN 14386); 39°55’ _ 57°50’ W, 1 D,51 fathoms

(93 m): 38°25' S$, 56°30’ W,3 D (MACN 16798); 39°35’

S, 57°10’ W, 1 A, 1 D (MACN 18342); 39°50’ S, 57°18’

W, 4 A, 52 fathoms (95 m) (MACN 21741); 39°00’ S,

57°10’ W, 2 D, 45 fathoms (S2 m) (MACN 15216);

37°15’ S, 54°50’ W, 1 A, B/V ALDEBARAN, March 2000,

commercial otter trawl, 111 m (MACN-In 36032):

36°37' S, 54°14’ W, 1 A, B/V AcpeBaran, March 2000,

otter trawl, 104 m (MACN-In 36031): 39°02’, 57°02’ W,

2 D, 46 fathoms (S4.1 m) (MACN-In 25118); 37°35’ S,

54°55’ W. 1 D, 105 fathoms (192 m) (MACN-In 25165-

2).

Distribution: Off Rio Grande do Sul state in Brazil

(Rios, 1994), Uruguay, to Tierra del Fuego, Argentina.

Remarks: The shell of Trophon acanthodes is some-

what similar to that of Coronium coronatum, which in fact

could be granted that generic position. The operculum

and the radula are different in both species. The oper-

culum is somewhat triangular in Coroniwm with a weak

rim instead of the characteristic thicker one of the Tro-

phon species. The radula of Trophon acanthodes presents

the intermediate cusp between the central and the lateral

Page 68 THE NAUTILUS, Vol. 119, No. 2

Figures 66-76. a ee Watson, 1582. 66-68. MACN-In 25118, 39°02", 57°02" W, 46 fathoms (84.1 m). 69-71.

MACN-In 36031, 37°15’ S 50’ Win 111 m. 72-73. MLP 26283, Mar del Plata. Scale bar for all shells = 1 cm.74. Two views

of the operculum of the specimen in Figures 69-71. Scale bar = 1 cm. 75. Ultrastructure of the shell. Scale bar = 200 jzm; detail,

large quadrangle. Scale bar = 50 zm. 76. Penis, critical-point dried (scale bar = 600 wm) with detail of the efferent conduct (scale

bar 150 jum

one of the rachidian teeth almost obsolete, while in Co- The protoconch in Coronium is multispiral and ex-

ronwm coronatum it is very conspicuous Castellanos tremely pointed, distinctive of the genus. All the studied

1986) drew a stereotyped radula of 7 acanthodes where specimens of T. acanthodes are worn; however, some of

denticles are wrongly placed on the margin of the rachi- them show the sinuated line of a typical Trophon pro-

dian tooth instead of the inne edge of the lateral cusp toconch

G. Pastorino, 2005

Page 69

Figures 77-78.

Trophon pelseneeri Smith, 1915

(Figures 79-93)

Trophon pelseneeri Smith, 1915: 92, pl. 2, figs. 6, 7; Rios, 1994:

115, pl. 38, fig. 484; Houart, 1991: 33.

Trophon sp.—Carcelles, 1944: 253.

Trophon orbignyi Carcelles, 1946: $1, pl. 12; Castellanos, 1970:

73, pl. 5, fig. 6.

Description: Shell small to medium in size (up to 35-

40 mm), fusiform, slender, chalky, pinkish with 2 weak,

diffuse reddish bands along the edge of last whorl la-

mellae; protoconcl symmetrical, cylindrical, smooth, of

2% whorls: teleoconch of 7 rectangular whorls, spire %

of total shell height. Spire angle about 45°, suture im-

pressed, subsutural shelf straight, aperture small, circu-

lar, interior glossy white; anterior siphonal canal long,

open, straight. with the tip adaxiz il curved, equal to

aperture height: umbilicus slightly pe n, sometimes only

a narrow slit; outer lip reflexed to form lamellae; inner

lip curved. with white adpressed callus. Axial ornamen-

tation of 9-11 regular, low lamellae, covering entire

whorl surface; lamellae forming a back-turned spine at

periphery. Spiral ornamentation of 2 to 3 very weak

threads in first whorls be coming more than 7 in last one,

sometimes only visible along edge of last whorl] lamellae.

Siphonal fasciolae slightly oblique, always present.

Operculum triangular (tear- oh ape -d), with terminal nu-

cleus. External surface covered by irregular growth lines.

Inner surface attachment area with horseshoe-shaped

scars: glazed rim weak but present (Figure 85).

Radula rachiglossate with rachidian teeth wider than

high, central cusp thin, in a different plane than laterals;

lateral cusps wider and shorter than central cusp, with

inner edge oblique; sharp straight denticle in the upper

third of the internal edge of the lateral cusp, external

edge with almost obsolete denticles numbering 3-4.

Base of rachidian tooth slightly sinuous, sliding beneath

base of next tooth. Marginal area with single conspicuous

Trophon acanthodes Watson, 1882. 77. MLP 26283, off Mar del Plata Radula of a large specimen, frontal view.

Scale bar = 50 pm. 78. Lateral view. Scale bar = 50 pm.

cusp. Lateral teeth with single, long and narrow cusp,

slightly attached basal plate (Figures 92-93).

Type Material: = [T. pelseneeri] BMNH 1915.4.18.276-7,

two syntypes from west of Malvinas Is. in 229 m (Figures

79-82) and [T. orbignyi] MACN-In 24421, Puerto Que-

quén, Buenos Aires, holotype (Figures $3-S5) and 10

paratypes.

Additional Material Examined: Macaé, Rio de Ja-

neiro, Brazil, August 1969, 2 A, 55 m (USNM 846550):

Macaé, Rio de Janeiro,. Brazil, : e 30 m (AMNH

187586); Rio de Janeiro, Brazil, (AMNH 241045):

off Rio de bare iro, Brazil Ae a ); 34°48'7" S,

54°21'9" W, 1 A, R/V ALpDEBARAN, Cruise 9901, Sta. 25,

27 January idea, 25 m, with Piccard trawl (MNHNM

15400).

Distribution: Rio de Janeiro, Brazil, to Uruguay and

Buenos Aires province, Mé ilvinas Is. Rios (1994) cited it

from dredgings from 55 to 225 m off the Brazilian coast.

Remarks: This is a rare species occasionally collected

by fishermen on the mussel banks off Buenos Aires

province. It was originally described from Sta. 3S of the

British Antarctic (“Terra Nova”) Expedition, west of

Malvinas (Falkland) Islands in 125 fathoms depth. How-

ever, together with this species, the author mentioned

several others not reported before or since from. this

latitude, but sae occur at Station 42 of the same ex-

pedition, off Rio de Janeiro. This leads to the supposi-

tion that the material from these stations could have

been mixed up (Searabino, 2003; 199).

Trophon amettei Carcelles, 1946

(Figures 94-100)

Trophon amettei Carcelles, 1946: $4, fig. 13; Carcelles and Wil-

liamson, 1951: 287

Description: Shell small in size (up to 30 mm), fusi-

Page 70 THE NAUTILUS, Vol. 119, No. 2

Figures 79-93. Trophon pelsenceri I. A. Smith, 1915, 79-80. BMNIT 1915.4.18.276-7 syntype. 81-82. BMNH 1915.4.18.276-7

other syntype. 83-85. T. orbignyi Carcelles, MACN-In 24421 holotype. 86-87. MNHNM 15400, B/T ALDrBaran, 34°d48'7" S

54°21'9" W, in 25 m. Seale bar lL cm. 88. Two views of the operculum of the specimen in Figures S6—-S7 Seale baa 2mm

$9-90. USNM 846550 apical and lateral view of the protoconch Seale bars 100 and 300 pom respectively 91. Detail of the

ultrastructure of the shell. Scale bar 100 jum. 92-93. Dorsal and lateral view of the radula of the specimen in Figures S6—S7

‘ le har (0)

Da ) Jem

G. Pastorino, 2005

Figures 94-100.

form, biconic, chalky whitish; protoconch elongate,

smooth, of

dered whorls, spire less than % of the total shell height.

2% whorls; teleoconch of 6 strongly shoul-

Spire angle about 45°, suture impressed; subsutural shelf

short but straight, aperture small, ovoid. interior glossy

white: anterior siphonal canal comparatively long, almost

same height as aperture, open and straight: umbilicus

closed, inner lip adpressed. Axial ornamentation of §

regular concave ee per whorl. Lamellae growing

attached to entire whorl surface and producing concave

spine along periphery. Spiral ornamentation of 2 cords

in first teleoconch whorls that soon become obsolete on

subse oe nt whorls. Last whorl with 6—10 cords on lower

part. Shell ultrastructure composed of two layers, similar

to T. geversianus. Operculum and soft parts unknown

Type Material: Holotype and 15 _ paratypes MACN-

In 23510) all from 45°09’ S, 66°27’ W anchorage Res-

tinga Aristizdbal, Chubut province, in 8 fathoms (11.28

m). rocky bottom associated with the

; calyptraeids Cre-

pidula cachimilla and Calyptraea pileolus

Remarks:

type locality

This is a rare species known only from the

concave lamellae, are 1 fact. unusuc il f tor the genus. The

has not been found again. The regular

Trophon amettei Carcelles, 1946, 94-96. MACN-In 23810, holotype, 45°09' S_ 66°27’ W in 11.28 m. 97-98.

MACN-In 23810, paratype. Scale bar = 1 cm. 99. MACN-In 23510, ultrastructure of the shell. Scale bar = 30 pm. 100. MACN-

In 23810, protoconch, uncoated SEM picture. Scale bar = 300 jzm

protoconch resembles those usually found in the genus

Fuegotrophon, however the typical fimbriate spiral or-

namentation is absent. Trophon pelseneeri has a com-

parable profile, however it is easily segregated: where

the latter has lamellae forming a back-turned spine at

periphery, T. amettei presents a characteristic and

unique concave lamellae. In addition T. pelseneeri has a

slightly open umbilicus while in T. amettei it is invariably

closed. Examination of the radula may beget a new ge-

neric allocation for this species

Trophon clenchi (Carcelles, 1953)

(Figures 101-114)

Murex clenchi Carcelles, 1953: 7, figs. 23-28: Castellanos

1970; SO, pl. 5, fig. 7; Fair, 1976: 31, fig. 15; Vokes, 1992b

24: Rios, 1994: 115

Poirieria (Poirieria) clenchi (Carcelles Vokes, 1970: 18

Sarina a Shell medium in size, delicate, thin,

che lly, tré isluce nt white prot toconch known onl from

the holot type, asyinine tric al. clobose, of 2 A 2 whorls: t te-

leoconch of 5 shouldered whorls; spire short, less than

% of total shell height. Spire angle about 45° (without

subsutural shelf shor

spines); suture impressed

a a SEE

Page 72

THE NAUTILUS, Vol. 119, No. 2

Figures 101-110. Trophon clenchi (Carcelles, 1953). 101-104. MACN-In 25146, holotype, 38°24! $, 55°36’ W in 89.61 m. 105-

107. MACN-In 36269, aproximately 45°10’ S$, 57°20’. Scale bar = 1 cm, 108. Holotype, protoconch. Scale bar = 1000 jzm. 109.

External and internal views of operculum of specimen in Figures 105-107, Scale bar = 0.5 cm 110. MACN-In 25146, paratype.

straight; aperture suboval, interior glossy white; anterior

siphonal canal very long (longer than the aperture

height deep and slightly curved in the beginning and

then straight open but narrow: outer lip sharp, inner lip

somewhat protruding adpressed. \xial ornamentation of

7-9 regular axial lamellae growing across entire whorl

surface, but attached to shell producing open, long and

regularly spaced spines along periphery Spiral Ormamen-

tation of 4—5 rounded cords on lower half of first whorls

becoming more than 20 in the last. Delicate growth lines

present on entire shell surface.

Shell ultrastructure composed of two layers following

the common pattern of the genus: innermost layer (40%

of total thickness of shell) composed of crossed lamellar

aragonite, outer layer (50%) of amorphous calcite.

Sometimes, depending on the fracture mode, a basal

aragonitic layer can be observed.

Radulae rachiglossate, rachidian teeth with three me-

lant

Pp.

. Pastorino, 2005

Figures 111-114. Trophon clenchi (Carcelles, 1953). 111. USNM 901774, 51°58" S, 56°38" W, R/V ELTaNIN Cruise 7, Sta. 558,

646-S45 m, ultrastructure of the shell. Scale bar = 100 pm. 112. Poirieria zelandica, ultrastructure, commarginal fracture surface.

Scale bar = 100 wm. 113. Trophon clenchi MACN-In 36269. Lateral view of radula ribbon. Scale bar = 100 jm. 114. Dorsal

view of radular ribbon. Scale bar = 50 juin.

dian cusps, the central one the larger and the lateral

ones with a denticle, attached to the upper third of the

interior margin of the lateral cusp; external margin with

2-3 obsolete denticles. Rachidian base sinuous, with the

base offset under the proximal tooth. Marginal cusps sin-

ale, never bifid. Lateral teeth curved, thin, with attach-

ment area also thin (Figures 113-114).

Operculum triangular or suboval tear-shaped, attach-

ment area elliptical, with horseshoe shape scars.

Type Material: 38°24’ S, 55°36’ W, off Mar del Plata,

in $9.6 m, holotype and paratype (MACN-In 25146).

Additional Material Examined: 51°58’ S$, 56°38’ W,

3 D, R/V ELTANIN Cruise 7, Sta. 558, 646-845 m

USNM 901774), 41°51’ S, 57°34" W, collected by Uru-

cuayan fishermen, June 2002, 1062 m, 1A, MACN-In

36270; approximately 45°10’ S, 57°20’ Uruguayan fish-

ermen, | A, MACN-In 36269: 54°50’ S, 63°50.5' W. 2.5

miles south Punta Fallows, Isla de los Estados, Tierra

del Fuego, 1 A, R/V Hero Cruise 715, Sta. 715/879, 28

October 1971, in 342-353 m (LACM 71-331).

Distribution: Known from off Buenos Aires province,

Patagonia and Isla de los Estados, in 90-1050 m depth.

Remarks: Carcelles (1953) described Murex clenchi

from two shells he received from the crew of the ARA

Banfa BLANCA, an Argentine Navy ship that occasionally

collected material for the collection of the Museo Ar-

gentino de Ciencias Naturales (MACN). Both holotype

and paratype were collected from the continental shelf

off Buenos Aires province. This material remained

housed at the MACN until it was sent on loan and it

appears to have been lost for almost 30 years (see Cas-

tellanos, 1986). The return of the material to MACN

made the type material again available for studies. Per-

haps because of these facts, the ordinary quality of the

illustration and the absence of soft parts, the species was

always reluctantly considered as the southernmost rep-

resentative of the genus Murex

E. H. Vokes, in a fundamental paper published in

1970, stated that Murex clenchi belongs in the genus

Poirieria sensu stricto. Rios (1994) considered M. clenchi

Page 74

THE NAUTILUS, Vol. 119, No. 2

as an anomalous specimen of Trophon acanthodes Wat-

son, a species that slightly resembles some specimens of

M. clenchi (e.g., the paratype).

The morphology of the shell as well as the radula and

penis of Murex clenchi allow the allocation in the genus

Trophon. In fact, this was already suggested by E. H.

Vokes (1992b). After studying the material of T) acan-

thodes housed at the MACN she proposed that M. clen-

chi belongs to Trophon sensu lato and not to Poirieria.

It bears no relationship with Poirieria despite some ap-

parent shell similarity. As it is shown in Figure 112, Poir-

ieria zelandica, type species of Poirieria, Tae no calcitic

layer on the shell but a thick aragonitic one instead (Fig-

ure 112a). In contrast, the entire group of Patagonian

Trophon (T. clenchi in particular), shows different de-

grees of development of the typical calcitic layer on the

distinct species.

The range of the very few lots studied falls within the

geographic distribution of other species of Trophon. Tro-

phon clenchi was known from two quite different geo-

graphic and bathymetric areas: off Buenos Aires prov-

ince (in about 90 m) and the slope off Patagonia. A new

lot is recorded here from Isla de los Estados, which sig-

nificantly increases the range of distribution of the spe-

cies. It is possible that hike other Patagonian species of

the genus, T. clenchi would be associ ‘ated to hard bot-

toms, thus hampering the collection of material. Its fra-

gility, size, and inadequate sampling of the area adds

other reasons for the scarcity of records. It is suggested

that these variables accounts for the disjunt recorded

distribution of this species. In fact, this species started

to be repeatedly collected recently, when the fishery of

the Patagonian toothfish (Dissostichus eleginoides) on

the Argentinean slope provided the opportunity to catch

accidentally entangled specimens (F. Scarabino, pers.

comm. ).

Trophon wilhelmensis Ramirez-Bohme, 1981

(Figures 115-119)

Trophon (Enixotrophon) wilhelmensis Ramirez-Bohme, 1981:

6, fig. la, b.

Description: Shell of medium size, about 52 mm

high, slender in profile, chalky white; Sa un-

iow teleoconch of 6 shouldered whorls: spire less

than % of total shell height. Spire angle less than 45°;

suture impressed, subsutural shelf straight, aperture

small, subcircular, interior glossy white; anterior siphonal

canal very long (longer a aperture height) deep,

straight or curved, always open; outer lip ri ather reflexed,

rounded, inner lip adpresse sd. Axial ornamentation of

12-15 regularly spaced, axial lamellae, slightly raised, al-

most attache d along the entire whorl, producing open,

conspicuous, regul: urly spaced spines at periphery; spines

becoming more close ly packed on last whorl. Spirs al or-

namentation of 4—5 weak rounded threads on the lower

part of the first whorls, becoming more than 20 in last

one. Growth lines present throughout shell, producing

wrinkly surface by intersection with spirals.

Operculum elliptical, nucleus terminal. External sur-

face covered by growth lines. Inner surface with mar-

ginal rim; Adachinent area, horseshoe-shaped scars pres-

ent.

Rachidian teeth of about 90 zm wide, central cusp

large; lateral cusps shorter than central cusp, inner edge

with conspicuous de nticle, external edge with 5 denticles

of equal size. Base of rachidian tooth straight, somewhat

sinuous, sliding beneath base of next tooth. Marginal

area with single cusp. Lateral teeth thin with single, Tong

cusp, slightly attached basal plate.

Type Material: Holotype in MNHN (unnumbered),

41°51' S, 74°30'5” W West of Chiloé Island in 250 m.

Additional Material Examined: Boca del Guafo,

43°39'36" S. 73°51'11" W, southern Chile, 6 July 2002,

1 A, 200 m (MNHWN unnumbered).

Distribution: Known only from the holotype and an-

other lot, both from the same area and depth, around

the Chiloe Is., Chile.

Remarks: Trophon wilhelmensis was recently de-

scribed from Chile with no mention of the similarity

with T. acanthodes. Both species are actually quite sim-

ilar. The species, known only from the holotype and an-

other specimen, may be distinguished from T. acantho-

des mainly by the upturne cd spines (compare Figures 66—

73 with Figures 115-117). In addition, slight diferentes

in the morphology of the rachidian reat: allow for fur-

ther differentiation of the two species. Notwithstanding,

future studies of specimens from intermediate localities

could demonstrate that these represent but a single spe-

cies.

Trophon bahamondei McLean and Andrade, 1982

(Figures 120-125)

Tropnon bahamondei McLean and Andrade, 1982: 10, figs. 24—

25.

Description: Shell of medium size, up to 50 mm,

slender, with narrow profile, chalky or creamy white;

protoconch unknown (worn in all specimens); teleo-

conch of 6 shouldered whorls; spire less than % of total

shell height. Spire angle about 45°; suture impressed,

subsutural shelf well defined, straight; aperture subcir-

cular, interior chalky white; anterior siphonal canal very

long (equal to aperture height), narrow, and curved,

open; outer lip sharp, mound. inner lip adpressed. Axial

ornamentation of 11-13 regular axial lamellae growing

on entire whorl surface, but attached to the shell and

producing open, short and regularly spaced spines along

periphery. Spiral ornamentation lac king. Growth lines

present throughout shell.

Ope srculum “elliptical, nucleus terminal. External sur-

face CC wvered by concentric, irregular crowth lines. Inner

surface attachment area reaching upper side or center,

with horseshoe-shaped scars (Figure 123).

Radula with rachidian teeth very wide (~114 jzm),

with narrow base, central cusp thin, large; lateral cusps

G. Pastorino, 2005

Figures 115-119. Trophon wilhelmensis Ramirez-Bohme, 1981. 115-117. MNHNS unnumbered, Boca del Guafo, 43°39'36" S,

73°51 11" W, southern Chile, 6 July 2002, 200 m. Seale bar = 1 em. 118-119. Radula of the specimen on Figures 115-117, 118.

Frontal view. Scale bar = 50 xm. 119. Lateral view. Scale

wider and shorter than central cusp, inner edge wi

sharp. curved denticle, external edge with denticles de-

creasing in size toward lateral edge where they disa

pear. Base of rachidian tooth sinuous, inserted in part

under base of subsequent tooth. Marginal area with si

yar = 40 jem

le conspicuous cusp. Lateral teeth with single, long and

very narrow cusp, $ ightly attached basal plate. Centr

cusps of rachidian teeth curved back in lateral view (Fig-

ures 124-125

Animal very similar to T. plicatus. Osphradium le

than half ctenidium length, asymmetrical, with 60—70

leaflets. Ctenidium with 140 leaflets less than two times

larger than those of osphradium. Tentacles well defined;

eyes deeply marked. Same type of accessory salivary

glands, twisted and non-compact

Male unknown, all specimens studied were females

Female similar to T. geversianus

Type Material: Holotype from off Pichilemu, Chile

34°27' S, 340 m (LACM 1982, but apparently on loan)

and two paratypes, 34°27’ S, 71°54’ W, 25 May 1976,

200-450 m (USNM 754739).

Additional Material Examined: 31°56’ S, 71°54’ W,

off Los Vilos, Chile, 2 A, 8 March 1977, 240-350 m

(LACM 72491): off Playa Blanca, Coquimbo, Chile, 400

m (ANSP 291065): off Coquimbo, Chile (LACM 75-88

Distribution: This is a deep water Trophon known

from latitudes 30° to 34° S off the coast of Chile

Remarks: Trophon bahamondei 1S apparently a Very

Page 76

THE NAUTILUS, Vol. 119, No. 2

Figures 120-125. Trophon bahamondei McLean and Andrade, 1952. 120-122. Paratype USNM 754739, 34°27’ S, 71°54" W,

Pichilemu, Chile, in 200-450 m. Scale bar = 1 em, 123. Operculum, external (right) and internal (left) views. Scale bar = 1 cm.

124. Dorsal view of radular ribbon. Scale bar = 30 xm. 125. Lateral view of rachidian teeth. Scale bar = 30 frm.

consistent species from the morphological standpoint. It

was only recently described despite its apparently wide-

spread presence in shrimp trawls. The deeper water

habitat of species (more than 200 m) probably rendered

the species less accessible in the past.

General morphology of shell and gross anatomy shows

at first glance some similarities with those of T. plicatus;

however close examination of shell and radula confirms

the presence of a different species.

Trophon parodizi new species

Figures 126-137, Table 1)

Description: Shell medium in size (up to 2

oO mm),

very thin, chalky: protoconch smooth, of 1% (1.

1.39) very asymmetrical whorls: transition to teleoconch

well defined; teleoconch of 4 convex whorls spire less

than ¥% total shell height Spire angle about 40°; suture

ImMpre ssed aperture subovoid. interior lossy white; all-

terior siphonal canal medium in size (less than half

height of aperture), narrow, open; umbilicus absent; out-

er lip rounded. Axial ornamentation of irregular, low,

rounded ridges occupying entire whorl surface, num-

bering up to 9 in early whorls, but vanishing and un-

dulate on last whorl. Spiral ornamentation poorly devel-

oped or almost smooth, when present consisting of ob-

solete, weak cords mostly developed on last whorl. T-

regular growth lines present throughout shell.

Shell ultrastructure composed of two layers; inner-

most layer thin (35% of shell thickness), composed of

colabrally aligned crossed lamellar aragonite, outer layer

thick (65% of shell thickness) with amorphous calcite.

Operculum oval, with terminal nucleus. External sur-

face covered by concentric, irregular, growth lines. Inner

surface attachment area with two or three horseshoe-

shape scars. Animal unknown.

Radula rachiglossan with rachidian teeth distinctive,

central cusp thin, large; lateral cusps almost same size

as central cusp, sharp denticle on upper third, irregular

external denticles present. Base of rachidian tooth sin-

G. Pastorino, 2005 Page 77

Figures 126-137. Trophon parodizi new species. 126-128. USNM 896397, holotype, 54°56" S, 65°03" W, in 229-265 m. 129-

130. USNM 896397, paratype. coated with ammonium chloride. Scale bar = 1 cm. 131. Ultrastructure, fracture surface commar-

ginal. Scale bar = 20 pm. 132. USNM 896397, operculum, external view. Scale bar = 0.5 cm. 133-135. Protoconch of the

paratype. three views. Scale bar for all figures = 400 pm. 136. Dorsal view of radular ribbon. Scale bar = 40 jum. 137. Lateral

view of rachidian teeth. Scale bar = 40 wm

THE NAUTILUS, Vol. 119, No. 2

Figures 138-144.

uous, sliding beneath base of next tooth. Marginal area

with a conspicuous cusp. Lateral teeth with single, long

cusps and narrow, slightly attached basal plate.

Type Material: Holotype and one paratype from

54°56° S, 65°03° W, Le Maire Strait, R/V ELTANIN Cruise

11, Sta. 969, 10 F ebruary 1964, 229-265 m, Blake trawl

(USNM 896397) (Map 1).

Material Examined: Only holotype and paratype.

Distribution: Known only from the type locality.

Etymology: This species is named after Juan José Par-

odiz, one of the pioneers of malacology Argentina,

Curator Emeritus at the Carnegie Museum, Pittsburgh,

and a good friend.

Remarks:

within Trophon. The small, very thin shell, almost com-

Trophon parodizi is an unusual species

pletely smooth and devoid of lamellae, is an exception

for the Patagonian Trophon. However, the radula and

protoconch match those typical of the genus. The rad-

ular morphology resembles that of T. plicatus where the

denticles of the lateral cusp of the rachidian teeth are

plac ed along the entire external e dge of the lateral c usp.

In 7 parodizi the lateral cusp is thinner and the denti-

cles are more clearly separated from it and placed mainly

along the marginal area as in T. bahamondei. In addi-

tion—a character also present in T. bahamondei—most

internal denticle lies against the lateral cusp and is clear-

ly differentiated from the rest

“Trophon” malvinarum Strebel 1908, 138-139. NHRM 1041, holotype

In 23944-2, 55°07’ S, 66°33’ W. 143-144. Protoconch. Scale bar = 400 xm

»592°29' S, 60°36! W. 140-142. MACN-

“Trophon™ malvinarum Strebel, 1908

(Figures 135-144)

Trophon malvinarum Strebel, 1908: 44, figs. 16 a-c; Carcelles

and Williamson, 1951: 289: Castellanos and Landoni,

1993: 10, pl. 2, fig. 29.

Description: Shell small, up to 30 mm, thick, fusi-

form, profile slender, chalky, whitish; protoconch glo-

bose, somewhat cylindrical, with two whorls: teleoconch

of 4% shouldered whorls, spire less than % of total shell

height. Spire angle about 45°; suture impressed; subsu-

tural shelf abaxially oblique, aperture subquadrate, an-

terior siphonal canal moderately long (same height as

aperture): umbilicus closed, outer lip polygonal to

rounded; inner lip slightly curved, cas sd. Axial or-

hnamentation of S—9 re soular, distinct low varices, which

never develop into full-fledged lamellae. Spiral orna-

mentation of about 3 cords, filling interspaces between

consecutive varices, and beginning at periphery of

whorls.

Radula and anatomy unknown.

Table 1. Measurements of the type specimens of Trophon

parodizi new species in mm

Species Leneth Width Whorls

[. parodizxi

Holotype 23 L] {

Paratype Ll 10.6 {

G. Pastorino, 2005

a. > TC

| age iv

T eee aeny ~

Buenos Aires a ,

y A

Golfo

San Matias

~ -Malvinas Is.

te rt eS

v A

sar Staten Is

nce — eh

Figure 145. Map showing the type locality of T. parodizi

new species ( @ ).

Type Material: 52°29’ S, 60°36’ W, West Falkland

(Malvinas) Is., Svenska Sudpolar Expedition, 11 Septem-

ber 1902. Sta. 58, 197 m (NHRM 1041).

Additional Material Examined: 55°07’ S, 66°33’ W,

1 D. $3 m (MACN-In 23944-2).

Distribution: Around Malvinas Is. and the Magellanic

region.

Remarks: “Trophon” malvinarum is a rare species

known from only one specimen other than the holotype.

It was originally described in the genus Trophon; how-

ever, the protoconch is somewhat shorter a symmet-

rical and the shell never develops true lamellae but low

varices. It probably could be better assigned in the genus

Urosalpinx. Assessment of its real attaites must wait

until estes with soft parts preserved can be studied.

CONCLUDING REMARKS

The geographic distribution of the species of the genus

Trophon sensu stricto is restricted to the southwestern

Atlantic and the southeastern Pacific Oceans. This dis-

tribution is certainly a consequence of their larval biol-

ogy. Planktonic larval development is unknown to occur

in the genus. Moreover, the larvae do not need to move

any significant distance either for feeding or reproduc

tive purposes—those from shallow water habitats live on

mussel banks on which they feed and mate. Such a con-

dition is reflected in the enormous morphological vari

ation shown by the shells of the different species, par-

ticularly in T. geversianus. While most of the species are

well known, some others were only collected in a single

location (e.g., T. amettei) and therefore their range “of

variation remains to be known.

Radular and anatomical features suggest that the en-

tire Tro} hon group from Patagonia is very homoge-

neous. An the species included in this genus have several

common radular features. The most femaieble are: the

intermediate denticle attached to the upper third of the

internal edge of the lateral cusp of the rachidian teeth;

a single marginal denticle in the external edge of the

base of the eaidian teeth; the attachment area of the

marginal teeth are always (no exceptions known) narrow,

thin, with the free part of same thickness, and the cen-

tral cusp of the rachidian is always thin and larger than

the laterals (see also Pastorino, 2002).

The Antarctic species so far assigned to Trophon and

the boreal ones recently included in the genus Boreo-

trophon (see Egorov, 1993) have the immer denticle be-

ie central and lateral cusp of the rachidian teeth al-

ways free, attached to the base of the teeth. In addition,

most of the radulae of these northeastern Atlantic spe-

cies—according to Bouchet and Warén’s revision

(1985)—have a lite: id attachment of the marginal teeth.

Conchological features are so variable that I consid-

ered them as secondary. However, the protoconchs are

actually very different and allow for the division in at

least two clearly defined oe There is no ornamen-

tation on the protoconchs of Patagonian representé itives

of Trophon, whereas most of the Loreal species of Bor-

eotrophon have a delicate pattern of irregular threads.

The anes species have also no ornamentation in the

protoconch with only one exception: T. scotianus Powell,

1951 which has apparently the same pattern observed in

North Atlantic Boreotroj yhon species.

There are several ae features that character-

ize the group of species living along the South American

coast. The accessory salivary glands, when known, are

usually tubular; the esophagus produces a typical esoph-

ageal loop after the cake of Leiblein and posteriorly

runs appressed to the left side of the gland of Leiblein;

Page SO

the esophageal glands in the mid-esophagus are incon-

spicuous, not exomally visible; and finally, the penises

are always dorso-ventrally flattened, with a large papilla

and a simple vas deferens either closed by the ov erlap-

ping sides of the penis or open.

The features mentioned above and the geological sto-

ry of the two areas, Antartica and Patagonia, allow for

the clear differentiation of these two groups.

ACKNOWLEDGMENTS

I thank the following people for making available spec-

imens and type material for study or advice when re-

quested: P. Greenhall and T. Nickens (USNM); K. Way

(BMNH), D. Reid (BMNH); A. Warén (NHRM); P.

Bouchet and V. Heros (MNHN); J. H. McLean and L.

Groves (LACM); P. Mikkelsen (AMNH); G. Rosenberg

and P. Callomon (ANSP); B. Sirenko (ZIL); O. Galvez

Herrera (MNHNS); C. Osorio (Santiago, Chile); F. Scar-

abino (DINARA) and C. Ituarte (MLP). P. Lozouet

(MNHN) took the photographs of specimens from Paris.

S. Horta (DINARA) and H. Racz-Lorenz (Montevideo)

kindly provided some of the specimens of Trophon clen-

chi that allowed part of the present work. Emily Vokes

(Tulane University) and Bruce Marshall (Museum of

New Zealand Te ‘Papa Tongarewa, Wellington) helped

with excellent reviews that Gonsiderabls improved on the

original manuscript. Finally special thanks to M. G. Har

asewych (USNM), who guided this work from the be-

ginning, and to M. Griffin for good advice.

Part of this study was conduc ted during a Postdoctoral

Fellowship granted by the Consejo Nacional de Inves-

tigaciones Centos y Técnicas (CONICET), Argenti-

na, to support my tenure at the National Museum of

Natural History, Smithsonian Institution, Washington,

DC. The study was also supported in part by a Research

Award from the NSF-USAP United States Antarctic

Program [Contract No. OPP-9509761], a Grant Award

from Conchologists of America and the Walter E. Sage

Memorial Aw ard, in addition to the projects PICTs 02-

01795 and 03-14419 from the National Agency for Sci-

entific and Technical Promotion, Argentina.

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THE NAUTILUS 119(2):83-89, 2005

Page 83

Latitudinal trends in shell characters of the neogastropod

Olivancillaria urceus (Gastropoda: Olividae) in the temperate

southwestern Atlantic Ocean

Alvar Carranza!

Walter Norbis

Seccién Oceanologia

Facultad de Ciencias

Igua 4225

11400 Montevideo

URUGUAY

ABSTRACT

Variation in shell characters of the neogastropod Olivancillaria

urceus (Réding, 1798) was analyzed at the central part of its

geographical range. Nine shell dimensions were measured

from specimens from seven localities between 32°10’ S and

40°33' S along the Atlantic coast of South America. er ae

effects of collection site were detected on all measured vari-

ables. The first three components in principal component anal-

ysis together explained nearly 0% of the observed variance in

morphome tric measures. Variables also differ in their allome-

tric trajectories among localities: all measured shell dimensions

displayed both positive and negative allometric coefficients

along the latitudinal axis. Allometric effects were then removed

to allow for determination of size-free variation in shape. High-

spired, elongated forms occurred more frequently in higher

latitudes. Spire width, maximum width, maximum width/height

and shell thickness decreased as latitude increased. Spire

length measured on the apertural side showed an opposite

trend. No relationship with latitude was found for fasciolar-

band length and spire length (measured along the abapertural

shell side). The existence of extreme forms within the analyzed

sample showed high phenotypic and ecological plasticity in the

populations assigned to O. urceus.

INTRODUCTION

Gastropod shell shape reflects a trade-off among func-

tional requirements, energetics of construction and

maintenance, rules governing growth, and the imprint of

evolutionary ancestry. The result of the interactions be-

tween these factors on the realized shell is also affected

by environmental features and tied to architectural con-

straints (Vermeij, 1993). Phenotypic plasticity in shell

characters appears to exert a strong influence on small

and large-scale morphological variation in marine snails

Trussel and Etter, 2001).

Different patterns in shell variation are found as dif-

' [email protected]

ferent spatial scales and taxonomical or ecological hier-

archies are considered: well-disseminated eco-geograph-

ical rules, suitable for many taxa, predict that animals at

high latitudes should have larger body sizes and life

spans than at low latitudes (Mayr, 1956: Atkinson and

Sibly, 1997). In most mollusks growth is faster at high

temperatures and in the presence of food than in eel.

nutrient-poor conditions (Vi ermeij, 1993).

Latitudinal variation in shell characters was detected

in northem hemisphere gastropods assemblages. This

variability was related to efficiency of calcium carbonate

utilization. It was shown that more heavily calcified and

ornamented shells occur with decrease in latitude

(Grauss, 1974).

When considering variation in shell features at the in-

traspecific level, it could be interpreted as a response to

environmental conditions, which may include biotic and

abiotic factors. Biotic interactions have been proposed

as driving forces for morphological changes. Certain

shell characters interpreted as anti-predatory features

such as low spire and elongate aperture were found to

be significantly different between comparable gastropod

assemblages from both sides of the tropical Atlantic and

Indo-West Pacific (Ver meij, 1978).

Within-location variation in shell shape and shore-lev-

el size gradients have been found for several species of

intertidal mollusks (Vermeij, 1972). This intra-popula-

tional variability is often coupled with shifts in shell

growth and has been related experimentally with food

supply and density-dependant processes: specimens of

Littorina littorea (Linnaeus, 1758) with plentiful supph

of seaweed grow faster and develop low-spired shells

than other group maintained under crowded conditions;

in the latter growth was slower and higher spired shells

were developed (Kemp and Bertness, 1984). De Wolf et

al. (1998) documented large-scale patterns of shell var-

iation in Littorina striata, a planktonic-developing peri

winkle from Macaronesia, confirming expectations based

Page 84

THE NAUTILUS, Vol. 119, No. 2

BRAZIL

Cassino

La Coronilla

La Paloma

Punta del Este

ARGENTINA

(Ax Mar del Plata

A Puerto Militar

ATLANTIC OCEAN

San Antonio

Figure 1. Map showing parts of the Atlantic coast of Brazil,

Uruguay, and Argentina that include the collection sites of Oli-

vancillaria urceus

on morphological patterns observed among other pros-

obranchs for increasing shell size and weight in southern

sites,

There is also ontogenetic related variation due to al-

lometric increase of spire length relative to other shell

dimensions (Vermeij, 1993). In this context, remotion of

allometric effects is needed in order to detect changes

in shell shape.

However, most of the studies dealing with morpho-

logical variability in gastropods are restricted to intertidal

rocky-shore snails, in particular those belonging to Lit-

torinidae. We know of no attempt made to analyze or

even describe patterns of intraspecific variation in neo-

gastropod shell features along a latitudinal axis in the

Southern Hemisphere.

The genus Olivancillaria d’Orbigny, 1839, is widely

distributed along the Atlantic coast of South America. It

comprises seven species occurring from tropical areas

(Espirito Santo, Brazil) to temperate regions (Golfo San

Matias, Argentina) (Burch and Burch, 1964; Klappen-

bach, 1964; 1965; 1966; Rios, 1994). Olivancillaria ur-

ceus (R6ding, 1798) is distributed along the entire lati-

tudinal range of the genus in subtidal soft bottoms, rang-

ing from the surf zone to at least 35 m depth (Juanicé

and Rodriguez-Moyano, 1976; Milstein et al., 1976; Es-

cofet et al., 1979; Scarabino, 1984) and exhibiting con-

spicuous differences in shell shape as noticed by Bara-

tinni and Ureta (1961). For these reasons, this species

is suitable for the examination of patterns of morpho-

logical variation in the region

In this paper we document large-scale variability in

hell features of Olivancillaria urceus in the temperate

Figure 2.

Dorsal, ventral, and apical view of Olivancillaria

urceus showing measurements taken for statistical analysis.

Seale bar = 1 cm. Abbreviations: LWL: last whorl length from

suture to anterior end; AL: aperture length, distance along out-

er lip from suture to anterior end: FB: distance between an-

terior and posterior ends of fasciolar band along internal side

of the aperture; MW: maximum linear distance from outer lip

to opposite side; MWH: maximum linear distance perpendic-

ular to MW to anterior end; ST: shell thickness; SW: diameter

of spire base from tip of callus above aperture to opposite point

on suture; SL2: lateral spire length from tip of callus above

aperture to protoconch tip on outer lip side in ventral view,

parallel to growth axis; and SLI; same length measured on the

opposite side

sector of it distribution range and analyze this species

variability along the latitudinal axis. We also discuss pos-

sible ecological implications of the observed pattern.

MATERIALS AND METHODS

\ total of 193 specimens collected in seven sites be-

tween 32° and 40° S along the Atlantic coast of South

America were analyzed. Collection sites were Cassino

Beach, Brazil (32°10' S, 52°20’ W, Site 1), La Coronilla

(33° 57’ §, 53°30’, Site 2), La Paloma (34°38" S, 54°08’

W, Site 3) and Punta del Este (34°36' S, W 58° 1S’ W,

Site 4), Uruguay, and Mar del Plata (38°02' S, 57° 32’

W, Sites 5), Puerto Militar (3S°44' S, 62°10’ W, Site 6)

and San Antonio Oeste (40°33' S, 64°50! W, Site 7), Ar-

gentina (Figure 1). Examined material is deposited at

Museo Argentino de Ciencias Naturales “Bernardino Ri-

vadavia” (Buenos Aires, Argentina) and Museo Nacional

de Historia Natural (Montevideo, Uruguay),

Nine shell dimensions were measured with vernier

A. Carranza and W. Norbis, 2005

Page 85

Table 1. Sample size, % of variation accounted by PCI and allometric coefficients for morphometric variables in each collection

site. Abbreviations: LWL: last whorl length from suture to anterior end; AL: aperture length, distance along outer lip from suture

to anterior end; FB: distance between anterior and posterior ends of fasciolar band along internal side of the aperture; MW:

maximum linear distance from outer lip to opposite side; MWH: maximum linear distance perpendicular to MW to anterior end;

ST: shell thickness; SW: diameter of spire base from tip of callus above aperture to opposite point on suture; SL2: lateral spire

length from tip of callus above aperture to protoconch tip on outer lip side in ventral view, parallel to growth axis; and SL1: same

length measured on the opposite side.

Locality

Variable l 2 3 4 bt 6 7

Sample size 27 17 39 18 43 2] 28

% of variation accounted

for by PCL 65.33 67.54 $4.36 71.81 94.23 87.82 75.44

Allometric coefficients

LWL 1.258 0.492 1.098 1.144 1.063 0.985 0.890

AL 1.451 0.37 1.166 0.965 1.072 1.028 L125

FB 1,544 0.617 1.207 1.086 1.142 1.05 0.948

MW L101 0.382 1.12 1.089 1.102 1.134 1.123

MWH 1.386 0.459 1.163 1.041 1.134 1.026 1.062

ST 1.214 0.485 1.143 1,159 1.118 1.043 1.151

SW 1.294 0.760 1.338 1.101 1.136 1.165 1.285

SLI 0.025 2.573 0.426 0.619 0.623 0.750 0.765

SL2 —0.274 2.858 0.340 0.796 0.610 0.819 0.650

calliper, within an accuracy of 0.05 mm. Abbreviations

for the variables are as follows: LWL: last whorl length

from suture to anterior end; AL: aperture length, dis-

tance along outer lip from suture to anterior end; FB:

distance between anterior and posterior ends of fasciolar

band along internal side of the aperture; MW: maximum

linear distance from outer lip to opposite side; MWH:

maximum linear distance perpendicular to MW to an-

terior end; ST: shell thickness; SW: diameter of spire

base from tip of callus above aperture to opposite point

on suture; SL2: lateral spire length from tip of callus

above aperture to protoconch tip on outer lip side in

ventral view, parallel to growth axis; and SLI: same

length measured on the opposite side (Figure 2). Col-

lection sites were grouped a priori for discretion of

differences among samples from different geographic

regions. For this we used the non-parametric Kruskall-

Wallis ANOVA median test. Allometric coefficients for

the seven collection sites were calculated for all vari-

ables. The method used was based on Jolicoeur (1963)

with additions from Kowalewski et al. (1997). Data were

log-transformed and subjected to principal component

analy sis (PCA). The first principal component (PC1) was

regarded as a size axis. Correlation between sampling

size and explained variance was examined by means of

regression analysis. The allometric coefficient for each

original variable was estimated by dividing the PC] load-

ing for that variable by the mean PC1 loading over all

variables. Estimation of 95% confidence intervals for the

cane coefficients was made by bootstrapping spec-

imens (2000 bootstrap replicates were made, not

jae

Size effects were then removed, following Lleonart et

al. (2000), by scaling all individual to the same size

(mean LWL) ) adjusting their shape to the one they would

have in the new size according to allometric change.

PCA was run on transformed dass and loads of the eight

remaining variables (i.e., all but total length) were cake

culated. This removal method is one of the most pow-

erful tools available, since it is in turn a theoretical gen-

eralization of the technique used by Thorpe (1975,

1976), which was recorded as one of the most efficient

methods in the empirical evaluation done by Reist

(1985).

A cluster analysis (single linkage, Euclidean distances)

was then performed to discriminate among possible

morphological groups using site mean value for each var-

iable. The cophenetic eourelanion coefficient (CCC) was

calculated to measure the internal distortion of the clus-

ter analysis following Sokal and Rolf (1962). Finally, pos-

sible relationships heteen latitude and shell characters

were examined by regression analysis.

RESULTS

All variables showed significant differences between sites

(non-parametric ANOVA: p < 0.01). PCA performed

with log-transformed data showed that variance ex-

plained by PC1 (Factor 1, regarded as size axis) differed

among sampling sites and varied between 65.33% (site

1) and 94.23% (site 5). Explained variance was not cor-

related with sample size (regression analysis, F = :0.010,

p-level = 0.922). Allometric coefficients calculated for

the nine variables also varied among collection sites. All

variables displayed both positive (b > 1) and negative

(b < 1) allometric trajectories (Table 1).

The first three factors in size-free PCA explained to-

gether 79.38% of the total variance. Variables MW,

MHW, ST, and SW grouped together with positive loads

on Factor 1. SLI and SL2 formed another highly cor-

Page 86

THE NAUTILUS, Vol. 119, No. 2

Table 2. Results of size-free PCA Analysis. % of variance ac-

counted by each factor and loading of each variable are shown

Abbreviations: LWL: last whorl length from suture to anterior

end; AL: aperture length, distance along outer lip from suture

to anterior end; FB: distance between anterior and posterior

ends of fasciolar band along internal side of the aperture; MW:

maximum linear distance from outer lip to opposite side;

MWH: maximum linear distance perpendicular to MW to an-

terior end; ST: shell thickness; SW: diameter of spire base from

tip of callus above aperture to opposite point on suture; SL2:

lateral spire length from tip of callus above aperture to pro-

toconch tip on outer lip side in ventral view, parallel to growth

axis; and SLI: sane length measured on the opposite side.

Factor loadings

Variable PC] PC2 PC3

% of variation 41.54 22.9 14.95

AL 0.422 —(0.474 0.550

FB 0.173 —(.487 0.716

M\W 0.874 0.2125 —0.063

MHW 0.888 0.083 0.034

ST 0.S58 0.226 —O.11]

SW O.S1] 0.246 —(0.044

SLI —(0.094 O.S15 0.446

SL2 —0.396 0.735 0.399

related group loading positively on Factor 2, and AL and

FB did the same on Factor 3 (Table 2)

Two different groups were discriminated when the

collection-site mean value for each variable were plotted

onto a hierarchical cluster, one of these groups com-

posed by individuals belonging to sites 6 and 7 (Figure

The calculated value for CCC was 0.93.

Latitudinal-related patterns of variation were found in

variables MW, MHW, ST, AL, and SW. This group of

variables showed a decreasing trend with increasing lat-

itude. On the other hand, SL2 tend to increase with

latitude, whereas SLI and F . ran wved no relationships.

The coefficient of determination (r?) explained less than

30% for all variables (Figure 4).

DISCUSSION

The present work documents shell shape variation in O.

urceus along a latitudinal gradient. The results show that

shell features vary between sites. The analyses presented

above show that northern forms are stunt, low- -spired,

and conical, whereas southern specimens (mostly those

from sites 6 and 7) are high-spired, narrower and elon-

gated.

a location variation may be due to local differ-

ences in habitat conditions, for example, contrasting

re morphodynamics, which are in turn associated

with different sediment features. However, the poor

geographical definition of collection sites did not allow

us to assign in all cases a particular individual to a par-

ticular habitat. Notwithstanding, intraspecific phenotypic

plasticity in a small spatial scale is known to occur in

mollusks. For example, observations made on the mur-

icid Nucella la villus (Linnaeus, 1758) showed that spec-

imens from very exposed shores are short-spired when

compared with those from sheltered localities (Cooke,

1895; 1915).

When considering variation at a geographic scale, two

possible scenarios are suggested to explain the among-

collection sites differences. These are either: (1) a grad-

ual response to shifts in environmental conditions along

the latitudinal gradient that generate a shape gradient

(in this scenario, macro-scale variation in physical pa-

rameters such as sea water temperature and calcium car-

bonate availability could be correlated with the observed

variation); or (2) different morphs associated with par-

Figure 3. Extreme forms of Olivancillaria urceus in the study area

Dorsal and apertural views of specimens from site 7. Scale bat

A. dorsal and apertural view of specimens from site 3. B.

Lom

A. Carranza and W. Norbis, 2005

Page

40,

MW = 45.70 - 0.60 * Lat“

= 0.286 ;

Maximum Width (mm)

DP p> Pp Pp

a SW = 38,74 - 0,64 * Lat

r= 0.281

Spire Width (mm)

—

fe)

> DPE P D> p>

DA AADIRE b DD

PERE | >

DAN >> pb>

AL = 36.68 -0.14" Lat

r= 0.050

Aperture Length (mm)

28 A

31 32 33 34 35 36 37 38 39 40 41 42

Shell Thickness (mm) Max. Width Height (mm)

Spire Length 2 (mm)

MWH = 42.97- 0.47 * Lat “

r’=0.284 ,

> DEE! >

DDE DE I D>

ST =39.92-0.55*Lat 4

’=0.244 “4

28 4 A

DOSRBR >

DE D>

B>pe> PP

SL2 = -1.39 + 0.22 * Lat A

r= 0.153

(oe)

PEP DP > >

DPD > b>

Se DE>

MP DPB >

D> PRPPP> DP DP

4, 4 z

1 32 33 34 35 36 37 38 39 40 41 42

Latitude

Figure 4.

and linear model fitted are shown.

ticular habitats, such as exposed sandy beaches or shel-

tered bays: broader specimens with short spire in high-

energy habitats and high-spired, elongated forms in pro-

tected habitats. In this case, likelihood of occurrence of

one particular morph depends on environmental char-

acteristics of geographic locations and may be consid-

ered local populational phenomena.

A related species, Olivancillaria vesica (Gmelin, 1791)

also shows two geographical forms, recognized as sub-

species. which are easily separable along its distribution

range (Klappenbach, 1966). This species shows an op-

posite latitudinal pattern: northern forms are narrower

and longer compared with the broader, “auriculated”

ear-shaped) southern forms. In accordance with (2), the

latter form extends along the southerm coast of Brazil

Olivancillaria urceus, relationship between standardized shell characters and latitude. Coefficients of determination (1°)

(Santa Catarina), Uruguay, and Argentina as far as

Puerto Quequén, Province of Buenos Aires, the same

geographical area in which O. urceus displays broader

and low spired shells. These broader shell forms could

be interpreted as associated with a large foot that en-

hances its ability of “anchoring” to the substratum in

higher energy environments, thus improving individual

fitness. However, more study is necessary on the taxo-

nomic status on O. vesica forms before any analysis of

its morphological variation.

It can be argued that differences in spire length and

width reflect shifts in growth patterns associated with

the adaptive process. It should be noticed that there is

a coincidence between occurrences of high-spired forms

(associated with slower growth mode) in higher lati-

Page 88

THE NAUTILUS, Vol. 119, No. 2

LINKAGE DISTANCE

Pto. Militar

San Antonio

Punta del Este

La Paloma

La Coronilla Cassino

Mar del Plata

COLLECTION SITES

Figure 5.

each morphometric variable.

tudes. This could be supporting the hypothesis that both

latitudinal and env ironmental effects may be coupled to-

gether in a combined effect that masks the isolated ef-

fects of each factor.

As demonstrated by PCA analysis, nearly half of the

variance is explained by PC] (42% of total variance),

which in turn is mainly affected by MW, MWH, ST. and

SW. This fact implies that changes in shell shape de-

tected within the analyzed sample are due mostly to dif-

ferences in its variables. Variation observed in SL2 and

SLI are of much less importance as form determinants

when considering total variance components. However,

height and shape of the spire and protoconch characters

have been regarded as important specific (L6pez et al.,

1988) and supraespecific (Tursch, 1988) features, for

which these characters deserves further analysis. Nev-

ertheless, it seems that macrogeographic variation in

shell morphology strongly depend on local conditions,

which make adequate sampling and data treatment very

difficult.

ACKNOWLEDGMENTS

The authors wish to thank Mr. Fabrizio Scarabino (Di-

reccion Nacional de Recursos Acuaticos, Montevideo),

Dr. Sergio Martinez and Dr. Alejandro Brazeiro (Facul-

tad de Ciencias, Montevideo), who provided useful bib-

Olivancillaria urceus, cluster analysis. Collection sites were grouped on the basis of similarities in the mean value for

liography and suggestions that helped us to improve this

manuscript. J. L ie sonart made his own designed software

available for statistical analysis on remotion of allometric

effects. Authors would also like to e xpress their gratitude

to the two anonymous referees for their great help in

reviewing and correcting the original version of the man-

uscript. A.C. thanks Msc. Estela Delgado for encourage-

ment and support and J. de los Santos for assistance with

the figures.

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pp:

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y 43°S.: consideraciones generales y esquema biocenol6-

gico. Memorias del Seminario sobre Ecologia Benténicas

y Sedimentacion de la Plataforma Continental del Atlan-

tico Sur, 1: 245-258. UNESCO, Montevideo.

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Juanicé, M. and M. Rodriguez-Moyano, 1976. Composicion

faunistica de la comunidad de Mytilus edulis platensis

dOrbigny, 1846, ubicada a unas 55 millas al SE de La

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Kowalewski, M., E. Dyreson, J. Marcot, J. Vargas, kK. Flessa

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THE NAUTILUS 119(2):90-91, 2005 Page 90

Errata

Due to an editorial lapse, the Introduction section was omitted from K6hler and Glaubrecht’s (2005) article in the

most recent issue of The Nautilus. The missing Introduction is printed below (literature references are in the main

article), with apologies to the authors and readers.

Fallen into oblivion—the systematic affinities of the enigmatic Sulcospira Troschel, 1858 (Cerithioidea:

Pachychilidae), a genus of viviparous freshwater gastropods f rom Java

Frank Koéhler

Matthias Glaubrecht

INTRODUCTION

The first volume of the well-known work “Das Gebiss der Schnecken zur Begriindung einer natiirlichen Klassifikation”

[Establishing a natural classification of snails from their dentition] by Franz Hermann Troschel (1810-1882) was

published in parts between 1856 and 1863. The work is an important historic landmark in the enduring challenge of

zoologists to create a natural classification of the living Sapel rds. In his work, Troschel (p. 117) described the genus

Sulcospira within the tribus “Pachychili) mainly based on features of the operculum and the radula (Figure Ly Ac-

cording to Robertson (1957), that description was wc bished in L858. Within the non-marine C ‘ereiioided, Sulcospira

represents one of the least known genera of Southeast Asian Pachychilidae, a group of viviparous freshwater gastropods

we have been focusing on in the last few years.

Recent research aiming to propose a phylogene tic systematics hypothesis and to establish a natural classification of

this limnic gastropod banaly will also allow a better understanding of their evolution, morphology, and ecology (Glau-

brecht, 1996: 1999: Kohler and Glaubrecht, 2001; 2002; 2003: Glaubrecht and Rintelen, 2003). Only recently, the

Pachychilidae Troschel, 1857, have been shown to represent a monophyletic group clearly distinct from the Thiaridae

Troschel, 1857, and from the other limnic Cerithioidea. This new conce pt has been suggested by analyses of mor-

phological data (e.g. Glaubrecht, 1996; 1999) and is corroborated by molecular data suggesting that Thiaridae (e.g.

Melanoides, Thiara, Tarebia) and Pachychilidae (e.g. Paracrostoma Cossmann, 1900, Pac shyc hils Lea, 1850) are not

very closely related to each other (see phylogenetic reconstruction in Lydeard et al., 2002: figs. 1, 2). This classification

conflicts with the traditional view of most earlier authors who treated pachychilid taxa as tae = Thiaridae

(among others, Thiele, 1929; Rensch, 1934; Benthem-Jutting, 1956; Brandt, 1968; 1974) or Pleuroceridae (e.g. Ponder

and Warén, 1988; Vaught, 1989).

However, our current knowledge of the phylogeny and systematics of freshwater Cerithioidea in general and the

Pachychilidae in particular is still limited, since many taxa remain poorly known. Only recently, systematic studies

using morphological as well as molecular genetic data have shed some light on the rele tionships of several other genera

withen the Pachychilidae. For instance, Kohler and Glaubrecht (2001) presented comparative morphological de ita on

taxa tre ditionally assigned to Brotia H. Adams, 1866, by various authors, revealing that this genus as previously per-

ceived actually comprises four lineages, each characte rized most conspicuously by distinet re productiv e morphologies.

Subseque nt studies including molecular phylogenetics data put special e mphasis on two of these lineages, Jagora

Kohler and Glaubrecht, 2003, endemic to the Philippines (KGhler and Glaubrecht, 2003) and Tylom dania F. and P.

Sarasin, 1595, endemic to Sulawesi (Rintelen and Glaubrecht, 1999; 2003), suggesting an independent generic status

for each. In addition, the properties of another Australasian pachychilid genus, Pse doy yotamis Martens, 1894, have

been extensively described by Glaubrecht and Rintelen (2003). Hence, a “nideable alin of crucial biological infor-

mation on pachychilids has been amassed, he Iping to facilitate a better understanding of pachychilid systematics,

phylogeny, and evolution.

Nevertheless, there remain a number of systematic and taxonomic problems and difficulties related to this group

of freshwater snails. One of them will be dealt with in this study: The t: wonomy and systematic position of Sulcospira.

Among the various generic names that have been introduced for Southe ast Asian pe achychilid taxa, Sulcospira is

clearly the oldest one. Its description predates that of other names, such as Brotia, Antime lania Fischer and Crosse,

1892, or Pse udopotamis; a complete annotated list of the introduced supraspecific names within the Southeast Asian

Pachychilidae is given in Kohler and Glaubrecht (2002). Consequently, every other supraspecific pachychilid taxon is

valid only with the reserve that it is not a junior synonym of Sulcospira. In spite of this significant taxonomic role of

Sulcospira, this taxon has been widely ignored, e spec ‘ially by modern systematists. Thus, another aim of this article is

to compile all available information on Sulcospira and to provide new data from our own examinations of the limited

Errata, 2005 Page 9]

material from museum collections. In addition, implications for pachychilid taxonomy and systematics are discussed

in relation to Sulcospira.

LITERATURE CITED

Kohler, F. and M. Glaubrecht. 2005. Fallen into oblivion—the systematic affinities of the enigmatic Sulcospira Troschel, 1858

(Cerithioidea: Pachychilidae), a genus of viviparous freshwater gastropod from Java. The Nautilus 119: 15-26.

In the recently published article by Ardila and Valdés (2004), please note the following corrections:

On page 134, caption of Figure 3, replace “Armina muelleri Thompson, Cattaneo and Wong, 1990”, for “Armina

muelleri (Ihering, 1886)”;

On the same page, the adjacent subsections “Type Material” and “Type Locality” should be replaced by:

Material Examined: MHNMC INV MOL3901, 32 mm length, alive, from off Salamanca Island, Colombia (11°5'46”"

N, 74°40'35” W), 20 m depth, hard bottom with pennatulaceans (Renilla reniformis and Renilla muelleri).

LITERATURE CITED

Ardila, N. E. and A. Valdés. 2004. The genus Armina (Gastropoda: Nudibranchia: Arminidae) in the southern Caribbean, with the

description of a new species. The Nautilus 11S: 131-138,

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THE NAUTILUS

Volume 119, Number 3

October 6, 2005

ISSN 0028-1344

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CONTENTS

one

MNAUTILUS

Volume 119, Number 3

October 6, 2005

ISSN 0028-1344

Cristian Ituarte

Shannon M. Carpenter

Paolo Mariottini

The Sphaeriidae (Bivalvia) from northwestern Argentina including three

EW Species Of PIStdiH ge socos oun Oo ee te MEA a Ha4 we ED EER Ee ER 93

Mysella pedroana, a commensal bivalve (Lasaeidae) on two decapod

Crustacean NOSstS. cs 448% an oe bathe Pe taaneahwag faced edemedan 105

Coralliophila trigoi (Gastropoda: Muricidae), a new species from the

Carlo Smriglio northeastern Atlantic Ocean . i. s406cacee cd Gk ded aaa ee ee vba eee 109

Emilio Rolan

Book REVIEW. 6 6.465.556 4645 babar coe bem oabbte habeas dhoudn cn tavesedananeacucapebhabaa 116

OC) 18 2005

THE NAUTILUS 119(3):93-104, 2005

Page 93

The Sphaeriidae (Bivalvia) from northwestern Argentina including

three new species of Pisidium

Cristian Ituarte

Division Zoologia Invertebrados

Museo de La Plata

1900 La Plata

Buenos Aires

ARGENTINA

[email protected]

|, n,n

ABSTRACT

Knowledge on the Sphaeriidae fauna in southern South

America is significantly improved with the description of three

new species of the genus Pisidium from Salta and Jujuy

provinces (northwestern Argentina). This paper provides the

first record of sphaeriids in restrictive high-altitude South

American environments, particul uly from very small water

courses found in “vegas”,

“cushion vegetation” or “cushion peat bogs” developing

between 2000-4000 m altitude in the Argentine pre-Andean

ranges. Furthermore, the geographic distribution range of

Pisidium chiquitanum Ituarte, 2001, only reported to date from

the type locality in sub-Andean regions of central Bolivia, is

considerably e nlarged.

INTRODUCTION

The Sphaeriidae fauna from Argentina is poorly known;

the first reports were those by Strobel (1874) on

Musculium argentinum (dOrbigny, 1835) and Pilsbry

(1911) describing several new gee of Pisidium C.

Pfeiffer, 1821, and Musculiwm Link, 1807, from

Patagonia. Later on, Doello-Jurado (1921) described

the first species of Eupera Bourguignat, 1854, from

Argentina, and Ituarte (1989, 1994) and Ituarte and

Dreher-Mansur (1993) described three new species of

Eupera from Iguazi, Uruguay and Parana River basins

in northeastern Argentina. Regarding the species di-

versity of Pisidium C. Pfeiffer, 1$21, twelve species are

known from Patagonia and Northeastern provinces

Ituarte, 1996, 1999, 2000).

Only two species of Sphaeriidae have been reported

from northwestern Argentina: Sphaerium lauricochae

Philippi, 1869), from Jujuy Province (Ituarte, 1995)

and Musculium argentinum from Mendoza Province

(Strobel, 1874). The species diversity of Pisidiwm

has essentially not been documented. In _ the

present paper, three new species of Pisidium

from lowland and high-altitude habitats in the pre-

Andean mountain ridges are described. Based on new

exce ptionally localized areas of

findings the knowledge on the geographic distribu-

tion of Pisidium chiquitanum Ituarte, 2001, is

updated.

MATERIALS AND METHODS

Materials for the present study were obtained during

three field trips to Northwestern Argentina (Tucuman

and Salta provinces in March 1999, Salta and Jujuy

provinces in December 2001 and March 2004, and

Catamarca Province in March 2004). Figure | shows the

location of collecting sites; more detailed information on

the source of specimens is given in the Systematics

section. The collected specimens were fixed immediately

after collecting in 70° alcohol after being relaxed through

a short rinse (around 20 seconds) in warm water (about

50°C). Specimens for scanning electron microscopy

(SEM) were cleaned by repeated rinsing in distilled

water followed by a short treatment (about 5 seconds) in

10% sodium hypochlorite solution. Soft anatomy was

studied after decalcification of valves through a 12-hour

rinsing in a 5% formaldehyde and 2% acetic acid

solution. Linear measurements (shell length [SL], shell

height [SH], shell width [SW] and presiphonal suture

length [PSS]), shape indices and coun ratios

(height index [HI= SH/SL]. convexity index

SH], ratio of hinge length [HiL] to shell length [HiL/

SL]), were calculated according to the criteria followed

by Ituarte (1996). For each calculation (n= 10, unless

otherwise stated), mean and standard deviation values

are given.

Type specimens are deposited at Museo de La Plata,

La Plata (MLP), Museo Argentino de Ciencias Naturales

“Bernardino Rivadavia”, Buenos Aires (MACN), Funda-

cin Miguel Lillo, Tucuman (FML) and Muséum

National d'Histoire Naturelle, Paris (MNHWN). Types of

Pisidium chiquitanum, housed at Museo de Historia

Natural “Noel Kempff Mercado”, Santa Cruz de La

Sierra, Bolivia (MHNB) and MLP were also used for

con iparatiy e pl 1 rpc SES.

Page 94

(

BOLIVIA

THE NAUTILUS, Vol. 119, No. 3

Balasto®/

\/@Hualfin®

Snel t \ 3 \

68° Ww

Figure 1.

SYSTEMATICS

Pisidium omaguaca new species

(Figures 2-15)

Diagnosis: Shell markedly oval, high and anteriorly

produced, beaks depressed, displaced backward, not

projecting from shell surface and only barely visible

above dorsal margin; ligament internal; anal and

branchial mantle openings present; two demibranchs

present, nephridia of closed type, with lateral lobe not

visible in lateral or dorsal views.

Description: Shell thin, translucent, small to medium

size (mean SL = 2.73 + 0.17, maximum observed size:

3.7 mm), rather high (mean HI = 85 + 1) (Figure 2),

not inflated (mean Ci = 58 + 3) (Figures 2, 3). Shell

| | \ \ \

Tt | Catamarca’

ca.

Location map. Stars in black indicate type localities of Pisidium omaguaca new species, Pisidium ocloya new species, and

Pisidium chicha new species.

outline markedly oval, anteriorly elongated. Anterior end

produced in a sharp curve, posterior end short, widely

rounded, sometimes slightly truncated and _ straight

(Figures 2, 4, 5, 8). Dorsal margin short, weakly

connected with anterior margin, which slopes markedly

towards anterior end; sometimes, a gentle angle marks

joining point of dorsal and posterior margins (Figure 8).

Beaks very low, depressed and wide, not raised from

shell surface, only slightly projected above dorsal

margin, displaced backward, located at about 59% of

SL (Figures 2, 4,5). Shell surface finely and somewhat

irregularly striated, glossy, amber.

Hinge plate strong, hinge line short (HiL/SL = 53 +

3), strongly curved. Hinge: Left valve (Figures 4, 6):

cardinal teeth well-developed, the inner one (Cs) thin,

short at base, bent upward, slightly oblique with respect

C. Ituarte, 2005 age 95

Figures 2-7. Pisidium omaguaca from Termas de Reyes, Jujuy. 2. Holotype (MLP 5496-1-1): outer view of right valve. 3-7.

Paratypes (MLP 5496-1-2). 3. Posterior view. 4. Left valve, inner view. 5. Right valve, inner view. 6. Left valve, detail of hinge

Right valve, detail of hinge. Scale bars = 500 um

to antero-posterior axis rounded at the tip, outer displaced forward posterior lateral tooth (PIL) minute

cardinal tooth (C4) a narrow, slightly wider at posterior straight and strong, cusp high, distally displaced. Right

end, uniformly curved blade, quite oblique, overlapping valve (Figures 5, 7): cardinal tooth (C3) not strong

C3 at posterior half: anterior lateral tooth (AIT) ver somewhat displaced forward, curved in the middle quite

strong. short, nearly straight, cusp high, pointed, narrow at anterior half, slightly enlarged in a posterior

Page 96 THE NAUTILUS, Vol. 119, No. 3

Figures 8-13. Pisidiwm omaguaca. 8. Paratype (MLP 5485). Right valve, outer view. 9. Paratype (MLP 5496-1-2). Dorsal view,

detail of the escutcheon, 10-11. Inner view of left and right valves of a specimen from Tiraxi, Jujuy (MLP 6535). 12, 13. Outer and

inner views of right and left valves of a specimen from a “peat bog” between Yavi (Salta) and Santa Victoria (Jujuy) (MLP6559). Scale

bars (8, 10-13) LOO00 Um; (9) = 200 Lm

elongated cup. Lateral teeth short and robust, inner never visible from outside (Figure 9). Escutcheon

anterior lateral (AI) curved cusp subcentral or slightly inconspicuous (Figure 9).

displaced anteriorly; outer anterior lateral tooth (AIII)

quite short, cusp distal; inner posterior lateral (PI) nearly Anatomy: Anal siphon and branchial inhalant mantle

straight, short cusp subcentral; outer posterior lateral opening present. Presiphonal suture about 9% of shell

PIII) minute, with distal cusp leneth (Figure 15). Eight or nine well-marked muscle

Ligament-pit enclosed, deep, inner margin straight o1 scars located away from pallial line correspond to inner

slightly concave (Figures 6, 7). Ligament relatively short radial mantle muscles. Muscle scars corresponding to

representing 20 + 1% of shell length, strong, internal, anal siphon retractors are coalescent with that of

C. Ituarte, 2005

Page 97

Via

Figures 14-15. Pisidium omaguaca new species. 14. Gross

anatomy. 15. Mantle muscles. (as: anal siphon; asr: anal

siphon retractor; ia: inhalant aperture; id: inner demibranch;

isr: inhalant siphon retractor; irm: inner radial mantle muscles;

n: nephridium: od: outer demibranch; pss: presiphonal suture).

posterior adductor muscle (Figures 4, 5). Bundles of

fibers of inner radial mantle njnaales strong, converging

anteriorly, except for two posterior bundles { (Figures 4,

5. £4).

Inner and outer demibranchs present. Outer demi-

branch much smaller, formed by 10-12 very short

descending filaments, reaching back to the 14"” filament

of inner demibranch ( (Figure 14). Up to three large

embryos (1.3 mm length) were found within each becod

pouch of a specimen 3.7 mm L. Nephridia of closed

type. dorsal lobe, usually subquadrate, completely

covering pericardial part of nephridium (Figure 14).

Type Locality: A small watercourse opening into Reyes

River at Termas de Reyes, 24°10'19" S, 65°29'27" W,

1754 m altitude, Jujuy Province, Argentina (Figure 1).

Type Material: Holotype (MLP 5496-1-1) and 42

paratypes from the type locality (16 paratypes MLP

5496-1]-2: 6 paratypes MLP 5485: 4 paratypes MACN-In

36361; 12 paratypes FML 14506; and 4 paratypes

MNHN.

Other Material Examined: Jujuy Province: numer-

ous specimens from the type local ity (MLP 5496-1-3)

Tumbaya, small pool with vegetations at the side of

national road No. 9 (23°47'28” §, 65°28'37” W), 2070 m

(MLP 6548); small stream flooding from springs in

highland areas covered with “cushion vegetation”,

between Yavi (Jujuy) and Santa Victoria (Salta)

(22°07'11" S 65°13'05” W), 4150 m (MLP 6559): small

pool at the side of Rio Grande River (22°58'14" S

65°27'01" W), 3950 m (MLP 6530); small springs at side

of Manzanito Rivulet, near Huertas (22°14'20" S

65°00'31" W), 2740 m (MLP6551): small stream at the

side of the road to Yala (24°07'20" S 65°24'16” W),

1430 m (MLP 6531). Salta Province: unnamed brook on

provincial road No. 57, near Cachi (25°05'24" S

66°07'33" W), 2340 m (MLP 6540).

Distribution: Highlands of Jujuy and Salta provinces,

Argentina, between 1400 and 4100 m altitude.

Etymology: The name refers to the Omaguacas,

ancient aboriginal inhabitants of the Quebrada de

Humahuaca, the spectacular 150 km long valley of the

Rio Grande River (Figure 1), which underwent a major

cultural change during the past 10,000 years.

Remarks: Pisidium omaguaca new species differs

from Pisidium meierbrooki Kuiper and Hinz, 1984, in

being higher and less obese, having lower and_ not

inflated be: iks, and a more broadly founded posterior

end. Pisidium omaguaca new species is similar to

Pisidium chiquitanum Ituarte, 2001, in soft anatomy,

but differs in having a more solid and higher shell, with

posterior end shorter and anterior half of dorsal margin

sloping markedly towards the anterior end.

Pisidium ocloya new species

(Figures 16-33)

Diagnosis: Shell rather trapezoidal and high, small size,

presence of only one (anal) mantle aperture and one

demibranch.

Description: Shell thin, translucent, of small to

medium size (maximum observed SL = 3.2 mm), high

(mean HI = 85 + 2), quite convex (mean Ci = 77 + 4),

shell outline rather trapezoidal. Anterior end somewhat

produced in a sharp curve, posterior end short,

truncate, somewhat oblique (Figures 16, 17). Beaks

full, wide at base, markedly raised from shell surface

and projected above dorsal margin, subcentral or

slightly displaced backward, located at about 58% of

SL (Figures 16, 19, 20). Shell surface fine ly and

irregularly striated (Figures 16, 18), glossy, whitish or

yellowish. ;

Hinge plate not strong, narrow in middle, hinge line

rather long (HiL/SL = 56 + 2), arcuate. Hinge on right

valve (Figures 20, 22): cardinal tooth (C3) delicate,

Page 98 THE NAUTILUS, Vol. 119, No. 3

Figures 16-22. Pisidium ocloya new species from Burrumayo River, Jujuy. 16. Holotype (MLP 5499-1): outer view of right valve

17-22. Paratypes (MLP 5499-2). 17. Posterior view. 18. Posterior view, detail of ligament. 19. Left valve, inner view. 20. Right

valve, inner view. 21. Left valve, detail of hinge. 22. Right valve, detail of hinge. Scale bars (16, 17, 19-22) = 500 um; (18)

200 Lm

rather weak, slightly curved, narrow on anterior half, posterio1 lateral (PI) gently curved, not long, cusp

nlarged into a well-marked, blunt posterior cup distally displaced; outer posterior lateral (PIIT) reduced

Lateral teeth robust, inner anterior lateral (AI) well- in size, with distal cusp. Hinge on left valve (Figures 19,

developed, cusp displac ed distally: outer anterior lateral 21)

cardinal teeth minute, inner one (C2) short and

tooth (AIII) shorter and weaker, cusp distal; inner high, horizontal with respect to antero-posterior axis,

SSS SSS SSS

C. Ituarte, 2005

Page 99

outer one (C4) short, oblique, slightly overlapping Co at

posterior end; anterior lateral tooth (AIT) very strong,

cusp high, distal; posterior lateral tooth (PII) shorter and

weaker, cusp high, distal.

Ligament-pit enclosed, deep, inner margin slightly

and evenly curved (Figures 21, 22), Ligament narrow,

relatively long, representing 24 + 1% of shell length,

visible from outside and somewhat protruded at anterior

half of ligament length. Escutcheon slightly marked by

a delicate lanceolate line (Figures 17, 1S).

anatomy: Only one demibranch (inner) present

(Figure 23). Only anal mantle opening present. Inner

radial mantle muscles weak, 6-7 bundles of few weak

fibers converging anteriorly, attached just above pallial

line (Figure 23): sometimes scars corresponding to inner

radial muscles coalescent with pallial line (Figures 19,

20). Anal siphon retractors attached immediately ven-

trally to posterior adductor muscle. Nephridium with

lateral loop visible dorsally (Figure 24).

Type Locality: Small flooded areas on the banks of

Burrumayo River (24°10'18” S, 65°22'43” W), 1201 m

altitude, in the neighborhood of Jujuy City, Jujuy

Province, Argentina; and unnamed brook opening into

Zapla River (24°16'03" S, 65°07'09”" W), 946 m altitude,

Zapla, Jujuy Province, Argentina.

Type Material: Holotype (MLP 5499-1) and 15

paratypes from the outskirts of Jujuy City (6 paratypes

MLP 5499-2; 10 paratypes FML 14505; 56 paratypes

from Zapla (36 paratypes MLP 6899-2; 10 paratypes

MACN-In 36362; 10 paratypes MNHN),.

Other Material Examined: Catamarca Province:

unnamed brook at national road No. 40 at La Ciénaga

de Abajo, between La Ciénaga and Belén (27°31'05" S,

66°59’08" W), 1520 m (MLP 7201); Jujuy Province:

numerous specimens from the type locality (MLP 7369):

unnamed brook on side of provincial road No. 4 at

Guerrero (24°11'13" S, 65°26'51"” W), 1650 m (MLP

7379); small spring on side of provincial road No. 4, near

Termas de Reyes (24°10'36"” S 65°28'18" W), 1730 m

(MLP 5497-1, MLP 7370-1); small water course on side

of national road No.9 at Tumbaya (23°51'26" S,

65 27'57" W), 2020 m; Los Cedros Rivulet, south to

“El Carmen” (24°28'0" S 65°17'08" W), 1190 m (MLP

6545-1); small spring near Tiraxi (23°59'57" S,

65°19'39" W), 1576 m (MLP 6553): on the road

between Termas de Reyes and Laguna Yala, flooded

areas at the bottom of hills (24°07'47" S, 65°28'58” W),

1920 m (MLP 7371). Salta Province: unnamed brook at

national road No. 40 at Molinos (25°18’'53” S,

66 -14'5S8” W), 2155 m (MLP 6526); small spring at road

No. 33. to Cachi at Los Laureles (25°06'27" S,

65° 36/10" W), 1360 m (MLP 6529): Tucuman Province:

flooded areas at side of the Nio River, near Rio del Nio

City 26 25.60’ S 64°55.60' W), SS6 m (MLP 7403): on

side of Medina River, on provincial road No. 305,

between El Tipal and Aserradero (MLP 7404-1); small

24 CS

Figures 23-24. Pisidium ocloya new species. 23. Gross

anatomy. 24, Detail of nephridium. (id: inner demibranch;

irm: inner radial mantle muscles; n: nephridium; p: pericar-

dium; pa: posterior adductor; r: rectum.

pond on side of Potrero de Las Tablas River, Raco,

880 m (MLP6992), La Angostura dam, on Los Sosa

River (26°55'21” S, 65°41'02” W), 2000 m (MLP 6897-

Distribution: Catamarca, Jujuy, Salta, and Tucuman

provinces, Argentina, between 880 and 2155 m altitude.

Etymology: The name of the new species refers to the

Ocloyas, ancient aboriginal inhabitants of the lands in

the surroundings of the type locality.

Remarks: Pisidium ocloya new species _ strikingly

differs from other northwestern Argentine Pisidium

species by its relatively small size and trapezoidal shell

outline. Compared with Pisidium vile Pilsbry, 1987,

a small species from the eastern drainage system of the

Rio de La Plata Basin, P. ocloya new species differs in

being larger and comparatively lower. Ituarte (1999)

described Pisidium huillichum from southern Chile,

another small-sized Pisidium species with one mantle

opening and one demibranch, which differs from P.

ocloya in having a non trapezoidal shell outline, lower

beaks, very strong lateral teeth, and strongly marked

commarginal ribs of the shell surface.

The height/length ratio, the convexity index and the

degree at which part of the ligament is protruded

showed a relatively wide variability in samples of

Pisidium ocloya from different localities (Figures 25-

33); larger specimens were in general more convex with

much inflated and pronounced beaks, and more sharply

defined trapezoidal outline.

THE NAUTILUS, Vol. 119, No. 3

Figures 25-33. Pisidiwm ocloya new species. 25-27. Specimens from Termas de Reyes, Jujuy (MLP 5497-1). 28-30. Specimens

from Zapla, Jujuy (MLP 6899-2). 31-33. Specimens from Cachi, Salta (MLP 6529). Scale bars = 500 um

Pisidium chicha new species

(Figures 34-43)

Diagnosis: Rather elliptic shell outline sub-centrally

located beaks, somewhat inflated and markedly raised

from shell surface, are diagnostic features. The presence

of only one mantle aperture and one demibranch is also

distinctive

Description: Shell small, maximum observed. shell

length 2.7, moderately high (mean HI 79 + 1), not

convex (mean Ci = 68 + 5): shell outline strikingly oval.

Dorsal and ventral margins broad, dorsal margin, slightly

ircuate, ventral margin uniformly curved; anterior end

evenly curve and only. slightly projected forward,

posterior end slightly truncated (Figures 34-37). Beaks

wide, widely rounded at tip, somewhat inflated, raised

above dorsal margin but low, sub-central, slightly

displaced backward, located at about 57-58% of SL.

Shell surface glossy, amber, sculptured with well marked

fine and rather regularly spaced striae (Figure 34).

Hinge plate narrow, hinge line somewhat short, HiL/

SL = 53 + 3% of SL (n = 8), widely curved. Hinge on

left valve (Figures 36, 38): cardinal teeth well-developed,

the inner one (Cy) thin, long, straight at base, bent

upward distally parallel] with respect to antero-posterior

axis, rounded at tip, outer one (Cy) a slender, slightly

C. Ituarte, 2005

Figures 34-41. Pisidium chicha new species. 34. Holotype (MLP 6S99-1-1): outer view of right valve, 35-41. Paratypes (MLP

6899-1-2). 35. Posterior view. 36. Left valve, inner view. 37. Right valve, inner view. 38. Left valve, detail of hinge. 39. Right valve

letail of hinge. 40. Right valve, detail of cardinal tooth and ligament. 41. Posterior view, detail of ligament. Scale bars (34-37

500 um; (38, 39) = 200 um: (40) = 250 um; (41) = 100 um

Page 102

THE NAUTILUS, Vol. 119, No. 3

curved blade, quite oblique, overlapping Cz on posterior

half; anterior lateral tooth (AII) very strong, straight,

cusp high, somewhat acute, displaced distally; posterior

lateral tooth (PII) short, strong, cusp high, distal. Hinge

on right valve (Figures 37, 39, 40): cardinal tooth (C3)

well-developed, narrow, and evenly curved on anterior

half, quite enlarged into triangular, slightly grooved cup

at posterior end; slightly hanging from inner margin of

hinge plate. Lateral teeth short and robust, inner

anterior lateral (AI) somewhat curved, cusp displaced

forward; outer anterior lateral tooth (AIIL) very short,

straight, with distal cusp; inner posterior lateral one (PI)

short, straight, slender and low, cusp sub-central; outer

posterior lateral one (PHI) minute, with distal cusp.

Escutcheon lanceolate, long, outline demarcated by

a very delicate line. Ligament-pit enclosed, inner margin

gently sinuous (Figures 40, 41). Ligament moderately

strong, internal, slightly visible from exterior, but not

protruded, representing about 23% of shell length.

Anatomy: Only one (anal) mantle opening present.

Only inner demibranch present (Figure 42). Inner radial

mantle muscles weak, inserted just above or coalescent

with pallial line. Nephridium with dorsally visible lateral

loop (Figure 4:3).

Type Locality: Unnamed brook flooding into Zapla

River in the neighborhood of Zapla City (24°16’01" S,

65°07'09" W), 946 m altitude, Jujuy Province, Argen-

tina, and small springs on bank of Manzanito Rivulet, on

the road from Santa Victoria East to Yavi near Huertas

(22°14'20" S, 35°00'31" W), 2740 m, Jujuy Province,

Argentina.

Type Material: Holotype (MLP6S99-1-1), 9 para-

types (7 paratypes MLP 6899-1-2; 2 paratypes MACN-

In 36363) from 24°16’ S, 65°12’ W; 62 paratypes from

22°14'20" S, 35°00'31" W (52 paratypes MLP 6550; 5

paratypes FML 14775; 5 paratypes MNHN).

Other Material Examined: Catamarca Province:

flooded areas at side of an unnamed river on the road

to Singuil, (27°38'25" S$, 65°57'23" W), 2000 m

(MLP7203). Jujuy Province: Los Cedros Rivulet, close

to Las Maderas Dam (24°28'40" S 65°17'08" W),

1190 m (MLP 6545-2). Tucuman Province: small pools

at the side of Medina River, on provincial road 305

between El Tipal and Aserradero (MLP 7404-2).

Distribution: Catamarca, Jujuy and Tucuman_ pro-

vinces, Argentina, between 940 and 2740 m altitude.

Etymology: The name of the new species alludes to

the Chichas, a small ethnic group that was a part of the

Omaguaca people, who inhabited the lands in the

neighborhood of the type locality.

Remarks: The shell shape, quite ovate and nearly

equilateral, and the marked sculpture of Pisidium chicha

new species are distinctive features that allow for easy

identification of the new species among other Pisidium

species from northwestern Argentina. A moderate shell

43. A —s

Figures 42-43. Pisidium chicha new species. 42. Gross

anatomy. 43. Detail of nephridium. (be: brooding embryos; id:

inner demibranch; n: nephridium; pa: posterior adductor; sr:

siphon retractor).

variation was observed: the specimens from Manzanito

Rivulet (MLP 6550) show shells slightly more convex

and higher than those of the specimens from Zapla

(MLP 6899-1); the striae were slightly coarser and more

marked in the former group. Specimens from Catamarca

Province (MLP 7203) show slightly inequilateral shells,

with posterior end slightly shorter. Pisidium chicha

shares with P. ocloya the same number of mantle

openings and demibranchs and the nephridium with

lateral lobe dorsally visible; but the former species

strikingly differs in having a smaller shell with quite an

oval shell outline. The presences of only one demibranch

and one mantle opening in P. chicha clearly separate this

species from P. chiquitanum. In addition, this latter is

a larger species of relatively similar shell outline but that

also differs from P. chicha in having an almost smooth

shell surface, lower, more backward displaced beaks,

and somewhat truncated posterior end.

Pisidium chiquitanum Ituarte, 2001

(Figures 44-47)

Pisidium chiquitanum Ituarte, 2001: 50; figs. 2-14 (La Siberia,

West of Comarapa, Santa Cruz de La Sierra, Bolivia,

holotype MHNB 34734),

Diagnosis: Rather elongate shell outline, — slightly

truncated at posterior end, low and posteriorly located

beaks, ligament position, internal but externally visible,

presence of branchial and anal openings, two demi-

branchs on each side and nephridia of closed type.

C. Ituarte, 2005

Description: Shell thin, small to medium size (mean

SL = 3.9 + 0.25; maximum observed size: 4.2 mm),

not verv high (mean HI = SO + 1), not convex (mean

Ci 61 = shell outline markedly oval elongate,

anteriorly produced, posterior end short, widely

rounded, or slightly truncated and nearly straight

Figures 44, 46, 47). Beaks low, depressed, slightly

projected above dorsal margin, located at about 62% of

SL (Figures 4446).

vellowish, with fine and low commarginal — striae,

Shell surface dull glossy, straw-

moderately more accentuated towards the shell margin

Figures 44, 46)

Hinge plate solid, hinge line rather long (HiL/SL

56 + 2). Hinge on right valve: Right cardinal tooth (Cs

strongly curved in middle, quite narrow on anterior hi i

enlarged into gently sulcated, rounded, or somewhat

elongate cup. Right lateral teeth robust, inner anterior

lateral (AD), widely curved, long, cusp sub-central or

slighth displaced forward: outer anterior lateral tooth

AIIL) quite short, cusp distal; inner posterior lateral (PI)

nearly straight, short, cusp sub-central; outer posterior

lateral tooth (PII) reduced in size with distal cusp. Left

valve (Figure 47): cardinal teeth short, inner one (Cs)

short, oblique with respect to antero-posterior axis, outer

one (C4) a narrow curved lame, quite oblique, over-

lapping C2 on posterior half; anterior lateral tooth (ATI)

strong, straight, cusp sub-central: posterior lateral tooth

PII) narrow and weak, cusp distal.

Ligament-pit enclosed, deep, inner margin slightly

sinuous, concave at posterior end. Escutcheon well

marked by a delicate line; ligament long, internal, but

visible from outside in anterior half through a very

narrow and sometimes rather long gap between valves,

never protruded. Ligament length is 23 + 1% of shell

ength.

Anatomy: Anal siphon and branchial mantle opening

oresent. Presiphonz . suture rather long, representing 11

+ 2% of St

yowertul siphonal serene present. Inner radial mantle

nuscles. S bundles as rule. inse srted aWe Ly from pe alli il

ine, scars of those corresponding to anal siphon

retractors coalescent with posterior adductor muscle

scars. Inner and outer demibranchs present. Outer

demibranch reduced in size, composed of 11-15 very

short descending filaments, reaching back to the 14— 16”

filament of inner demibranch. Ne phridia of closed type.

dorsal lobe variable in shape, aed subquadrate,

with lateral loop not visible in dorsal view

Material Examined: Holotype (MHNB 34734) La

Siberia, West of Comarapa, Santa Cruz de La Sierra

Bolivia; Tucuman, Argentina: 27°01'24” S, 65°39'29" W

MLP 6554): Cerro Munoz, Santa Cruz, 26°54’ S,

65°46'42” W, 2400 m (MLP 6991): La Angostura dam,

26°56' S, 65°41'03" W, 1800 m (MLP 6527): Jujuy,

Argentina: Los Laureles, 25°06'27" S, 65°36'10" W,

1360 m (MLP 6528): Los Toldos. Santa Victoria De-

parisien 1770 m (MLP 6993): small brook near Tiraxi

23°59'57” S, 65°19'39” W), 1576 m (MLP 6552): small

Anal siphon well-developed, pair of

Figures 44-47.

Paratype

Pisidium chiquitanum Ituarte, 2001. 44, 45.

(MLP 5362). 44. Outer view of right valve. 45.

Posterior view. 46, 47. Specimen from Tiraxi, Jujuy (MLP

6552). 46. Outer view of right valve. 47. Inner view of left

500 um

valve. Scale bars =

spring at the side of provincial road No. 4, near Termas

de Reyes (24°10'36" S$, 65°28'18" W) (MLP 5497-

Distribution: Ranging from sub-Andean regions in

Siberia (west of Comarapa) in central Bolivia (1SO0 m

altitude) southward to northwestern Argentina (between

1360 and 2400 m altitude)

Remarks: Pisidium chiquitanum can be easily identi-

fied among South American Pisidium species by its oval

shell outline with low beaks and internal (however

visible from the outside) ligament. It is also character-

ized by two, inner and outer, demibranchs on each sid

two siphonal openings, and nephridia of closed type

SS SS

Page 104

THE NAUTILUS, Vol. 119, No. 3

Pisidium chiquitanum resembles Pisidium meierbrooki

ee and Hinz, 1984, from Peru and Bolivia, which is

the only known species from tropical South America

with both, branchial and anal, siphonal openings

(Ituarte, 1995). Pisidium meierbrooki differs from. P.

chiquitanum in having a more convex shell (according to

data in Kuiper and Hinz, 1984, the Ci varies between 77

and 80), fuller and more backward displaced beaks. As

pointed out by Ituarte (2001)Pisidium chiquitanum is

similar to specimens from Ecuador and Peru reported

by Kuiper and Hinz (1984), as Pisidium casertanum

(Poli, 1791), an Eurasian species extremely variable in

shell shape, currently reported as cosmopolitan (Burch,

1975; Kuiper, 1953; Kuiper and Hinz, 1984: Holopainen

and Kuiper, 1982), However, these specimens are larger

than P. chiquitanum, having more central beaks and ae

produced anterior end. P. chiquitanum also differs from

P. casertanum in having less convex shell, lower and

narrower beaks, decidedly displaced backward. The

specimens from northwestern Argentina slightly differ

from the ones from Bolivia in being generally higher,

with beaks slightly less displaced in posterior Grection

(Figure 46).

ACKNOWLEDGMENTS

The author is grateful for the warm friendship and kind

support received from colleagues Gabriela Cuezzo,

Fatima Romero, and Carlos Molineri during the three

field trips to northwestern Argentina. This study was

funded by grant: PIP 554/98 ae the Consejo Nacional

de Investigaciones Cientificas y Técnicas (CONICET),

Argentina. The author is rese ucher of the CONICET.

LITERATURE CITED

Burch, J. B. 1975. Freshwater Sphaeriacean clams (Mollusca:

Pelecypoda). Malacological Publications, Hamburg, Mi-

chigan, 96 pp.

Doello-Jurado, M. 1921. Una nueva especie de Eupera del Rio

de Ja Plata. Physis 5: 72-75.

Holopainen, I. J. and G. J. Kuiper. 1982. Notes on the

morphometry and anatomy of some Pisidium and

Sphaerium species (Bivalvia, Sphaeriidae). Annales Zool-

ogici Fennici 19: 93-107.

pn J. G. J. 1983. The Sphaeriidae of Australia. Basteria 47:

—52,

Gee J. G. J. and W. Hinz. 1984. Zur Fauna der

Kleinmuscheln in den Anden. Archiv fiir Molluskenkunde

114[1983]: 137-156.

Ituarte, C. F. 1989. Los géneros Byssanodonta dOrbigny,

1846 y Eupera Bourguignat, 1854 (Bivalvia: Sphaeriidae)

en el area parano-platense. Descripcion de Eupera

iguazuensis n. sp. del rio Iguazu, Misiones, Argentina.

Neotropica 35: 53-63. ;

Ttuarte, C. F. 1994. fpel guaraniana n. sp. (Pelecypoda:

Sphaeriidae) del rio Uruguay, Argentina. Gayana, ser.

zool. 58: 1-7.

Ituarte, C. F. 1995. Nuevos registros de Pisidium Pfeiffer,

1821 y Sphacrium Scopoli, 1777 (Bivalvia: Sphaeriidae) en

Chile, Bolivia y Noroeste argentino. Neotropica 41: 31-41.

Ituarte, C. F. 1996. Argentine species of Pisidium Pfeiffer,

1821, and Musculium Link, 1807 (Bivalvia: Sphaeriidae).

The Veliger 39: 189-203.

Ituarte, C. F. 1999. Pisidium chilense (VOrbigny, 1846) and

new species of Pisidium C. Pfeiffer, 1821 from southern

Chile (Bivalvia, Sphaeriidae). Zoosystema 21: 249-257.

Ituarte, C. F. 2000. Pisidium taraguyense and Pisidium

pipoense, new species from Northeastern Argentina

(Bivalvia: Sphaeriidae). The Veliger 43: 51-57.

Ituarte, C. F. 2001. Pisidium chiquitanum new species from

Santa Cruz de la Sierra, Bolivia (Bivalvia: Sphaeriidae).

The Nautilus 115; 50-54.

Ituarte, C. F. and M. ©. Dreher-Mansur. 1993. Enupera

elliptica n. Sp., una nueva especie en el rio Iguazu,

Misiones, Argentina. Neotropica 39: 11-16,

Pilsbry, H. A. 1911, ‘Non-marine Mollusca of Patagonia.

Reports of the Princeton University Expedition to

Patagonia 1S96—1S99, 3(Part 5): 513-633.

Strobel, P. 1874. Materiali per una malacostatica de terra e di

acqua dolce dell’ Argentina. Pisa.

THE NAUTILUS 119(3):105—LOS, 2005

Page 105

Mysella pedroana, a commensal bivalve (Lasaeidae) on two

decapod crustacean hosts

Shannon M. Carpenter

Santa Barbara Museum of Natural

History

Department of Invertebrate Zoology

2559 Puesta del Sol Road

Santa Barbara, CA 93105-2998 USA

[email protected]

ABSTRACT

Mysella pedroana (Dall. 1898), lives commensally on Isocheles

/ }

pilosus | Holmes, 1900), and Blepharipoda occide ntalis Randall,

1839. Because of their small size at reproductive maturity,

specimens attached to Isocheles pilosus were prev iously

referred to in the literature as an undescribed species.

However, preliminary comparisons between the variability of

hinge dentition and internal structure of those specimens and

small individuals of Mysella pedroana indicate that they

represent instead a single species. Variability of characters

including size at reproductive maturity and shell morphology

suggests that Mysella pedroana is a highly variable species.

INTRODUCTION

A small bivalve living attached to hermit crabs has

perplexed biologists for over a decade. It was referred to

as Mysella sp. H by Valentich-Scott and Barwick (2001),

as an undescribed species. It was shown to be

reproductively mature at 1.0 mm in length.

Bivalves in the family Lasaeidae have been taxonom-

ically problematic (Dall, 1898; Gage, 1966a; 1966b;

O Foighil and Eernisse 1988: Morton and Scott,

1989). These bivalves combine features of immaturity

induced by changes in environmental factors, perhaps

due to their commensal nature. Mysella pedroana

Dall, 1895), is no exception. It is found both as

a commensal and free-living (Scott, 1987); however,

due to its prevalence on hosts, it may prefer a eomniensal

habitat.

Mysella pedroana was previously thought to be host-

specific and found only on ee sand crab Blepharipoda

occidentalis Randall, 1839, (Burch and Burch, 1944;

Boss, 1965a; Lafferty, (on Boyko and Mikkelsen,

2002). Originally described by Dall in 1898 from a single

valve, M. pe -droana was then redescribed (Boyko and

Mikkelsen, 2002) and associated with B. occidentalis as

host. The discovery of its occurrence on another host,

the hermit crab Isocheles pilosus (Holmes, 1900),

previously undocumented variation in shell morphology,

and small size at reproductive maturity has led to this

present study,

Blepharipoda occidentalis (Decapoda: Albuneidae) is

found in the northeastern Pacific from Stinson Beach,

Marin County, California, USA, to Bahia Santa Rosalia,

Baja ee Mexico (Morris et al., 1980) both

intertidally and subtidally, burrowing in sand. Isocheles

pilosus (Decapoda: Diogenidae) ranges from Bodega

Bay Harbor, Sonoma County, California, USA, to Estero

de Punta Banda, Baja California, Mexico (Rickets et al.

1985). Like B. occidentalis, it is intertidal, but also occurs

in mud flats, bays and estuaries including depths

offshore up to 55 meters. Isocheles pilosus most

commonly inhabits shells of Polinices or Kelletia (pers.

comm. Scott, 2004) and crawls on the sand or buries

with only its eyes and mouth visible (Fager, 1965).

Mysella pedroana is either attached to the crab’s setae or

in its branchial chambers.

MATERIALS AND METHODS

Preserved specimens identified by Valentich-Scott and

Barwick (2001) as Mysella sp. H from the collections in the

Santa Barbara Museum of Natural History (SBMNH)

were studied. Additional material included SBMNH

specimens associated with preserved specimens of Iso-

cheles pilosus and B. occidentalis, along with specimens

from two living I. pilosus and one B. occidentalis collected

7 Sands Be ch, S Santa Barbara. Specimens removed from

pilosus are deposited as SBMNH 351472-351480 and

means Material of M. pedroana examined includes

SBMNH 348251, 345553, 348252, and 348253.

Of the 145 av a. specimens, 35 were opened and

Riche or gross anatomy. Characters examined were

internal structure, shell dentition, and shape. Measure-

ments were made with vernier calipers and anatomical

observations were performed under a dissecting micro-

scope. Scanning electron micrography was performed at

SBMNH with a Zeiss EVO 40 XVP with a variable-

pressure secondary electron detector.

THE NAUTILUS, Vol. 119, No. 3

Figure I.

Living Mysella perdroana were placed in Petri dishes

with sea water and coarse sediment for comparison of

movement in different media. Specimen with brood

was stained with crystal violet in distilled water prior to

examination.

RESULTS

Mysella pedroana (Dall, 1898)

Description: SHELL (Figures 1—3): Shell morphology

variable from ovate to subtrigonal, thin, more elongate

anteriorly; umbones opisthogyrate; beaks range from

central to posterior; shell surface white with poorly

defined commarginal striae; periostracum thick and

yellow, variable in texture, rough and dehiscent to

smooth and adherent; prodissoconch line present;

maximum shell length 11.0 mm, mean height to length

ratio 77% (herein) (SD +4.8; range 11.0-1.0 mm) to

80% (Boyko and Mikkelsen, 2002).

Muscle scars apparent in larger specimens with

anterior adductor scar elongate and posterior ovate;

pallial line entire; hinge with prominent subumbonal

resilium; two lateral cardinal teeth on right valve with

longer anterior tooth and slight groove, posterior shorter;

teeth diverging into an inverted V; left valve with thin

Figures 2-3. Mysella pedroana, hinge dentition. 2. From host Isocheles pilosus, SBMNH 351473, SEM. Scale bar =

3. From host Blepharipoda occidentalis, SBMNH 351478, SEM

\ crowth series of commensal Mysella pedroana, SEMNH 351473. Scale bar = | mm.

grooves (lamellae) that interlock with the right valve.

Extensive variation shown in smaller specimens (length

1-3.4 mm), which may exhibit a posterior tubercular

tooth on the right valve with a similar posterior

tubercular tooth on the left valve.

Gross Anatomy: Mantle papillate, more so anteriorly;

ctenidium encompasses large area of internal space and

serves as a brood chamber: presence of eggs observed in

specimens from 1-11 mm in length; labial palps small.

OBSERVATIONS ON Live SPECIMENS: Activity was enhanced

following regular replacing of cold sea water. Clams

were observed crawling on surface of Petri dish with

shell positioned vertically, in the manner of a gastropod;

dug in sand with anterior end, process took 15-20 sec.

Detached M. pedroana took 3 min. to reattach to

setae on the ventral surface of I. pilosus. However,

when I. pilosus setae were easily accessible and M.

pedroana was placed next to them, attachment took 30-

45 sec.

Host INFORMATION AND PREVALENCE: Blepharipoda occi-

dentalis had a carapace length of 4.06-4.55 cm (n = 3)

and all three had Mysella pedroana present. Isocheles

pilosus had a carapace length of 1.9-2.57 cm (n = 6)

with three of the six crabs having bivalves present.

200 Lm.

Scale bar = 1 mm

S. M. Carpenter, 2005

Page 107

Figures 4-5.

Three B. occidentalis were examined with 100%

prevalence. Of the six preserved museum specimens of

I. pilosus, only one was infested. However, two live

I. pilosus were examined and fifty-nine M. pedroana

specimens were retrieved.

The live B. occidentalis examined had two M.

pedroana that were found on the external surface near

the antennae and on the second pereopod.

Host Isocheles pilosus: Mysella specimens on I. pilosus

were found in the branchial chambers, attached to the

chelae, the junction between the chela and carapace, the

ventral setae and branchial chambers (Figure 4). The

largest specimens were on the chelae (1 mm) with

smaller individuals on the ventral surface (0.7—1 mm)

and juveniles (<0.7 mm) anteriorly in the right and left

branchial chambers. Those on the ventral surtaee and in

the branchial chambers were attached with byssus

(Figure 5). The few on the chelae were observed

crawling or attached by byssal threads to the surface

spines of the chelae. Upon preservation in 70% ETOH

M. pedroana retained its byssal threads.

DISCUSSION

This study suggests that shell shape of Mysella pedroana

is more variable than previously appreciated. These

variable characteristics include texture of the periostra-

cum (rough and dehiscent to smooth and adherent),

dentition, “shape of the shell. These differences have

been attributed to environmental conditions in other

molluscan shells (Wellington and Kuris, 1983; De Wolf

et al., 1998).

Shell variation probably led to the redescription of

Mysella pedroana as Rochefortia golischi by Dall in 1916

(Burch and Burch, 1944); these were later synonymized

by Scott (1957). However, this variation is also common

to many commensal species and makes it difficult to

Myse o pe droana. 4. Host Isocheles pilosus showing attachment between the chela and carapace (arrows), SBMNH

345553, Scale bar = 2.2 mm. 5. On host Isocheles pilosus with byssus (arrow), SBMNH 351472

. Seale bar = 0.8 mm.

identify them based on shell morphology (O Foighil and

Eernisse, 1988: Morton and Scott, 1989).

Due to shell variation and the small size at re-

productive maturity of M. pedroana, these smaller

bivalves on I. pilosus were thought to be a new species,

referred to as Mysella sp. H (Valentich-Scott and

Barwick, 2001). However, minimum size of brooding

adults of M. pedroana had been previously reported at

1 mm (Valentich-Scott and Barwick, 2001) and sper-

matogenesis has been confirmed in specimens of

1.2mm. Previous observations for individuals of M.

pedroana undergoing spermatogenesis were 4.7 mm in

length (pers. comm. Kevin Lafferty, 2004). Reproductive

maturity at small sizes has been reported also for

Pseudopythina macrophthalmensis at 2.0 mm (Jespersen

et al., 2001) and Mysella bidentata with egg production

at 1.7 mm (O Foighil et al., 1954).

The prevalence of Mysella pedroana was higher on B.

occidentalis than on I. pilosus. Those on B. beotdentalis

were also larger in the gill chambers compared to only

juveniles found in the gills of I. pilosus. Further

examination of both hosts would be required to see if

there is a host preference.

Reattachment to a host is possible for the species.

Both Lafferty (pers. comm. 2004) and Valentich-Scott

(pers. comm. 2004) noted that individuals could drop

their byssus and disassociate from the host if perturbed.

Individuals of M. pedroana could use this procedure to

move from host to host or to become free-living.

The above observations on morphology and symbiont-

host relationships permit the identification of this

bivalve as Mysella pedroana. Previously reported host

specificity for commensal bivalves may be due to the

scarcity of studies or experimental observations (Boss,

1965b; Morton and Scott, 1989). This species was

previously thought to be host-specific (Boyko and

Mikkelsen, 2002), an assumption here shown not to be

valid.

Page 108

THE NAUTILUS, Vol. 119, No. 3

ACKNOWLEDGMENTS

I would like to thank to Paul Valentich-Scott for his

guidance, knowledge and for introducing me to the

world of bivalves "The Hearst Fonndaton Internship

through the Santa Barbara Museum of Natural History

made this research possible. Scanning electron micro-

scope facilities funded by NSF grant number

MRI0420726, Daniel Geiger assisted with SEM images.

Michael Caterino, Henry Chaney and Armand Kuris for

reviewing preliminary drafts. Shane Anderson provided

living eae: Patricia Sadeghian contributed her knowl-

edge of crustaceans. Paula Mikkelsen and Kevin Lafferty

whom offered additional data. Kelvin Barwick contrib-

uted many specimens. Two anonymous reviewers

offered critical observations which greatly improved

the manuscript.

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THE NAUTILUS 119(3): 109-115, 2005

Page LO9

Coralliophila trigoi (Gastropoda:

Muricidae), a new species

from the none ers Atlantic Ocean

Paolo Mariottini'

Dipartimento di Biologia

Universita di “Roma Tre”

Viale Marconi 446

00146 Roma

ITALY

mari [email protected]

Carlo Smriglio

Via di Valle

00167 Roma

ITALY

Aurelia 134

[email protected]

Emilio Rolan

Museo de Historia Natural,

Campus Universitario Sur, 15752

Santiago de Compostela

SPAIN

[email protected]

ABSTRACT

Based on shell characters and with further support from mo-

lecular data, Coralliophila trigoi, a new species of gastropod of

the family Muricidea, is here described from the northeastern

Atlantic Ocean. The new taxon, consisting of several specimens

mainly collected along the Atlantic Spz inish coast, has previ-

ously been misidentified in the literature as Coralliophila bas-

ilea (Dautzenberg and H. Fisher, 1896). Coralliophila trigoi

new species is conchologically similar to Coralliophila meyen-

dorffii (Caleara, 1845), and Coralliophila panormitana (Mon-

terosato, 1869), but it can be easily separated from them mainly

because it is differently sculptured. The new species is com-

pared with other members of the genus Coralliophila from the

same geographical area and Me .diterranean Sea. Molecular se-

quencing of the intemal transcribed spacer 2 region (ITS2) of

the nuclear rDNA and part of the ditochondeal gene for 12S

rDNA confirm the validity of the new species.

INTRODUCTION

The coralliophilines form a monophyletic group of neo-

gastropods that includes approximately 200-250 de-

scribed species he based on their shell morphol-

ogy, in at least 7-LO “genera”, distributed worldwide in

temperate and tropical oceans. The subfamily Corallio-

philinae Chenu, 1859 (for the phylogenetic relationship

of this muricoidean groups, see Oliverio and Mariottini,

2001a) includes species invariably associated with cni-

darians, which are generally used as food by the gastro-

pods. Shell variability, absence of radula, absenes of a

preserved protoconch (often eroded in adults and even

in young specimens), together with a limited knowledge

of the anatomy, represent constrains to the understand-

ing of the taxonomic status of this group of neogastro-

pods. Their classical systematics above the species level

is at present far from. being stable (Clover, 1982; Bou-

chet and Warén, 1985: Kosuge and Suzuki, 1985: Oliv-

erio, 1989: Vaught; 1989; Olivers. in press). Data from

‘ Author for correspondence

mitochondrial and nuclear genes (12S rDNA and ITS2

rDNA, respectively) have been recently utilized in the

proposal of a molecular framework for the phylogeny of

these muricids (Oliverio and Mariottini, 2001a; Oliverio,

Cervelli and Mariottini, 2002). Data from both sequence

= secondary structure show that Rapaninae Gray, 1853

=Thaidinae Jousseaume, 1585) are their sister group

iat asewych et al., 1997; Oliverio and Mariottini, 200 1a;

Oliverio, Cervelli and Mariottini, 2002), indicating a

monophyletic radiation of the Coralliophilinae. The

state-of-the-art knowledge about feeding, anatomy, sex-

ual strategies, parental care, and protoconch of corallio-

philines was recently reviewed by Richter and Luque

(2002). The authors ‘reported the available data on pro-

toconch and larval development of many coralliophilines

belonging to ten different genera, including Corallio-

phila H. and A. Adams, 1853. We had the opportunity

to examine several shells of a coralliophiline that we

could allocate to any of the Atlantic and Mediterranean

species of this subfamily. These shells, mostly collected

along the coast of Galicia, Spain, were prev iously mis-

identifie d in the literature as Coralliophila basilea

(Dautzenberg and H. Fisher, 1896) (Rolan, 1953; Rolan,

Lopez and Gutiérrez-Garcia, 1995). After comparisons

with other species, we realized that they represent an

undescribed species, possibly related to Coralliophila

meyendorffti (Caleara, 1845) and Coralliophila panor-

mitana (Monterosato, 1569).

In order to verify the taxonomic validity of Corallio-

phila trigoi, we carried out ee sequencing of the

ater transcribed spacer 2 region (ITS2) of f the nucle-

ar rDNA and of part of the mitochondrial gene for 12S

rDNA. Genomic DNA was extracted from the

foot of two freshly collected individuals with standard

methods (SDS-proteinase K digestion, phenol/chloro-

form extraction, ethanol precipitation (Oliverio and Mar-

iottini, 2001b)). Mitochondrial rDNA was amplified

through the ieee rase chain reaction (PCR) with prim-

ers 12S-I and 12S-IIT (Oliverio and Mariottini, 2001a)

Nuclear ribosomal ITS2 was amplified using the primers

dissected

Page 110

THE NAUTILUS, Vol. 119, No. 3

Table 1. Collecting data and DDBJ/EMBL/GenBank accession number for specimens assayed in molecular systematics.

Species/individuals

Accession number

Coralliophila neritoidea

Coralliophila brevis

Coralliophila mejendorffii

Coralliophila panormitana

Coralliophila trigoi new species,

speci. #1]

Coralliophila trigoi new species,

specim. #2

La Maddalena Is. (

Cape Circeo (Latium, Italy

Taiwan, 23°10’ N, 120°05’ E, 5 m depth

Collecting locality and depth 128 ITS2

AJ293679 AJ420258

La Maddalena Is. (Sardinia, Italy), 41°15’ N, 009°26' E, 30 m depth — AJ293676 — AJ420256

Sardinia, Italy), 41°15’ N, 009°26' E, 3-7 m depth AJ297517 — AJ293661

), 41°11" N, 013°04" E, 70 m depth AJ293681 — AJ420259

Camarifias, Galicia, Spain, northeastern Atlantic Ocean, 15-50 m depth — AJ937305 — AJ937307

Camarifias, Galicia, Spain, northeastern Atlantic Ocean, 15-50 m depth —AJ937306 — AJ937308

its-3d and its-4r complementary to conserved regions of

the ribosomal coding portions on the 5.8S and 28S

tRNAs (Oliverio and Mariottini, 2001b). PCR-amplified

products were directly sequenced by an automated se-

quencer. Nucleotide sequences were first aligned by

hand and the alignment progressively optimized : accord-

ing to secondary structure homology. Phylogenetic anal-

yses_ were performed using PAUP? 4b10 ( (Swofford,

2002). GenBank accession auimber 12S and ITS2) of

the Coralliophila trigoi sequences are reported in Ta-

ble 1.

Institutional abbreviations used: MNCM, Museo

Nacional de Ciencias Naturales, Madrid, Spain; MZB,

Laboratorio di Malacologia, Museo di Zoologia

dell Universita di Bologna, fialy.

Abbreviations used for collections: CS-PM, Carlo

Smriglio and Paolo Mariottini (Rome, Italy); ER, Emilio

Rolén (Vigo, Spain); FS, Frank Swinnen (Lommel, Bel-

gium); Ir J uan Trigo (Brion, A Coruna, Spain); MO,

Marco Oliverio (Rome, Italy).

SYSTEMATICS

Superfamily Muricoidea Rafinesque, 1S15

Family Muricidae Rafinesque, 1815

Subfamily Coralliophilinae Chenu, 1859

Genus Coralliophila H. and A. Adams, 1853

Type Species: Fusus neritoideus Lamarck, 1816,

Ency. Meth., pl. 435, figs. 2a—b. (=Purpura violacea Kie-

ner, 1836), by subsequent designation (Iredale, 1912).

Recent, Indo-Pacific.

Coralliophila trigoi new species

(Figures 1-8, 13-14, 17-21)

Description: Shell of large size, up to 35 mm length.

Protoconch usually erode d in adult specimens (proto-

conch observed in only one juvenile Daas n, albeit

worn and lacking the embryonic sti we), Protoconch in-

dicative of plz inktotrophic larval development, multispi-

ral, composed of protoconch I and If. Protoconch IL of

about 2% whorls, with a diameter of about SOO jm,

showing two strong spiral keels, only one visible above

the suture of the first whorl, crossed by axial ribs forming

nodules at intersections. The protoconch- teleoconch de-

marcation is well-defined, marked by a varix. Teleoconch

shape biconical, elongate ovoid-fusiform, solid, rather in-

flated. Spire relatively high, conical, 4-5 rather convex

whorls, shoulder rounde a Suture not very evident, par-

tially covered by the sculpture, which is formed of 15—

20 spiral cords, regularly ordered, rarely alternating with

smaller spiral ponds: Spiral sculpture consisting of ribs

rounded in cross-section, all of similar width, densely

covered with imbricating, fine, and long lamellae. Axial

ribs 7—11, large, generally weak, crossing the spiral

cords. Siphonal ae short, narrow, open, moderately

curved. Aperture large, oval, representing about half of

the shell height, w /hite or cream-white inside. Umbilicus

absent. Outer lip thin and crenulate. Shell color uni-

formly reddish- or pale-brown. Operculum oval, oblong,

horny, concentric, with lateral-terminal nucleus, weddishe

brown.

Type Material (Figures 1-8): Holotype (Figures 1-2),

30.6 * 20.4 mm, MNCN 15,05/46458; paratype A (Fig-

ures 3-4), 31. ° x 19.4 mm, MZB 31023; paratype B

(Figures 5-8), 27.3 x 16.7 mm, CS-PM; paratype C,

27.4 X 18.2 mm, CS-PM: paratype D, 24.2 * 16.5 mm,

ER; paratype E, 26.5 * 16.6 mm, ER; paratypes A-E,

from type locality; paratype F, 30.9 * 18.5 mm, ER,

Malpica, Spain; paratype G, 23.5 * 16.3 mm, ER, Ca-

melle, Spain; paratype H, 24.8 < 17.3 mm, JT; paratype

I, 21.2 x 12.3 mm, JT; pe aratype L, 26.9 * 17.1 mm, ”

paratype M, 21.4 * 12.3 mm, JT; paratype N, 31.2

18.0 mm, JT; paratype O, 18.5 * 12.5 mm, JT; ears

P. 7.4 * 5.1 mm, CS-PM; paratypes H—P from Porto da

es Sagres, Portugal; paratype Q, 30.2 * 18.1 mm,

ER, Lira A Coruna, Spain; paratype R, 26.4 * 17.2 mm,

ER, Malpica, Spain; paratype S, 27.3 * 16.2 mm, CS-

PM; paratype T, 25.1 * 16.2 mm, CS-PM; paratypes

Q-T from Lira A Coruna, Spain; paratype U, 39.2 x

23.2 mm, CS-PM, A Guarda, Spain; paratype V, 30.1 x

18.6 mm, CS-PM, type locality; paratype W, 29.8 x 17.4

mm, CS-PM: paratype Z, 25.5 < 16.2 mm, CS-PM;

paratypes W-Z from Malpica, Spain.

Type Locality: Camarifias, Galicia, Spain, northeast-

ern Atlantic Ocean, 15-50 m depth.

Distribution: Known from Galicia, Spain, to Algarve,

Portugal, in the Atlantic Ocean, and from Malaga and

Almeria (Alboran Sea, Spain) in the Mediterranean.

P. Mariottini et al., 2005

Figures 1-8. Coralliophila trigoi new species. 1-2. Holotype, 30.6

19.4 mm, MZB 31023. 5-8. Paratype B, 27.3

Habitat: Several live collected specimens were found

attached at the base of host cnidarians Calliactis paras-

itica (Couch) (a sea anemone).

Etymology: This species name is dedicated to our

friend Juan Trigo, who supplied some of the specimens

of the new species

Molecular Sequencing: Given the often misleading

information conveyed by characters of shell morphology

in this group Oliverio and Mariottini, 2001b), we veri-

20.4 mm, MNCN 15.05/46455. 3-4. Paratype A, 31.6

16.7 mm, CS-PM. From type locality, depth 15-50 m. Scale bars = 1 em

fied the validity of the new species using a molecular

appre vach. We sequenced the internal transcribed spacer

2 region (ITS2) of the nuclear rDNA and part of the

mitochondrial gene for 12S rDNA from individuals of

the new taxon and compared them with available se-

quences of C, meyendorffii, C. panormitana, and Cor-

alliophila brevis (Blainville, 1832), plus Coralliophila

neritoidea (Lamarck, 1816) as outgroup. Parsimony anal-

ysis of the aligned sequences ol nuclear and mitochon-

Sed

drial DNA resulted in the tree reported in Figure 2%

THE NAUTILUS, Vol. 119, No. ¢

1s)

Figures 9-16. Shells of Coralliophila species. 9-10. Coralliophila squamosa (Bivona, 1838), 35.9

Galicia, Spain, 15-50 in depth. 11-12. Coralliophila panormitana (Monterosato, 1869), 19.2

20.9 mm, CS-PM,. Camarinas

12.1 mm, CS-PM, Malaga, Spain, 40 m

depth. 13-14. Coralliophila trigoi new species, 27.2 * 18.1 mm, MO, Marbella, Spain, 30-50 m depth. 15-16. Coralliophila meyendorffii

Calcara, 1845), 26.8

The two related species C. meyendorffii and C. panor-

mitana, were more closely related to each other than to

the two specimens of the new species

Other Material Examined: Coralliophila trigoi: 5

spec. from the tvpe locality: 3 spec., Laxe: | spec, A

Guarda 3 spec. amelle 3 spec Malpica, Galicia, Spain

ER: 2 spec. Sagres \loarve, Portugal, CS-PM; 2 spec

\lmeria; 3 spec. Marbella, Spain, MO; Coralliophila

meyendorffii: 10 spec. La Maddalena Isl., Sardinia Is-

land: 1 spec. San Pietro Isl., Sardinia Island; 6 spec. San-

ta Marinella; 3 spec. Ponza Isl.; 1 spec. Capo Palinuro

» SPE Le Castella: 5 spec Elba Isl.. Italy, CS-PM: 6

14.7 mm, FS, Punta del Carmen, Lanzarote Isl., Canary Islands, 20 m depth Seale bars = 1 cm

spec. Punta del Carmen, Lanzarote Isl., Canary Islands,

FS Coralliophila panormitana: 5 spec. Marina di Ca-

merota, Italy; 1 spec. Malaga, Spain; 1 spec. Portimao,

Portugal, CS-PM: Coralliophila squamosa: 23 spec. from

the type locality: 6 spec Laxe; § spec. Camelle; 7 spec.

Malpica, Galicia, Spain, ER; 10 spec. Ria de Vigo, Ga-

licia, Spain, JT; 1 spec. Malaga, Spain; 1 spec. San Pietro

Isl., Sardinia Island: 1 spec. Ventotene Ish: ] spec. Ma-

rina di Camerata; 2 spec. Le ( rastella: 2 spec. Civitanova

Marche, Italy, CS-PM

Remarks: Based on shell characters of the teleoconch

we take the conservative approach of conserving this

P. Mariottini et al., 2005

Figures 17-26.

species in the genus Coralliophila s. |., as traditionally

ormulated (see also Oliverio, in press). The new species

was collected in Galicia together with the Atlantic-Med-

terranean Coralliophila squamosa (Bivona, 1838). This

atter species is generally collected in the Mediterranean

Sea at depths ranging from 100 to 600 m (Figures 9-

10). The new taxon is clearly conchologically distinguish-

able from all other eastern Atlantic and Mediterranean

species of Coralliophilinae. Coralliophila trigoi shows a

certain resemblance with the Mediterranean Corallio-

phila panormitana (Monterosato, 1869), but the latter is

smaller, having a different sculpture that includes a larg-

er number of spiral cords (22-24), with smaller scales

Figures 11-12

Coralliophila trigoi is similar to Coralliophila meyen-

dorffii (Calcara, 1845) (Figures 15-16, 22-26), but it dif-

fers by its more rounded shape, by having the length/

width and length/aperture lensth ratios smaller (1.60

and 1.68 vs. 1.72 and 1.86), by possessing a larger num-

ber of primary spiral cords (15-20 vs. 13-15), which are

narrower in width and differently sculptured and by a

reddish- or pale-br« yWwTh color Coralliophila meyendorffii

1S generally milky-white Furthermore, the protoconch

Shells of Coralliophila species. 17-20. Coralliophila trigoi new species, 23.6

14.0 mmm, CS-PM, Algarve

Portugal. 18, 19. Details of shell sculpture. 21. Coralliophila trigoi new species, detail of the larval whorls, paratype P, 7.4 * 5.1

mm, CS-PM, Porto da Baleeira, Sagres, Portugal, 15-25 m depth. 22-25. Coralliophila meyendorffti (Calcara, 1545), 25.6 16.0

mm, CS-PM, La Maddalena Isl., Sardinia, Italy, 1 m depth. 23, 24. Details of shell sculpture. 26. Coralliophila meyendorffi, detail

of the larval whorls, specimen size 2.2 * 1.3 mm, CS-PM, La Maddalena Isl., Sardinia, Italy, 1 m depth. Scale bars =

or Figures 21, 26, scale bars = 200 xm

I cm, except

[1 of Coralliophila trigoi shows a number of whorls (2%

and a diameter (SOO xm) different from the values on

Coralliophila meyendorffii (3% and 650-750 jum, re-

spectively; see Figures 21, 26). The main shell morpho-

logical differences between these two species are sum-

marized in Table 2. Coralliophila meyendorffii is a lit-

toral, widely distributed species, occurring in the Med-

iterranean Sea (Figures 22-26), along the Atlantic

African coast and Canary Islands (Figures 15-16). It is

worth mentioning that Coralliophila meyendorffii preys

on a variety of anthozoans, including the cnidarian Cal-

liactis parasitica (personal observations). Interestingly,

Coralliophila trigoi seems to be distributed along the

\tlantic coast of Spain and Portugal, but restricted to

the Alboran Sea (Malaga, Almeria) within the Mediter-

ranean basin. The collecting depth is slightly deeper (50

m) than the bathymetric range (littoral) of ¢ oralliophila

meyendorffti Although the

velopment (as indicated by t

phila trigoi could bring abou

bution than that currently

planktotrophic mode. of de-

he protoconch) of Corallio-

it a wider geographic distri

Known for the species the

taxon has not yet been found in the Macaronesian Is

lands and the West African coast

Page 114

THE NAUTILUS, Vol. 119, No. 3

C. neritoidea

C. brevis

C. panormitana

100/97 C. meyendorffii

C. trigoi #1

100/100

C. trigoi #2

Figure 27. Cladogram of parsimony analysis for the studied

coralliophiline taxa. This topology was recovered under maxi-

mum parsimony analysis of the 12S + ITS2 dataset (either

including or excluding gap positions). Numbers at the branch

represent ee support in MP analyses (gap included and

gaps as missing). Locality data and GenBank accession num-

bers (128 and ITS2) are reported in Table 1.

Coralliophila trigoi has been previously misidentified

as Coralliophita pasilee (Rolan, 1983: 236, fig. 210; Ro-

lan, Lopez and Gutiérrez-Garcia, 1995: 30, fig.2 2), but it

is easily distinguishable from this taxon. palo:

basilea has a more turreted shell and a coarser spiral

sculpture. Among the fossil coralliophiline records, the

only species that slightly resembles the new taxon is Cor-

alliophila burdigalensis (Tournouér, 1874), a species

from the Upper Oligocene and Lower Miocene of Aq-

uitaine (France); but the fossil species is smaller and has

a different shell outline ane sculpture (Lozouet and

Renard, 1998: 173, figs. 2. 1-10).

Parsimony aay sis of the oe d sequences of nuclear

and mitochondrial DNA resulted in the tree re ported in

Figure 27. Accordingly, the two related species C. mey-

endorffii and C. panormitana were more closely related

to each other than to the two specimens of f the new

species. The same results (not shown here) were ob-

tained including additional (yet shorter) sequences of C.

meyendorffii from other Mediterranean localities (Sar-

dinia, Sicily, and Southern Spain). This is a clear indi-

cation that the spe cimens of the new form constitute a

distinct, isolated gene-pool and strongly support our de-

cision to describe it as new.

ACKNOWLEDGMENTS

We would like express our deep gratitude to Mr. Juan

Trigo, Brion, A Coruna, Spain, for kindly sending us

Table 2. Comparison of shell characters between Corallio-

phila trigoi and C. meyendorffi.

Shell characters C. trigoi C. meyendorffii

Protoconch diameter 800 jzm 650-750 jum

Protoconch number of

whorls 2.5 3.5

Teleoconch primary spiral

cords 15-20 13-15

Teleoconch axial ribs 7-11 8-10

LengthAvidth ratio 1.60 + 0.09 1.72 + 0.09

Length/aperture length

ratio 1.68 + 0.10 1.86 + 0.10

Size range 17.6-35.4 mm = 19.5-34.3 mm

ees of Coralliophila trigoi. Dr. Andrea Di Giulio

(Department of Biology, University of “Roma Tre”,

Rome, Italy) is acknowlec Iged for SEM photographs,

which were carried out at the LIME (Inter-Department

Laboratory of Electron Microscopy, University of “Roma

Tre”). Sincere thanks are due to Dr. Antonio Bonfitto

(Zoological Museum, University of Bologna, Italy) for

generously providing bibliography. We are crateful to

Drs. Marco Oliverio and Maria Vittoria Modiea (De-

partment of Animal and Human Biology, University of

“La Sapienza”, Rome, Italy) for help with molecular

work. M. Oliverio also proy ided valuable advices and dis-

cussion.

LITERATURE CITED

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Atlantic bathyal and abyssal Neogastropoda excluding Tur-

ridae_ (Mollusca, Gastropoda). Bollettino Malacologico,

Suppl. 1: 1-296.

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the Coralliophilidae family. Privately printed, i-ii, 18 pls.

+ 18 unnumbered text pages.

Harasewych, M. G., S. L. Adamkewicz, J. A. Blake, D. Saudek,

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eee

Iredale, 1912. New generic names and new species of ma-

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odique ‘de »s trois regnes de la nature”. 1-16, pls 391-488,

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Kosuge, S. and M. Suzuki. 1985. Ilustrated catalogue of La-

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Oliverio, M. 1989. Famiglia Coralliophilidae Chenu, 1896 in

Mediterraneo. La Conchiglia 246-249: 45-54.

Oliverio, M. In press. Coralliophilinae (Neogastropoda: Muri-

cidae) from the South West Pacific. In: P. Bouchet and V.

Heros (eds.) Tropical Deep-Sea Benthos. Memoires du

Muséum national d'Histoire naturelle, Paris.

Oliverio, M. and P. Mariottini. 200la. A molecular framework

for the phylogeny of Coralliophila and related muricoids.

Journal of Molluscan Studies 67: 215-224.

Oliverio, M. and P. Mariottini. 2001b. Contrasting morpholog-

ical and molecular variation in Coralliophila meyendorffii

(Muricidae, Coralliophilinae). Journal of Molluscan Stud-

ies 67: 243-246.

Oliverio, M., M. Cervelli and P. Mariottini. 2002. ITS2 rRNA

evolution and its use in the phylogeny of muricid neogas-

tropods (Caenogastropoda, Muricoidea). Molecular Phy-

logenetics and Evolution 25: 63-69.

Richter, A. and A. A. Luque. 2002. Current knowledge on Cor-

alliophilidae (Gastropoda) and phylogenetic implication of

anatomical and reproductive characters. Bollettino Mala-

cologico, Suppl. 4: 5-15.

Rolan, E. 1983. Moluscos de la Ria de Vigo I. Gasterépodos.

Thalassas, Anexo 1: 1-383.

Rolan, E., D. Lopez and G. Gutiérrez Garcia. 1995. Nuevas

citas de moluscos de Galicia. Noticiario SEM 25; 20-21.

Swofford, D. L. 2002. PAUP®. Phylogenetic Analysis Using

Parsimony (°and Other Methods). Version 4 [1998], beta

4.0b10 [2002]. Sinauer Associates, Sunderland, Massachu-

setts.

Vaught, K. C. 1989. A classification of the living Mollusca.

“American Malacologists Inc., Melbourne, 143 pp.

THE NAUTILUS 119(3):116-117, 2005

Page 116

Book Review

Land Snails of British Columbia

Forsyth, Robert G. 2004. Land Snails of British

Columbia. Royal BC Museum Handbook. Royal BC

Museum, Victoria, 192 pp. ISBN 0-7726- 5218- X. Black

and white illustrations, plus a section of color photo-

graphs.

Even as “cutting edge” systematics shifts its focus

from morp hological to molecular data sets, the need for

identification mianaals.] yased on real-world field marks

and accessible characters has never been greater. When

zoologists with landcare agencies or environmental

consultants ask for references on land snails, it hardly

helps to refer them to ponderous, outdated, and hard to

find tomes such as Pilsbry (1939-1945). So, manuals like

Land Snails of British Columbia are a welcome resource.

Land Snails of British Columbia is the latest in a Royal

BC Museum series of natural history books that began in

1942. It is the first work to describe and illustrate the 92

species of terrestrial mollusks known to occur in British

Columbia, Canada. Because all of the species treated

occur outside that province as well, its utility extends

beyond those borders. Identification aids include

a paragraph of physical description, usually brief, to

the point, and adequate; a sentence or two of

comparison with other, potentially confusing species;

clear illustrations (the line-and- stipple drawings of many

of the smaller kinds are noteworthy); and keys to, first,

the genera of snails and slugs, and then to the species in

each multi- species genus. For slugs, where the di-

agnostic characters are often internal, excellent, pre-

dominantly original, drawings of the distal genitalia are

provided. (The simple dissection needed to access these

features should be within the range of all to whom this

book is directed.) The language of the descriptions is

simple and intelligible, and a gloss: ury of technical terms

is included. The de -scriptive material is followed by

a summary of each species’ distribution, concise com-

ments on natural history, etymology of the generic and

spe cific name, remarks the it, among other things, upd ite

the species’ taxonomic history (e.g., since Pilsbry, 1939—

1948) or point out special conservation status, and selected

references pointing to the extensive and well researched

bibliography. All of this adds up to a highly functional

volume that will serve its intended audience well.

A six-page section of color photographs of living snails

and slugs, mainly by Kristina Ovaska, is not only

attractive but also helpful, particularly in showing the

habitus of livin

¢ slugs, something not always well

conveyed. by drawings.

The information is amalgamated (the author's word)

from published literature, his own observations, scien-

tific collections, and personal communications with

other workers. The distributions are based in part on

unpublished records from the Royal BC Museum and

other collections. Specific localities are not cited: three

online resources by the same author (Forsyth, 1999,

2005a, 2005b) ) provide more detail for specific regions.

A significant innovation is the introduction of Euconu-

lus (Euconulus) praticola (Reinhardt, 1883) to the North

American fauna. Its distinctness from E. (E.) fulvus

(Miiller, 1774) is well documented by characters of shell

and external anatomy. The former catchall “Zonitidae” is

correctly parsed into Pristilomatidae, Gastrodontidae, and

Daudebardiidae for regional genera, in keeping with

recent work by Hausdorf and others. Monadenia, iconic

of northwest American forests, is correctly assigned to

Bradybaenidae rather than the redundant and poorly

argued Monadeniidae of Nordsieck (1987) and Schileyko

(1996). In these and other ways, the author shows himself

to be well “booked up” on the current and evolving

literature of land snail systematics.

Overall, the taxonomy is simplified, appropriately for

a general interest manual. Synonyms are merely listed,

not referenced, as few pote mtial users will need that

entry into the primary literature. Subspecies are treated

in the remarks, not in the major taxon headings or in the

taxonomic checklist (pp. 23-28). If experience in Cali-

fornia is any guide, upon further study some—perhaps

most—nominal subspecies will be shown to be species:

and examination of data not available to prior mono-

graphers may uncover local endemics, possibly in former

glacial refugia such as the Queen Charlotte Islands. For

the present, however, this handbook provides a reliable

source of biodiversity information. Author Forsyth is

fully aware of the dynamic character of taxonomic

ani ilysis and astutely states (p. 2), “this book is a work in

progress.” By clear ly showing the state of our know ledge

at this time, he has contributed greatly to that process.

LITERATURE CITED

Forsyth, R. G. 1999. Terrestrial Gastropods of the Columbia

Basin, British Columbia. Royal BC Museum, Victoria. http://

www.livinglandscapes.be.cé Vcbasin/molluses/contents. html

[html ve rsion] or http:/Avww.livinglandscapes.be.ca/cbasin/

molluses/pdf/molluse3.pdf {pdf version ]. ania August

20, 2005.)

Forsyth, R. G. 2005a. Terrestrial Gastropods of the Peace

River Region — Northern Rockies of British Columbia.

Royal BC Museum, Victoria. http://livinglandscapes.be.

ca/prnr/prnr_snails/index.html [html version] or http://

livinglandscapes.be.ca/prnr/snails/prnr_snails.pdf [pdf ver-

sion|. (Accessed August 20, 2005.)

Forsyth, R. G. 2005b. Terrestrial Gastropods of the Upper

Fraser Basin of British Columbia. Living Landscapes,

Royal BC Museum, Victoria. http://livinglandscapes.be.ca/

Book Review, 2005

Page 117

uppertraserbasin/ufb_snails/index.html [html version| or

http://livinglandscapes.be.ca/uppertraserbasin/snails/

ufb_snails.pdf [pdf version]. (Accessed August 20, 2005.)

Nordsieck, H. 1987. Revision des Systems der Helicoidea

(Gastropoda: Stylommatophora) Archiv fiir Mollusken-

kunde 118: 9-50

Pilsbry, H. A. 1939-1948. Land Mollusca of North America

north of Mexico). Academy of Natural Sciences of

Philadelphia, Monograph 3, 1 (1): i-xvii + 1-573 + i-ix

(1939); 1 (2): iwi + 574-994 + i+ix (1940); 2 (1): i-viii + 1-

520 + i-ix (1946); 2 (2): i-xlviii + 521-1113 (1948),

Schileyko, A. A. 1996 Guamampa n.g. (Gastropoda

Pulmonata), a bradybaenid land snail with monadeniid

characters. Bulletin du’) Muséum National d’Histoire

Naturelle, Section A, Zoologie, Biologie et Ecologie

Animales 1S: 401-408

Barry Roth

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Volume 119, Number 4

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ONT KN)DR_13

CONTENTS ISSN 0028-1344

Ellen E. Strong A morphological reanalysis of Pleurocera acuta Rafinesque, 1831,

and Elimia livescens (Menke, 1830) (Gastropoda: Cerithioidea: Pleuroceridae) 119

Richard L. Squires New Late Cretaceous (Santonian and Campanian) gastropods from

LouElla R. Saul California and Baja California, Mexico... 0.0... ee 133

M. G. Harasewych Daffymitra lindae, a new genus and species of Volutomitridae

Yuri I. Kantor (Neogastropoda) from the Bellingshausen Abyssal Plain ............... 149

Sven N. Nielsen Exilia alanbeui, a new species from the Neogene of central Chile:

the first record of Exilia (Gastropoda: Ptychatractidae) from South America 153

Patricia Miloslavich Spawn of Amphissa sp. and Cosmioconcha sp. (Caenogastropoda:

Ana Karinna Carbonini Columbellidae) from the Colombian Caribbean ........0.0.0.0.0.0.0.0.0.000. 157

Juan Manuel Diaz

Néstor E. Ardila

Thomas J. DeVries Pterorytis pacanana new species (Gastropoda: Muricidae): circumstantial

evidence for late Pliocene El Nino events in southern Peru ............ 164

Eliézer de Carvalho Rios A new species of Falsimargarita (Gastropoda: Vetigastropoda: Trochidae)

Luiz Ricardo L. Simone from the South Atlantic Ocean... 0. 169

PACU Iy ois. os. oh aye ae ns eee iad ela G Gd oR ob HG ew hd @ ache Sadak bY Did oa des wathe Da hassle io wae 163

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THE NAUTILUS 119(4):119-132, 2005

A morphological reanalysis of Pleuwrocera acuta Rafinesque,

1831, and Elimia livescens (Menke, 1830)

(Gastropoda: Cerithioidea: Pleuroceridae)

Ellen E. Strong

Department of lave rtebrate Zoology

National Museum of Natural History

Smithsonian Institution

P.O. Box 37012-MRC 163

Washington, DC 20013-7012 USA

[email protected]

ABSTRACT

Pleurocera acuta and Elimia livescens have been the subject of

several anatomical and ecological studies and are two of the

most thoroughly documented species of North American

Pleuroceridae. Yet significant gaps still remain in our un-

derstanding of their structure. Consequently, the anatomy of

these two species is re-described, allowing a re-interpretation

of pallial oviduct homologies; features not previously portrayed

in the literature (midgut and Kidney) are newly described.

These taxa are characterized by the presence of an ovipositor,

a kidney with a subdivided internal lumen that invades the

pallial roof, a prostate with a highly folded anterior spermato-

phore-forming region, and a pallial oviduct with spermato-

phore bursa but lacking a seminal receptacle. This analysis

verifies the degree of similarity between the two species, but

a number of differences were identified including features of

the ovipositor, pallial oviduct, prostate, anterior esophagus,

midgut, kidney, pericardium and nervous system. Comparison

to other pleurocerids confirms that species distributed

Western North America (Juga) and Asia (Hua, Semisulcospira)

share the presence of a seminal receptacle—a feature that is

lacking in all described Eastern North American species.

INTRODUCTION

The freshwater Pleuroceridae Fischer, 1855, comprises

one of the most species-rich assemblages of limnic

mollusks occurring in North America and Eastern Asia.

In North America, they are represented by eight genera

(Athearnia Morrison, 1971, Elimia H. and A. Adams,

1854, Jo Lea, 1831, Juga H. and A. Adams, 1854,

Leptoxis Rafinesque, 1819, Lithasia Haldeman, 1540,

Pleurocera Rafinesque, 1$18, and the extinct Gyrotoma

Shuttleworth, 1845) and estimates of 159. species

currently considered valid; of these, 34 are extinct and

59 are listed as critically imperiled (G1) or imperiled

G2) (Johnson et al., 2005).

The current concept of the family Pleuroceridae (e.g.

Bouchet and Rocroi, 2005) can be traced to the works of

Thiele (1928, 1929) who recognized 6 subfamilies within

the heterogeneous “Melaniidae” (an invalid name for

Thiaridae Gill, 1871), including the Pleurocerinae. An

alternative, highly polyphyletic view promoted by

Morrison (1954) caused great confusion for more than

four decades concerning the extension and indepen-

dence of the Pleuroceridae and Pachychilidae P. Fischer

and Crosse, 1892 (e.g. Ponder and Warén, 1988). Recent

work on the systematics of limnic lineages within the

Cerithioidea Fleming, 1822 (e.g. Glaubrecht, 1996,

1999: Kohler et al., 2004) has resolved some of this

confusion and supports the distinctiveness of these

families on both morphological and molecular grounds.

However, molecular data ( (Lydeard et al., 2002) do not

support monophyly of the Pleuroceridae as currently

defined, but suggest that a clade of eastern North

American species are more closely related to Melanopsis

ge ie H. Adams and A. Adams, 1854) than to

a clade of western North American (Juga) and eastern

Asian pleurocerids (Semisulcospira Boettger, 1586, Hua

Chen, 1943).

As noted by Woodard (1934), anatomical data are

critical in refining the phylogenetic relationships and

classification of these species. However, all taxonomic

treatments of the family have been based primarily on

conchological characters in the absence of a broad

comparative understanding of morphology. Although the

subject of numerous ecological studies (e.g. Dillon,

2000) and comparably many descriptions exist for the

radula, operculum and life history of North American

pleurocerids, surprisingly little is known about their

basic biology and anatomy. Scant information on the

anatomy (besides radula and operculum) has been

provide sd for Elimia Shai (Say, 1829) (Woodard,

1934), E. potosiensis (Lea, 1841) (Jones and Branson,

1964) and Pleurocera canaliculata (Say, 1821) (Magru-

der, 1935b):; among western North American forms, only

reproductive anatomy is known for several species

(Prozorova and Rascheph bkina, 2004). In addition, the

Page 120

THE NAUTILUS, Vol. 119, No. 4

classically cited work of Dazo (1965) on the natural

history, ecology, distribution and anatomy of Pleurocera

acuta Rafinesque, 1831 and Elimia livescens (Menke,

1830) has long stood as the most comprehensive

morphological snaey of any pleurocerid gastropod. As

such, that study has formed the basis for hypotheses of

homology in recent higher-order phylogenetic studies

based on morphology (e.g. Houbrick, 1988; Glaubrecht,

1996). Yet, Dazo’s study ¥ was completed long before our

present understanding of the structure, function and

homologies of canbhioidean reproductive anatomy was

in place, particularly through the work of R. S. Houbrick

on marine species. We alee have a much more thorough

understanding of midgut structure and its utility oy

revealing phylogenetic affinities of cerithioideans (Glau-

brecht and Strong, 1999: Strong and Glaubrecht, 1999,

unpublished data).

Given the persistent paucity of anatomical data

available for the family, the goal of this contribution is

to reevaluate the morphology ‘and putative homologies of

the two species described by Dazo, and to place them

within the emerging fr amework now available for limnic

cerithioideans. This j is critical for ongoing morphological

and phylogenetic studies of coithioide: in gastropods,

and in particular for clarifying the monophyly and

systematic affinity of the family Pleuroceridae.

MATERIALS AND METHODS

This study is based on collections of individuals of

Pleurocera acuta and Elimia livescens living sympatri-

cally in the Mukwonago River at the outflow from Lower

Phantom Lake, Mukwonago, Waukesha County, Wis-

consin (42°51.402 N: 88°19.767 W). Populations were

sampled in May and June; only reproductively mature

individuals were used for observations of reproductive

anatomy. Individual specimens were cracked, preserved

in 95% ethanol and were not relaxed. Voucher material

is deposited in the National Museum of Natural History

in Washington (USNM).

As stated above, Dazo (1965) provided a rather

detailed account of external features (operculum, shell,

ovipositor, color patterns of the head-foot) as well as of

the radula and all internal organ systems (alimentary,

nervous, excretory, respiratory, \ vascular, and reproduc-

tive systems). Baker (1928) also provided information on

the ope erculum, jaw, 1 radulae, and external anatomy of the

two species: additional scattered accounts relating to the

radula, external anatomy and life history are also

available (e.g. Jewell, 1931; Howe, 1938; Goodrich,

1945). As such, the following reanalysis emphasizes

anatomy of the soft parts. As the two species are

overwhe Imingly similar, a thorough account is provided

for Pleurocera acuta — the type species of the type genus

for the family; only details that differ are noted for

Elimia livescens. However, the internal kidney structure

of P. acuta, while displaying the same configuration as E.

livescens, is too occluded with excretory tissue to allow

a clear representation. As such, only the internal

structure for the latter species is illustrated.

Specimens were examined using a Leica MZ 12,5

binocular microscope with camera lucida; visualization

of structures was enhanced through the use of aqueous

toluidine blue. Typically four to five specimens were

aaa for each organ system investigated, especially

for those systems showing high 1 evels of individual and/

or seasonal variation (i.e. reproductive system); a mini-

mum of two individuals were examined for particularly

complex structures (i.e. midgut, nerves). Descriptions of

midgut morphology are given with the stomach opened

dorsally and the style sac uppermost. Unless otherwise

indicated, an incision is made along the extreme right,

upper and lower margins, and the ior deflected laterally

to the left. Terminology follows Strong and Glaubrecht

(2002, 2003).

RESULTS

Pleurocera acuta Rafinesque, 1831

Material Examined: Wisconsin: Mukwonago River:

USNM 1081522, 1081524.

External Anatomy: Operculum sub-ovate, corneous,

dark reddish-brown in color, with 3.5 whorls; paucispiral

with | large, eccentric nucleus of approximately 2.75 to 3

whorls (Figure 1). Final whorl moderately inflated.

Nucleus occupying slightly under 1/2 (~43%) of total

length.

Foot ovate with narrow propodium; anterior pedal

gland oe along anterior margin (Figure 2, ap).

Ovipositor (ovp) located on side of neck below right

cephalic tentacle. Ciliated egg groove extending Shore

distance up side of neck from ovipositor, sh: allowing past

mantle margin; groove fading near anal aperture. Two,

partially juxtaposed, parallel folds, extending into

ovipositor pore from aperture (*); one fold at upper

posterior edge, second at lower anterior edge of

ovipositor (when viewed laterally), forming obliquely

flattened H-shaped lumen. Folds unequal in size;

anterior fold along floor significantly larger than

posterior fold along roof. Ovipositor pore expanding

medially into head-foot, then curving and narrowing

slightly posteriorly. Two vertical lane of “H” unequal

Sich that posterior limb forming rather narrow channel

along posterior wall. Anterior limb inflated and expand-

ing medially into foot, forming large flattened, sub-

triangular chamber. Folds diminishing toward blind tip

of pore. Short grooved tract extending ventrally from

ovipositor aperture toward foot sole, but not reaching

edge of foot.

Most individuals with straight sr ieee but some

with curved anterior tip (Figure 3, os). Hypobranchial

gland highly developed with pendulous, transverse folds

(Figure 4, hg).

Alimentary System: = Forecurt. Buccal mass short and

stout, extending to base of cephalic tentacles (Figure 3,

E. E. Strong, 2005 Page 12]

Zp ", int

555

*) indicate folds extending into ovipositor. 3. Mantle cavity and anatomy of cephalic hemocoel. Dorsal view, anterior is below

Hypobranchial gland removed for clarity. 4. External view of organs in visceral mass. Dotted line indicates extent of pericardium

under main kidney chamber. Abbreviations: ap, anterior pedal gland: b, bladder; bm, buccal mass; em, coc ineel muscle; et,

ctenidium: e, esophagus: hg, hypobranchial gland; int, intestine; kd, main kidney chamber; me, mantle edge; nr, circum-esophageal

nerve ring; op, operculum; sea uapintaes Ov, Ovary; Ovp, Ovipositor; per, pericardium; po, pallial oviduct: ae ane intestinal sinus;

r, rectum; rt, buccal mass retractor muscle; sg, salivary gland: sn, snout; sp, supra-esophageal ganglion: ss, style sac; sto, stomach

Scale bars = 1 mm.

Figures 1-4. Anatomy of Plewrocera acuta. 1. Operculum. 2. Ovipositor and egg groove. Right lateral view of side of foot. Asterisks

Page 122

THE NAUTILUS, Vol. 119, No. 4

bm). Odontophore occupying majority of buccal cavity

with small, glandular subradular organ protruding before

radula. Small jaws present at anterior ends of dorsal

folds: epithelium of buccal cavity overlying dorsal folds

glandular (stippled region). Shallow, non-glandular

buccal pouches extending underneath dorsal folds

adjacent to buccal ganglia at rear of buccal cavity.

Radular sac short, curving upward behind base of buccal

mass. Robust buccal retractors (rt) inserting onto lateral

walls of cephalic hemocoel adjacent to cerebral ganglia

(nr). Short, glandular mid-ventral fold forming smi all flap

just behind ‘odontophore in anterior esophagus, flanked

laterally by two ventro-lateral folds. Ventro-lateral folds

converging short distance behind mid-ventral fold and

continuing through mid-esophagus (e). Mid-esophagus

long, hearing paired longitudinal ventral and dorsal

folds. Epithelium between dorsal and ventral folds

weakly glandular and irregularly striated; septate esoph-

ageal ele and lacking. Dorsal and ventral folds converging

and fusing at distal end of mid- esophagus before

continuing into posterior esophagus and subdividing

into numerous folds equal in height. Long, tubular

salivary glands (sg) opening dorso-laterally to buccal

cavity ‘alongside odontophore, passing through circum-

esophageal nerve ring (nr), almost reaching posterior

esophagus.

MIpcuT.

midgut floor (Figure 5, e). Marginal fold (mf) extending

anteriorly from esophageal aperture alongside major

typhlosole (t1), then turning posteriorly bordering right

margin of sorting area (sa). Groove present along

midline of marginal fold (mf) for much of its length:

groove fading ‘proximally and distally. Sorting area

elongate- triangular, tapering posteriorly; posterior tip

curving slightly to the left around wedge-shaped sorting

area pad ( sap). Accessory marginal fold (amf) ) emerging

from esophageal aperture, paralleling marginal fold wae

curving around posterior tip of sorting area; fold

bifurcating at posterior end of gastric Charer to form

two folds. Fine parallel striations extending anteriorly

from esophagus up face of major typhlosole. Midgut roof

to the left of sorting area coarsely folded and einsieulane

ized (eu). Gastric Shiela (gs) ) small and delicate, strongly

concave, with narrow, tubular posterior end and more

flaring, flattened anterior end; shield continuous with

cuticle of stomach roof and crystalline style ea (p).

Glandular pad (gp) large and broadly rounded. Cres-

centic ridge (er), bounding deep crescentic groove,

extending from esophageal aperture and fusing to right

side of elandular pad. Paired digestive gland ducts (dd)

opening to deep pocket near proximal tip of crescentic

ridge. Shallow caecum (ec) extending ventrally under

glandular pad behind gastric shield. Single longitudinal

fold (ef) extending from caecum (ce) ne posterior

end of gastric chamber. Prominent fold (wu) extending

from right side of style sac lip, along floor ne crystalline

style pocket, to base of major ty phlosole: fold bounding

u-shaped depression below lip . style sac (ss). Style sac

Esophagus opening under ledge on left side of

and intestinal groove communicating along entire length.

Crystalline style present.

Hinpcur. Proximal intestine (Figure 4, int) passing

below distal tip of style sac (ss), then extending

posteriorly alongside style sac to main gastric ch siber

(sto). Intestine curving anteriorly, with broad loop

overlying proximal style sac. Intestine extending under

posterior end of main kidney chamber (kd), entering

pallial roof alongside bladder (b) and pallial gonoduct

(Figure 3, po), continuing forward to papillate anus near

mantle margin (1).

Reno-pericardial System: Kidney oe three

interconnected chambers (Figure 4, 17, kd, b). Main

chamber (kd) ) lying along dorsal os ‘of body whorl,

anteriorly surrounding pericardium (Figure 4, per),

crossing axis of body from right to left and extending

short ditadce into pallial roof at base of mantle cavity.

Chamber occluded anteriorly (within pallial roof) with

excretory tubules. Posterior ly, main chamber with small,

narrow lumen, dorsally enclosing intestine. Second

chamber (see exposed chamber in Figure 17) extending

between pericardial chamber to right body wall below

intestine, forming small bladder (Figure 4, b). Chamber

mostly occluded by vertical sheets of excretory tissue

radiating from afferent renal vessel (see Figure 17, arv),

and communicating to mantle cavity via large nephro-

pore (np). Sheets of excretory tissue branching and

anastomosing, and fusing to right lateral floor, roof and

walls: vertical sheets highly branched anteriorly and

forming comparatively dense honeycomb of excretory

tissue. Bladder penetrating connective tissue along right

side of body, short distance into mantle roof. Excretory

tissue separating small ventral chamber below, within

pallial portion ( (dottec | line). Size of ventral chamber, as

well as branching pattern and number of excretory

sheets of tissue variable between individuals. Small

aperture just behind afferent renal vessel connecting

main chamber and bladder (arrow). Nephridial gland

absent.

Pericardium voluminous (Figure 4, per), extending to

right side of body (dotted line).

Nervous iia Circum-esophageal nerve — ring

(Figure 3, nr) lying immediately behind buccal mass,

at base of hs alic tentacles. Cerebral ganglia (Figure 6,

ce) connected by short, stout commissure, each pro-

ducing seven nerves (optic, statocyst, tentacular, and

four labial nerves). Buccal connectives short, innervating

buceal ganglia lying ventro-laterally at base of buccal

cavity immediately behind buccal retractor muscles.

Pleural ganglia (pl) lying behind and below cerebral

ganglia connected to cerebral ganglia by short, thick

connectives. Pedal ganglia (pe) with two prominent

anterior nerves and five smaller accessory nerves. Small

statocysts (st) with approximate ‘ly 10-15) statoconia

present dorsally alongside pedal ganglia behind pedal

connectives. Sub- esophage val ganglion (sb) joined to left

pleural ganglion by thickened connective (co); connec-

E. E. Strong, 2005

Figures 5-6.

Anatomy of Plewrocera acuta. 5. Midgut anatomy. Dorsal view, anterior is uppermost. 6, Circum-esophageal nerve

ring. Frontal view on the left, right lateral view on the right. Abbreviations: amf, accessory marginal fold: ¢, caecum: ef, caecal fold:

ce, cerebral ganglion: co, thickened connective between left pleural and sub-esophageal ganglia: er, crescentic ridge: cu,

cuticularized region of stomach roof; dd, duct of digestive gland; e, esophageal aperture; gp, ¢ nd ates pad; gs, gastric shield; int,

intestine; mf, marginal fold; p, crystalline style pocket; pe, pedal ganglion; pl, pleural ganglion; pn, pallial nerve from left pleural

ganglion; sa, sorting area; sap, sorting area pad; sb, sub- esophageal ganglion; sp, connective to supra-esophageal ganglion; ss, lip of

style sac; st, statocyst; tl, major typhlosole: u, u-shaped fold; z, zygoneury. Scale bars = 1 mm.

tive producing 1-3 small nerves (n=2). Zygoneury (z)

formed between sub-esophageal and right pleural

ganglia. In addition to zygoneury (z), sab: esophageal

ganglion (sb) producing one other prominent nerve and

connectives to the left pleural (pl) and visceral ganglia.

Long connective uniting right Sosa and supra-

esophageal ganglia (Figures 3, 6, sp), the latter lying

on left side mantle floor near midline of osphradium.

Dialyneury formed between pallial nerve of left pleural

ganglion ‘and osphradial nerve of supra-esophageal

ganglion at junction of mantle roof and floor. Single

visceral ganglion present between pericardium and

kidney at base of mantle cavity, above posterior

esophagus on the right. Ganglion producing two

prominent nerves.

Reproductive System: Mate. Narrow vas deferens

(Fisure 8. vd) emerging ventrally from testes, continuing

forward along ventral midline of whorl. Short, distal

portion of vas deferens thickened and forming straight

seminal vesicle. Vas deferens narrowing and curving

dorsally to enter posterior end of prostate (Figure 7, pr)

at base of mantle cavity. Prostate glandular, forming

flattened tube, opening to mantle cavity through broad

slit along entire length except for a short fused segment

at base of mantle cavity (arrows). Lateral lamina

epithelium thin along aperture; short distance into

lumen, glandular tissue forming flattened longitudinal

shelf. Glandular tissue diminishing in thickness at

anterior and posterior ends of gonoduct. Shelf undercut

along much of its length by flattened sinus (Figure §,

dotted line). Glandular tissue forming central rounded

mass flanked by shallow trough along sonoductal groove.

Medial lamina unevenly glandular, oath irregular troughs

and ridges; glandular tissue diminishing in thickness

anterior @ and posteriorly (Figure 9). Epithelium of

medial lamina posterior 1/5 smooth and concave, partially

separated from anterior region by prominent curving

ridge; ridge embracing corresponding rounded elandular

mass in lateral lamina. Ridge continuous anteriorly with

curving trough formed oe opposing ridges of tissue.

Trough fading anteriorly, and becoming fl: inked by short,

shallow flap running along gonoductal groove.

Femare. Gonad (Figure 4, ov) dorsally pease cues

digestive gland (except for a narrow ventral strip) from

tip of visceral mass to posterior end of midgut (sto).

Oviduct emerging ventrally from ovary. Renal oviduct

(Figures 10, 11, ovi) ) deflected dorsally behind mantle

cavity before entering base of glandular les il oviduct.

Pallial oviduct, with proximal albumen (ag) and distal

capsule glands (eg). Albumen and capsule glands

forming narrow bands with opposing flattened surfaces

bounding gonoductal groove (Figure 11, gg); non-

glandular portions of medial and lateral laminae formed

by thickened connective tissue (et), Proximal segment of

albumen gland at base of ahs cavity under pallial

kidney extension rather short and straight ( (Fieures 10,

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THE NAUTILUS, Vol. 119, No. 4

Figures 7-11.

Reproductive anatomy of Pleurocera acuta. 7. External, left lateral view of prostate. Anterior is to the left. Arrows

indicate extent of opening to gonoductal groove. 8. Internal aspect of prostate lateral lamina. Dotted line designates extent of sinus

under glandular shelf. 9. Internal aspect of prostate medial lamina. Anterior is to the right. Note parallel folds and groove at anterior

end, representing presumptive spermatophore forming region. 10. External, left lateral view of ee oviduct. Anterior is to the left.

Arrows indicate extent of opening to gonoductal groove. 11. External, right lateral view of pallial oviduct. Anterior is to the right.

Abbreviations: ag, albumen gland; eg, capsule g gland; et, connective tissue; gg, gonoductal groove; ovi, renal oviduct; pr, prostate;

sg, sperm gutter; spb, spermatophore bursa; vd, vas deferens. Scale bars = 1 mm.

11, ag). Anterior to pallial kidney chamber, albumen

gland curving under distal tip of bursa to the right, then

arcing dorsally. Capsule gland comprising approximately

anterior 1/4 of pallial oviduct; externally capsule gland

not inflated (Figure 10, eg). Broad aperture along entire

length of pallial oviduct except for a short fused segment

at base of mantle cavity (Figure 10, arrows). Above

aperture, sperm gutter (sg) opening in medial lamina at

anterior tip of pallial oviduct and deepening posteriorly;

gutter leading to short, blind spermatophore bursa

(spb). Seminal receptacle absent.

Elimia livescens (Menke, 1830)

Material Examined: Wisconsin: Mukwonago River:

USNM 1081521, 1081523.

External Anatomy:

, Operculum ovate, corneous, dark

reddish-brown in color, with three whorls; paucispiral

with small, basal nucleus of approximately 2.5 whorls

(Figure 12). Last whorl expanding rapidly. Nucleus

comprising approximately 1/5 of total length.

Foot broad and rounded, with wide propodium and

long anterior pedal gland along anterior margin

(Figure 13, ap). C iliated egg groove extending short

distance up side of neck from ovipositor; groove fading

near base of tentacle. Parallel folds extending into

ovipositor pore from aperture (*) roughly equal in size.

Grooved tract extending ventrally from ovipositor to sole

of foot at junction of propodium and mesopodium, just

behind termination of anterior pedal gland.

Osphradium with curved anterior tip; some individu-

als with mostly straight osphradium (Figure 14, os).

ela ae gland well developed with deep, trans-

verse folds ( Figures 14, 15, hg).

Alimentary System: FrorEGUT. Epithelium between

dorsal and ventral folds glandular and irregularly to

transverse ly striated. Dors: l and ventral folds diminish-

E. E. Strong, 2005

Page 125

Figures 12-15. Anatomy of Elimia livescens. 12. Operculum. 13. Ovipositor and egg groove. Right lateral view of side of foot.

Asterisks (*) indicate folds extending into ovipositor. 14. Mantle cavity and anatomy of cephalic hemocoel. Dorsal view, anterior is

below. 15. Extemal view of organs in visceral mass. Dotted line indicates extent of pericardium under main kidney chamber.

Abbreviations: ap, anterior pedal gland: b, bladder; bg, buccal ganglion; et, ctenidium; e, esophagus; hg, hypobranchial gland; int,

intestine; kd, main kidney chamber; me, mantle edge; nr, circum-esophageal nerve ring; op, operculum; os, osphradium; ev, ovary;

Ovp, ovipositor: per, pericardium: po, pallial oviduct; r, rectum; rt, buccal mass retractor muscle; sg, salivary gland; sp, supra-

esophageal ganglion: ss, style sac; sto, stomach; t, cephalic tentacle. Scale bars = 1 mm

Page 126

THE NAUTILUS, Vol. 119, No. 4

Figures 16-17.

Anatomy of Elimia livescens. 16. Midgut anz tomy, Dorsal view, anterior is uppermost. 17. Kidney anatomy.

Internal view of bladder. Lateral view, anterior is to the right. Right we all of bladder and adhering sheets of excretory tissue remov ed

to reveal interior; cross-hatching indicates cross-section through sheets of tissue. Arrow indicates opening in septum allowing

communication between bladder and main kidney chamber.

Dotted line anteriorly indicates extent of ventral chamber.

Abbreviations: amf, accessory marginal fold; arv, afferent renal vessel; e, caecum: ef, caecal fold; er, crescentic ridge; cu,

cuticularized region of stomach roof; dd, duct of digestive gland; e, esophageal aperture; gp, glandular pad; gs, gastric shield; int,

intestine; kd, main kidney chamber; mf, marginal fold; np, nephropore; p, crystalline style poc ‘ket; sa, sorting area: sap, sorting area

pad: ss, lip of style sac; t1, major typhlosole; u, u-shaped fold. Scale bars = 1 mm.

ing at distal end of mid-esophagus (Figure 14, e) but

continuous into posterior esophagus. Posterior esopha-

gus narrow, bearing numerous folds of equal height.

Long, tubular salivary glands (sg) just reaching transition

to posterior esophagus.

Mipcut. Groove along midline of marginal fold lacking

(Figure 16, mf). Single, weak, caecal fold (ef) along right

side of midgut behind gastric shield, opposite caecum.

Hinpcur. Proximal intestine (Figure 15, int) passing

below distal tip of style sac (ss), then extending

posteriorly alongside style sac almost reaching main

gastric chamber (sto). Intestine curving anterior rly with

loop partially overlying proximal style sac. Intestine

ue nding under posterior end of main kidney chamber

(kd), entering pallial roof alongside blad der (b) and

pallial gonoduct (po), continuing forward to papillate

anus near mantle margin (Figure 14, r).

Reno-pericardial System: — Bladder (Figure 15, b) largely

occluded by vertical sheets of excretory tissue radiating

from afferent renal vessel (Figure 17, arv) and fusing to

right lateral floor, roof and walls. Sheets of excretory tissue

loosely and regularly branching and anastomosing;

posterior sheets less branched than those anteriorly.

Pericardium rather narrow and short (F igure 15, per).

extending to intestinal loop (dotted line)

Nervous System: Buccal ganglia (Figure 14, bg) lying

dor so-laterally at base of buccal mass between buccal

retractor muscles (rt) and salivary glands (sg). Thick-

ened connective (Figure 18, co) between left pleural and

sub-esophageal ganglia producing 1-2 nerves (n=2). In

addition to zygoneury (z) and connectives to left pleural

and visceral gangli ia, sub- ae ae ganglion producing

2-3 prominent nerves (n=2 _ Small statocysts (st) with

approximately 20-30 ee

Reproductive System: Mace. Flattened longitudinal

shelf of glandular tissue within lateral lamina diminish-

ing in thickness at anterior and posterior ends of

prostate (Figures 19, 20, Deep longitudinal cleft

opening in glandular shelf along gonoductal groove at

midpoint of gonoduct and extending anteriorly; cleft

closing a short distance back from anterior tip of

gonoduct (Figure 20, cl). Medial lamina thinly and

rather evenly glandular along its length; glandular

tissue slightly diminishing anteriorly ( Figure 21). Epi-

thelium of medial lamina posterior ‘third smooth

and strongly concave, separated from anterior 2/3 by

prominent curving ridge; ridge embracing correspond-

ing groove in glands of lateral lamina at proximal end of

cleft. Epithelium of medial lamina anterior 2/3 irregu-

larly and variably crossed by oblique and longitudinal

ridges,

E. E. Strong, 2005

Page 127

Figure 18. Circum-esophageal nerve ring of Elimia lives-

cens. Frontal view on the left, right lateral view on the right.

Abbreviations: ee, cerebral ganglion; co, thickened connective

between left pleural and sub- esophageal ganglia; pe, pedal

ganglion; pl, pleural ganglion; sb, a esophageal g ganglion; sp,

connective to supra- esophi eal ganglion; st, statocyst; Z,

zygoneury. Scale bar = 1 mm.

Femate. Renal oviduct (Figures 22, 23, ovi) deflected

dorsally behind mantle cavity before entering base of

glandular pallial oviduct. Proximal albumen ole nd rather

long and initially curved, then forming straight segment

along base of mantle cavity under pallial kidney

extension. Capsule gland comprising approximately

anterior 1/3 of pallial “avidick externally capsule gland

markedly saflated: Above aperture, short distance bak

from anterior tip of oviduct (~1/5 of length), sperm

gutter (sg) sama: in medial lamina and: deepening

posteriorly : gutter leading to short, blind spermatophore

bursa (spb).

DISCUSSION

Whatever may be discovered in the future regarding the

structure and relationships of species currently placed

within the Pleuroceridae, given that Plewrocera acuta is

the type species of the type genus for the family, this

description will necessarily remain as the standard for

the application of the name.

COMPARISON OF PLEUROCERA ACUTA AND ELIMIA LIVESCENS

Baker (1925) commented on the considerable uniformity

of structure in the genitalia of these two species and that

the soft parts do not seem to show the same degree of

differentiation as the shells. Although this statement was

based on external observations, Dazo (1965) similarly

noted that, except for differences in size, the internal

anatomy of the two species is quite similar or often

identical in all organ systems; the most. significant

differences were those relating to size and shape of the

operculum, snout, tentacles, foot, and radula.

Yet, notions of similarity and how similar two entities

must be to be characterized as “identical” are subjective

concepts. The thorough documentation of these two

species has been provided to allow a more objective

means of assessing the degree of similarity between the

two. Of course, any aimatounical re ndering will maintain

some element of subjectivity.

With this in mind, the present analysis confirms that

Pleurocera acuta and Elimia livescens are remarkably

similar, both in overall organization and in many details.

This level of similarity is perhaps not une xpecte od given

the sister group relationship between the two genera

supported in one molecular analysis (Holznagel and

Lydeard, se However, it should be noted. that

monophy ly of these genera has not been demonstrated

unambiguously ( (e.g. ‘Sides, 2005), but awaits confirma-

tion within a more comprehensive phylogenetic frame-

work. Thus, the present results may indicate a closer

systematic affinity than currently appreciated.

Yet, a number of differences between the two species

are apparent. Externally, the two differ in development

of the hypobranchial gland, but this is difficult to

quantify. They also differ in the position of the ovipositor

and its relationship to the foot sole. This is consistent

with described differences in the mode of egg capsule

transfer to the substrate; in Plewrocera acuta, only the

everted walls of the ovipositor function in oviposition

(van Cleave, 1932), but in Elimia laqueata—a species

with an ovipositor configuration identical to E. livescens

(see below)—both the everted lips of the ovipositor and

the margins of the finely noued tract guide the ova to

the sibetrane (Woodard, 1934).

The most significant differences in midgut structure

are length of caecal fold, and the presence of a groove

along the marginal fold; however, these variations may

be attributable to preservation artifacts. The significance

of other minor differences (size and/or shape of caecum,

glandular pad, major typhlosole, gastric shield, crescen-

tic ridge, field of parallel striations above esophageal

aperture) can only be ascertained once a broader

sampling of species from both genera have been

examined, But these may also be attributable to

preservation artifacts and/or intra- specific variation.

Internal structure of the kidney differs only in that the

vertical sheets of excretory tissue are more highly and

densely branched within the bladder in Ple UurOCera

acuta. It was observed that the amount of such excretory

tissue varied between individuals in a species and with

maturity. Although a highly qualitative character, the

degree of difference between the two species surpasses

that of intra- specific variation.

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THE NAUTILUS, Vol. 119, No. 4

Figures 19-23. Reproductive anatomy of Elimia livescens. 19. External, left lateral view of prostate. Anterior is to the left. Arrows

indicate extent of opening to gonoductal groove. 20. Internal aspect of prostate lateral lamina. Note deep cleft at anterior end (el),

representing presumptive spermatophore forming region. 21. Internal aspect of prostate medial lamina. Anterior is to the right. 22.

External, left lateral view of pallial oviduct. Anterior is to the left. Arrows indicate extent of opening to gonoductal groove. 23.

External, right lateral view of pallial oviduct. Anterior is to the right. Abbreviations: ag, albumen gland; eg, capsule | gland; cl, deep

cleft; et, connective tissue; gg, gonoductal groove; ovi, renal oviduct; sg, sperm gutter; spb, spermatophore bursa; vd, vas deferens.

Seale bars = 1 mm.

The configuration of the nerve ring and visceral loop is

largely idence between the two. One difference is the

dumber of nerves issuing from the sub-esophageal/left

pleural connective and from the sub- esophageal gangli-

on. However, given the observed intra-specific variation,

these difiesences likely fall within the range of individual

variation.

In contrast to the results presented here, Dazo (1965)

reported 9 cerebral nerves, only a single nerve from the

visceral ganglion, and an inconstant number of accessory

pedal nerves: however, Dazo did confirm the unique

thickened left pleural/sub-esophageal connective. Dazo

also commente a on the unlikely generality of Rose-

water's (1961) findings that pleurocerids differ primarily

in the lengths of the cerebral commissure and left

pleural/sub-esophageal connective (n=6 for 9 species).

Indeed, these lengths were found to be sometimes

conspicuously different between individuals examined in

the present study.

Males of the two species differed in the pattern of

folds within the anterior region of the prostate. Regard-

less, the anterior region is inferred to be the site of

spermatophore formation rather than the comparatively

smoother posterior region. This conclusion seems

justified given the similarity in the configuration of the

folds as compared to overall form and shape of the

spermatophore (Jewell, 1931; Dazo, 1965). Limnic

cerithioideans in the family Paludomidae Stoliezka,

1868, have separated the glands in this anterior region

to form a hollow tube that has been implicated in

spermatophore formation (Glaubrecht and shone.

2004). That discovery further supports the notion

that sperm packets are produced anteriorly and suggests

that this function may be homologous in different

lineages.

For a summary of these and other morphological

differences, see Table 1

COMPARISON TO OTHER PLEUROCERIDS

With minor rer eee published accounts agree on the

main patterns of pleurocerid anatomy. Thus, pleurocer-

ids have long been known to be dioecious and oviparous

(except Se misulcospira) ) with an ovipositor involved in

the ee of the egg capsules (Stimpson, 1564).

Members of the family are also aphallate, with open

gonoducts, a gonad thi it dorsally surrounds the digestive

eland, and produce see nt-shaped spe rmatophores

(e.g. Stimpson, 1564; Jewell, 1931; Woodard, 1934, 1935;

Maprude r, 1935b; Jones and Branson, 1964; Dazo,

1965). Like other cerithioideans, pleurocerids possess

E. E. Strong, 2005

Table 1. Summary of morphological differences between Plewrocera acuta and Elimia livescens.

Pleurocera acuta

Elimia livescens

External Anatomy:

Propodium

Ovipositor ventral groove

Curved anterior tip of osphradium

Alimentary System:

Ventral folds at posterior end of mid-

esophagus

Groove along marginal fold

Length of caecal fold

Hindgut loop

Reno-Pericardial System:

Bladder excretory tubules

Pericardium

Nervous System:

Buccal ganglia

Statoconia

Nerves from sub-esophageal/left pleural

connective

Nerves from sub-esophageal ganglion

Reproductive System:

Spermatophore-forming region

Sperm gutter

Junction of renal and pallial oviduct

Proximal albumen gland

Narrow

Does not extend to foot margin

Sometimes present

Fused

Present

Long

Extends to main gastric chamber

Densely and highly branched

Extends to right body wall

Ventro-lateral

~ 10-15

1-3

2 connectives and 2 nerves (including

zygoneury)

Parallel folds and trough in medial lamina

Extends to anterior tip of oviduct

Renal oviduct curves dorsally to straight

segment of albumen gland

Short, straight segment between renal

oviduct and posterior end of bursa

Broad

Extends to foot margin

Often present

Untused

Absent

Short

Does not reach main gastric chamber

Loosely and regularly branched

Extends to inte cael loop

Dorsal

~20-30

1-2

2 connectives and 3—4 nerves

(including zygoneury)

Deep cleft in lateral lamina

Does not extend to anterior tip of

pallial oviduct

Renal oviduct ventrally joins curved

portion of albumen gland

Initially curved, with long, straight

segment to posterior end of

spermatophore bursa

two types of glands within the pallial oviduct — a feature

not previously documented among eastern North

American species.

The gut is characterized by the presence of tubular

salivary glands, a crystalline style and a style sac in

restricted communication with the cen intestine

(Magruder, 1935a, b; Itagaki, 1960; Dazo, 1965). The bi-

lobed nature of the kidney has been noted before

(Magruder, 1935b; Itagaki, 1960), but the internal sub-

division of the organ had not been previously documented.

The nervous system is consistent with many other

cerithioideans (left dialyneurous, long connective between

the right pleural and supra-esoph nBdeal ganglia, single

visceral ganglion) (e.g. Strong, 2003; Strong and Glau-

brecht, 2002, 2003), uk is distinguished by the presence

of an enlarged connective between the left pleural and

sub- -esophageal g ganglia and a zygoneurous connection on

the right—the ites a highly homopk istic character in the

Cerithioidea (see review in Houbrick, 1988). However, the

presence/absence of zygoneury among pleurocerids re-

quires confirmation as it has been de spicted as dialyneury

in several other species (Magruder, 1935b; Itagaki, 1960).

Only the former study confirmed the presence of the

enlarged left pleural and sub- sh ae connective.

Additionally, published descriptions (Magruder, 1935b;

Itagaki, 1960; Dazo, 1965; present study) disagree on the

“umber of nerves produced by various ganglia, but as

noted above, this can be highly vageule even within

species. However, the eumber of statoconia reported by

Magruder (30-40; 1935b) is significantly more than the

auimaber shee herein, possibly exceeding the level of

intra-specific variation and, thus may be an informative

phylogenetic character.

Several significant discrepancies among previous

descriptions of pleurocerid anatomy are now resolved.

Wood. ard ( 1934, 1935) observed a so-called “cytophore

organ” at the base of the mantle cavity, apparently

content with the sperm duct. Similarly, Dazo (1965)

reported the presence of a cytophore organ in males of

both Plewrocera acuta and Elimia livescens. Based on the

present analysis and Woodard’s description of the

internal structure and position of this organ, it is clear

that Woodard misidentified the kidney bladder as a part

of the reproductive tract. However, it is not clear why

the cytophore organ was described as lacking in females.

The intimate connection between the bladder and

proximal pallial gonoduct was correctly depicted in

Hua by Prozorova (1990).

Dazo (1965), as well as

(Woodard, 1934: Jones and Branson,

other workers

1964), have

several

Page 130

THE NAUTILUS, Vol. 119, No. 4

Table 2.

Summary of morphological differences between Pleurocera acuta and Elimia livescens compared to other limnic

gastropods cli issified in the Pleuroceridae and Me lanopsidae. Details from Itagaki, 1960; Bilgin, 1973; Houbrick, 1988; Nakano and

Nishiwaki, 1989; Glaubrecht, 1996; Strong and Glaubrecht, unpubl. data.

Pleuraceura acuta

Elimia livescens

juga

Semisulcospira Melanopsidae

External Anatomy:

Ovipositor pore Simple, weakly glandular

Simple, weakly

Not applicable Complex, highly glandular

glandular

Alimentary System:

Salivary glands Tubular > Tubular Tubular/branched

Salivary gland position Pass through nerve ring ? Pass through nerve Pass through/anterior to

ring nerve ring

Esophageal gland Absent ? Absent? Present

Digestive gland ducts 2 ? 2 |

Caecum Small ? Small Deep and spiral

Reno-Pericardial System:

Bladder Small, pallial e i Small, pallial

Nervous System:

Dialyneury, Zygoneury Zygoneury if Dialyneury? Zygoneury

Reproductive System:

Seminal vesicle Straight ? Straight Coiled

Pallial oviduct Open Open Closed Open

Seminal receptacle Absent Present Present Present

Reproductive strategy Oviparous Oviparous Viviparous Oviparous

reported the presence of a seminal receptacle, but no

mention of a spermatophore bursa was made. The

present ee has confirmed that the sperm storage

structure in Pleurocera acuta and Elimia livescens is

a bursa paced on the presence of unorientated sperm.

Examination of specimens of E. laqueata (USNM

1081558) confirmed that the structure reported as

a seminal receptacle is indeed a bursa—no_ seminal

receptacle is present; judging from the description of

Jones and Branson (1964), the same holds true for E.

potosiensis. The pallial position of the bladder was also

confirmed in E. laqueata (pers. obs.). It is interesting to

note that the sperm gutter extends farther anteriorly in

E. laqueata than in E. livescens. Additionally, the straight

segment of the albumen gland between the tip of the

bursa and the junction of the renal oviduct is lacking in

E. laqueata. Instead, the albumen gland arcs dors: illy to

join the renal oviduct just behind the tip of the bursa. In

all other respects, the overall structure of the pallial

oviduct in’ E. laqueata is consistent with features

described here, as well as in the close association

between the ovipositor and the junction of the

propodium and mesopodium.

A final discrepancy is the presumed site of spermato-

(1934, 1935)

distal prostate as smooth and the highly folded proximal

phore formation. Woodard described the

portion as the site of spermatophore formation. Jones

and Branson (1964) did not distinguish a spermatophore-

forming region in Elimia potosiensis. In the present

study, it is the highly folded distal region that is inferred

to be the site of sperm itophore formation,

SYSTEMATICS OF PLEUROCERIDAE

As mentioned above, although confusion has long

existed, the distinctiveness of Fie Pachychilidae from

other limnic lineages including the Pleuroceridae has

now been clarified based on morphol ygical and molec-

ular data (e.g. Glaubrecht, 1996, 1999; Lydeard et al.,

2002: Kéhler et al., 2004). However, the paraphyly of

eastern and western North American and Asian pleur-

ads with respect to the Melanopsidae based on

molecular data (Lydeard et al., 2002) remains at issue.

The analysis of Houbrick (1988) did not include

sufficient taxon sampling to adequately assess mono-

phyly of the two fi cas 's, but a sister-group relationship

between the two was supported.

Although an in depth analysis of monophyly and

affinity of the two families is beyond the scope of this

study, several morphological features may be informa-

tive in clarifying these relationships. ‘As noted by

Prozorova (1990) the reproductive anatomy of eastern

North American pleurocerids differs from species in

western North America (fuga) and Asia (Semisulcospira)

in that both Juga and Semisulcospira possess a seminal

receptacle in addition to a spermatophore bursa. The

latter genus has modified the pallial oviduct into a closed

brood pouch (Itagaki, 1960; Nakano and Nishiwaki,

1989: Prozorova, 1990: Rashchepkina, 2000; Prozorova

and Raschepkina, 2001, 2004).

In sddiece to the synapomorphies recovered in the

analysis of Houbrick (weakly developed hypobranchial

ole ind, zygoneury, long left ple ural/sub-esophageal con-

E. E. Strong, 2005

Page 13]

nective), midgut anatomy is broadly congruent in the

two families, dif ffering in several significant respects from

that of the other limmic lineages (e.g. Paludomidae,

Pachychilidae, Thiaridae) (Bilgin, 1973; Kohler and

Glaubrecht, 2001; Strong and Glaubrecht, 2002, 2003,

unpubl. data). They also share similarities in reno-

pericardial (presence of a bladder) and reproductive

anatomy (open pallial gonoducts, presence of a seminal

rece eptacle) the latter, in particular, are undoubtedly

symplesiomorphic. But other aspects of the anatomy are

consistent within each family and clearly differentiate

the two when sufficient information is available. Thus, in

addition to features of the radula and shell, melanopsids

may be distinguished by the presence of an esophageal

gland, salivary glands that lie anterior to the nerve ring

(although variable in the family), a single digestive gland

duct and spiral caecum in the midgut, md a oiled

seminal vesicle. Of course, the extent to which these

features represent shared derived or homoplastic

features remains to be discovered in the context of

a phylogenetic analysis.

CONCLUSIONS

The present study has provided the first detailed

description of the midgut and kidney for any pleurocerid

snail, and_ has elaahed the internal structure and

homologies of the pallial gonoducts of eastern North

American forms. This comparative analysis has confirmed

the high degree of morphological similarity between

Picipocet acuta and Elimia livescens, but has also

revealed a number of differences in detail; the extent to

which these features support monophyly of the genera

remains to be established. The fact that the presence/

absence of a seminal receptacle distinguishes eastern and

western North American/Asian pleurocerids is confirmed.

Pleurocerids and melanopsids are broadly similar in

features of the midgut and share a similar configuration of

the pallial oviduct, but can be distinguished by saatowical

characters of the alimentary (salivary glands, esophageal

gland. digestive gland ducts, caecum) and reproductive

(seminal vesicle) systems. However, comprehensive

anatomical treatments of western North American and

Asian pleurocerids are needed to fully assess the

morphological distinctiveness of the two ‘families. The

clarification of the distribution of these features, within

the context of a phylogenetic analysis, should aid in

refining the monophyly of the Pleuroceridae and their

affinity, to other freshwater lineages.

ACKNOWLEDGMENTS

I thank Charles Lydeard for organizing a gastropod

morphology w vorkshop at University of Alabama, Tusca-

loosa, in September 2003 that inspired this study. I also

thank Rex Hanger (University of Wisconsin-W hitewater)

for assistance with collecting the samples of Pleurocera

acuta and Elimia livescens, Paul Johnson (Alabama

Department of Conservation and Natural Resources) for

supplying comparative material of Elimia laqueata, and

Jonathan Slaght (University of Minnesota) for providing

translations of several Russian texts. I am indebted to

Marilyn Schotte (USNM) for inking the anatomical

drawings. Arthur Bogan (North Carolina State Museum

of Natural Sciences), Philippe Bouchet (Muséum

national d'Histoire naturelle, Paris) and John Wise

(College of Charleston) provided valuable comments

that improved the quality of the manuscript.

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THE NAUTILUS 119(4):133-148, 2005

Page 133

New Late Cretaceous (Santonian and Campanian) gastropods

from California and Baja California, Mexico

Richard L. Squires

Department of Geological Sciences

California State University

Northridge, CA 91330-5266 USA

[email protected]

County

LouElla R. Saul

Invertebrate Paleontology Section

Natural History Museum of Los Angeles

900 Exposition Boulevard

Los Angeles, CA 90007 USA

lousaul@earthlink. net

ABSTRACT

Three new genera and six new species of shallow-marine Late

Cretaceous gastropods are reported from various formations in

California and from one formation in Baja California, Mexico.

7 jeanae new species, of early Campanian age, is the

earliest known species of this trochid genus. Nerita (sub-

genus?) orovillensis new species is the second known Early

Campanian neritid from California. The cerithioid Bullamir-

ifica new genus is represented by three species: Bullamirifica

verruca new Lae of Coniacian age; Bullamirifica elegans

new species of early Campanian age; and Bullamirifica ainiktos

(Dailey and Popenoe, 1966) of middle to late ( Campanian age.

The latter species has the most widespread distribution, with

occurrences in southern California and northern Baja Cali-

fornia. Minytropis melilota new genus and species of Santonian

age, and Paxitropis dicriota new genus and species of Late

Santonian to early Campanian age are high-spired trichotro-

pids. As presently known, Bullamirific. a, Minytropis, and

Paxitropis were endemic to the study area.

INTRODUCTION

This study is based largely on specimens collected by

Eric Géhre of Oroville, California. Over the years, he

has amassed a sizeable collection of shallow-marine

mollusks from the lower Campanian Pentz Road

member of the Chico Formation near Pentz, Butte

County, northern California (Figure 1). His collection

has yielded several new species of gastropods, and some

of these were described by Groves (2004) and Squires

and Saul (2004). In part, this present study concerns

three additional new species and a new genus of

gastropods found in his collection. They are the trochid

Tegula jeanae new species, the neritid Nerita (sub-

genus?) orovillensis new species, and the cerithioid

Bullamirifica elegans new genus and species.

Inspection of the literature, as well as examination of

the collections at the Natural History Museum of Los

Angeles County, allowed us to incorporate two additional

species into Bullamirifica. These are Bullamirifica

verruca new genus and species from the Coniacian

Member IV of the Redding Formation in the Oak Run

area, northern California, and Bullamirifica anikitos

Dailey and Popenoe (1966) new combination from the

middle ¢ Campanian Pigeon Point Formation southwest of

San Francisco, northern California: the middle upper

Campanian Punta Baja Formation, Baja California,

Mexico; and the upper Campanian n Jak uma Formation,

southern California (Figure 1). “Cimolithium miya-

koense” (Nagao, 1934) and “Vicarya (Shoshiroia) yabei”

Kamada, 1960, reported by Perrilliat-Montoya (1968)

from Baja California, Mexico (see Figure 1, formation 6),

are judged by us to be synonyms of Bullamé ‘ifica ainiktos.

Also included in this present study are new tricho-

tropid gastropods found in the collections at the Natural

History Museum of Los Angeles County. They are

Minytropis melilota new genus and species from the

Santonian part of the Redding and Chico formations of

northern California, and Paxitropis dicriota, new genus

and species from the of upper Santonian part of the

Redding Formation, northern California; the lower

Campanian part of the Chico Formation; and the lower

Campanian part of the upper Holz Member of the Ladd

Formation, southern California (Figure 1).

The geologic age of each new species described in this

paper is shown in Figure 2. The entire interval of time

that encompasses all these species is Coniacian to late

Campanian, or about 19 million years. The new species

are locally common, except for Tegula jeanae, Nerita

(subge mus?) orovillensis, and B. verruca.

The classification system used here generally follows

that of Hickman and McLean (1990) for the tegulines,

Ponder (1988) for the trichotropids, and Ponder and

Warén (1988) for the other taxa.

Study localities are listed in Appendix 1. Abbreviations

used in the text are: CAS: California Academy of Sciences,

Page 134

THE NAUTILUS, Vol. 119, No. 4

1-Redding Formation

2-Chico Formation

3-Pigeon Point

Formation

4-Jalama Formation

5-Ladd Formation

6-Punta Baja

Formation

Figure 1. Location of formations bearing the new taxa.

San Francisco; IGM: México Museo del Paleontologia del

Instituto de Geologa; LACMIP: Natural History Museum

of Los Angeles County, Invertebrate Paleontology

Section; UCLA: University of California, Los Angeles

(collections now housed at LACMIP); UCMP: University

of California Museum of Paleontology (Berkeley); USGS:

United States Geological Survey.

STRATIGRAPHY

Except for the Punta Baja Formation, which is discussed

below, the ages and depositional environments of all the

formations and members containing the new taxa

discussed in this paper can be found in the following

papers: Member IV of the Redding Formation, Squires

and Saul (2003a); Musty Buck Member of the Chico

Formation, Saul and Squires (2003); Pentz Road

member (informal) of the Chico Formation, Squires

AGE (m.y.) 80 75

UPPER CRETACEOUS

Sant-

onian

Coniacian Campanian

lower; middie j; upper

polarity

chrons 3 +— C33 ——+——4# (32

eanae

Tegula f

orovillensis

[227777]

Nerita

—— SSS

verruca !

see elegans |

f !

Bullamirifica :

eins

—————_| > = 1. > oh

Minytropis melilota 1

Paxitropis dicriota

Figure 2. Chronostratigraphic positions of the new taxa.

Ages of stage boundaries and magnetostratigraphy from

Gradstein et al. (2004, fig. 19.1).

and Saul (1997); Ten Mile Member of the Chico

Formation, Squires and Saul (2003b); upper Holz Shale

of the Ladd Formation, Squires and Saul (2001); Pigeon

Point Formation, Elder and Saul (1993) and Squires and

Saul (2003b); and Jalama Formation, Squires and Saul

(2003b). The locales of these formations are shown in

Figure 1. Stratigraphic information mentioned below

concerns additional pertinent biostratigraphic details.

The age of the Jalama Formation used here is slightly

younger than used in our previous papers because we

had to adjust its chronostratigraphic position based on

the latest published ( Gradstein et al., 2004) absolute-

time and global-paleomagnetic data correlations.

Punta Baja FORMATION

Perrilliat- Montoya (1968) reported specimens of gastro-

pods, herein assigned to Bullamirifica ainiktos, from the

“Rosario Formation” at Punta Baja, near El Rosario,

northern Baja California, Mexico. The 5-140 m_ thick

Punta Baja Formation (Figure 1) overlies fluvial deposits

of the La Bocana Roja Formation, and the angular

unconformity between these two formations is canyon-

shaped (Boe shIke and Abbott, 1986). This canyon is filled

with conglomerate, sandstone, and siltstone reported by

Kilmer (1963) to have been deposited in shallow-marine

depths not exceeding 60 me is rs. Boehlke and Abbott

(1986) have a differing viewpoint and reported that the

ie posits represent buaebicite »s that accumulated in bathyal

depths. They reported, furthermore, that shallow-marine

mollusks are common, but Kilmer’s collection at UCMP

does not contain very many specimens. The Punta Baja

Formation is unconform: ably overlain by terrestrial

deposits of the La Escarpa Member of the El Gallo

R. L. Squires and L. R. Saul, 2005

Formation, which, in turn, is overlain by the Rosario

Formation.

Based on molluscan fossils collected by F. H. Kilmer,

Saul (1983: 21-22. fig. 9) reported the ammonite

Metaplacenticeras cf. pacificum (Smith, 1900) and the

gastropod Turritella chicoensis pescaderoensis Arnold,

1908, from the siltstone in the Punta Baja Formation.

Although these two mollusks were reported by Saul

(1983: 65-66) to be of late Campanian age, more recent

biostratigraphic studies (Elder and Saul, 1996: fig. 1)

depicted both of these taxa as ranging in age from late

middle Campanian to earliest late Campanian. Adjust-

ments for the most recently published (Gradstein et al.,

2004) absolute-time and global-paleomagnetic data

correlations place these ammonite and turritellid zones

in the middle late Campanian. Recent examination by

the junior author of additional Punta Baja Formation

mollusks revealed three specimens of the bivalve Calva.

The best preserved specimen is from LACMIP loc.

12582 and is Calva (Egelicalva) crassa Saul and

Popenoe, 1992, whose geologic range is early late

Campanian to early Maastrichtian elsewhere on the

Pacific slope of North America (Saul and Popenoe,

1992). The other two Calva je aie are worn and

broken, from UCMP loc. B-3388. These two specimens

are similar to Calva (Calva) peninsularis (Anderson and

Hanna, 1935), whose geologic range is latest Campanian

to early Maastrichtian elsewhere on the Pacific slope of

North America (Saul and Popenoe, 1992).

Boehlke and Abbott (1986: fig.

the Punta Baja Formation to the early Campanian based

entirely on calcareous nannofossils. They also reported

that the benthic foraminifera in this formation corre-

spond to the F2-lower E foraminifera zones of Goudkoff

(1945), but they did not rely on the foraminifera for their

age call. Almgren (1986: table 2) reported that the F2-

lower E foraminifera zones are essentially correlative to

the early Campanian to late Campanian. It is important

to mention that the Alcalde Shale in the Co: ilinga area

along the west side of the San Joaquin Valley, central

California, is correlative to the E zone (Almgren, 1986:

ee 3). As depicted in Saul (1983: fig. 10), the Alcalde

Shale contains Metaplacenticeras ct. M. pacificum, and

Almgren (1986) assigned the Alcalde Shale to the early

late Campanian.

In summary, the Metaplacenticeras, Turritella, Calva,

and benthic foraminifera data strongly support a middle

late Campanian age for the Punta Baja Formation. The

calcareous nannofossils, however, support an early

Campanian age. It seems probable that the older

calcareous nannofossils are reworked, and this would

be consistent with the depositional environment of the

formation.

SYSTEMATIC PALEONTOLOGY

Superfamily Trochoidea Rafinesque, 1815

Family Trochidae Rafinesque, 1515

4) assigned the age of

Subfamily Tegulinae Kuroda, Habe and Oyama, 197]

Genus Tegula Lesson, 1835

Type Species: Tegula elegans Lesson, 1835, by

monotypy; Recent, west coast of Central America to

the Gulf of California, Mexico.

Although Wenz (1938), Keen (1960), and

Davies (1971) reported the geologic range of Tegula to

be Miocene to Recent, Bandel and Stinnesbeck (2000)

reported a species of Tegula of Late Cretaceous

(Maastrichtian) age from central Chile. Kiel and Bandel

(2001) reported a tentatively identified Tegula from

upper Campanian strata in northern Spain. The early

Campanian new species described below represents the

confirmed earliest record we know of for Tegula. For the

Pacific slope of North America, the previous earliest

record of Tegula was given by Addicott (1973: 17, pl. 8,

figs. 2, 4), who reported it from the Wygal Sandstone

Member of the Temblor Formation, southwestern

margin of the San Joaquin Valley, Kern County, central

California. Squires (2003: table 2.1, fig. 2.1) placed this

member in the lower Oligocene Matlockian Stage.

Discussion:

Tegula jeanae new species

(Figures 3-5)

Diagnosis: A Tegula with low to moderate spire.

Whorls convex, smooth, and bearing one spiral groove

on posterior third of last whorl. Anomphalous. Last

whorl with raised lip along basal edge, base sunken

between this lip and columellar lip, which bears at least

one denticle and one much smaller denticle adapically.

Description: Shell medium (up to 16.6 mm height

and 21.7 mm diameter, same specimen). Turbiniform

with spire low to moderately elevated. Protoconch

unknown. Teleoconch consisting of three whorls. Suture

impressed, slightly channeled. Whorls convex, some-

times slightly concave short distance anterior of suture;

blunt angulation anterior of medial part of last whorl.

Whorls smooth: spiral groove eee of medial part

of last whorl. Aperture oblique, peristome discontinu-

ous. Anomphalous, umbilical area covered by broad

callus. Base wide and smooth, peripheral (abaxial)

margin coincident with raised lip along edge of last

whorl; area depressed between this lip and columella.

Outer lip strongly prosocline. Columellar lip with at least

one oblique denticle; much weaker second denticle

sometimes present immediately posterior of main

denticle. Growth lines strongly prosocline, forming wide

bands.

Holotype: LACMIP 13322,

22 mm in diameter.

LACMIP 13323.

18.7 mm in height,

Paratype:

Type Locality: LACMIP loc. 24337

Geologic Age: Early Campanian.

Page 136

THE NAUTILUS, Vol. 119, No. 4

Figures 3-8.

11.6 mm, diameter 16.8 mm, 7-8. Paratype LACMIP 13325, total shell height 10 mm, diameter 13.7 mm.

Apical view.

Distribution: Chico Formation, Pentz Road member

(informal), near Pentz, Butte County, northern Califor-

nia.

Etymology: Named for Jean Géhre, mother of Eric

Gohre, who collected and donated the type material to

LACMIP.

Discussion: This new species is known from two

specimens, both showing good preservation. The new

species is remarkably aay to Tegula (Chlorostoma)

funebralis (Adams, 1855), from Pliocene and Pleistocene

strata of southern California (Grant and Gale, 1931) and

from the Recent of Vancouver Island, British Columbia,

to central Baja California, es (McLean, 1978). The

new species differs from T. ) funebralis by having

a smooth shell rather than oe ornamented by weak

nee ribs. The similarity between the new species and

T. (C.) funebralis is even stronger if the specimens of the

a are worn.

Tegula ovallei (Philippi, 1887: - fig. 4; Bandel

and Stinnesbeck, 2000: 761, pl. 1, the only other

positively identified Cretaceous Te mule chi it we know of,

is from Maastrichtian strata in central Chile. The new

species differs from T. ovallei by having a smooth shell

rather than being ornamented by granulated spiral

ridges

Kiel and Bandel (2001; 139, pl. 1, fig.

a tentatively identified Tegula? simplex (Quintero and

Revilla, 1966: 49 pl. §, fig

1) reported

from upper Campanian

New tegulid and neritid gastropods. Specimens coated with ammonium chloride. 3

LACMIP loc. 24337. 3. Paratype LACMIP 13323, apertural view, height 13.9 mm, diameter 17.9 mm, 4-5. Holotype 13322, height

18.7 mm, diameter 22 mm. 6-7. Nerita (subgenus?) orovillensis new species. 6. Holotype LACMIP 13324, a view, height

-5. Tegula jeanae new species,

7. Abapertural view. 8.

strata in northern Spain. The new species differs greatly

from T.? simplex by having a less elevated spire, smooth

shell, blunt Aiea than a sharp angulation anterior of the

medial part of the last whorl, hioad callus covering the

umbilical region, wider aperture, very much stronger

denticles on the columella, raised lip along the basal

edge of the last whorl, and sunken base between this

raised lip and the columellar lip.

Family Neritidae Rafinesque, 1S15

Genus Nerita Linnaeus, 1755

Type Species: Nerita peloronta Linnaeus, 1758, by

subsequent designation (Montfort, 1810); Recent, south

Florida, West facies: and Bermuda.

Discussion: Nerita sensu lato ranges from een

Cretaceous (Hauterivian), and the earliest record i

from the Ono Member of the Budden Canyon

Formation, Trinity County, northern California (Saul

and Squires, 1997). The new species described below

represents the first record of an early Campanian Nerita

from the study area.

Subgenus?

Nerita (subgenus?) orovillensis new species

(Figures 6—S)

Diagnosis: A Nerita with approximate ‘ly 18 to 19

beaded spiral ribs. Columellar lip with four or five

obscure teeth.

R. L. Squires and L. R. Saul, 2005

Page 137

Description: Shell medium small (up to 11.6 mm in

height and 16.3 mm in diameter, same specimen),

broader than high, globose. Last whorl rapidly exp: roe

ing. Protoconch unknown. Teleoconch consisting of 2

to 2.75 whorls. Uppermost spire very low. ee

obscure. Earliest 1.5 teleoconch whorls apparently

smooth, rest of teleoconch covered with approximate ly

1S narrow spiral ribs bearing small beads; interspaces

between ribs approximately as wide as interspaces.

Beads on ribs becoming smaller and slightly elongate

on base of last whorl, especially in pariet ‘all region. Spiral

rib adjacent to suture can be slightly stronger say other

ribs. Aperture large, nearly circular. Outer lip flared,

interior smooth. Columellar lip with five somewhat

obscure teeth, most posterior tooth widest and longest.

Deck area broad, sloping, and sharply demarcated from

base of last whorl. Growth lines prosocline.

Holotype: LACMIP 13324,

16.8 mm in diameter.

Paratype: LACMIP 13325.

LACMIP loc. 24337

11.6 mm_ in height,

Type Locality:

Geologic Age: Early Campanian.

Distribution: Chico Formation, Pentz Road member

(informal), near Pentz, Butte County, northern Califor-

nia.

Etymology: Named for Oroville, California.

Discussion: The new species is based on two speci-

mens. The external surfaces are moderately well pre-

served, but the columellar lip and especially the deck

area are poorly preserved.

The new species is remarkably similar to Nerita

(Theliostyla) crooki Clark (1938: 700, pl. 4, figs. 1, 2)

from the Markley Formation east of San Francisco,

Solano County, northern California. Squires (2003: table

2.1. fig. 2.1) assigned this formation to the middle

Eocene ( ee Stage”). The new species differs from N.

(T.) crooki by having fewer and wider teeth on the

columellar lip, fewer ribs on the last whorl with relativ ely

wider interspaces, and ribs near the middle of the last

whorl not noticeably broader than adjacent ribs.

The new species is also very similar to Nerita

umzambiensis Woods (1906: 311, pl. 37, figs. 14-15;

Bandel and Kiel, 2003: 51-52, pl. 1, figs. 4-5) from

the Santonian/Campanian Umzamba Formation in

southeastern South Africa. The new species differs

from N. umzambiensis by having fewer teeth on the

columella lip. ribs on the base of the last whorl, and

a deck area sharply demarcated from the base of the last

whorl.

The new species somewhat resembles Nerita (The-

liostyla?) kennedyi Squires and Saul (2002: 185-187,

figs. 31-34) from the upper lower to lower middle

Eocene (“Domengine Stage”) Santiago Formation,

northern San Diego County, southern California. The

new species differs from N. (7.2) kennedyi by having

beads that are not elongate, wider interspaces between

the ribs, and fewer, stronger, and wider teeth on the

columellar lip.

The only other early Campanian neritid known from

the Pacific slope of North America is Neritina (Dostia)

cuneata (Gabb, 1864: 137, pl. 21, fig. 97) from lower

Campanian strata at Tuscan Springs on Little Salt Creek,

Tehama County, northern California. Gabb’s species

might also be present in 1) upper Campanian and/or

lower Maastrichtian strata in the Pozo area, San Luis

Obispo County (Vedder, 1977) and 2) Maastrichtian

strata along the western edge of the San Joaquin Valley,

California ‘Chloods and Saul, 1986). The new species is

vastly different from Neritina (Dostia) and does not have

its patelliform shape nor its distinctive collabral sculp-

ture.

Superfamily Cerithioidea Férussac, 1819

Family Indeterminate

Discussion: The new genus described below is most

likely a cerithioid, on the basis of its sigmoidal growth

lines, high spire, sculpture, short siphonal canal (slightly

twisted), smooth columella, and smooth interior of the

outer lip. Some specimens of the new genus have

a narrow spire, like that found in cerithioids, but other

specimens of the new genus have a buccinid-like shell.

The strongly sigmoidal “growth lines of the new genus,

however, are ankike that found on buccinid shells. It is

possible that the new genus belongs to a new cerithioid

family.

Genus Bullamirifica new genus

Type Species: Bullamirifica elegans, new species;

Early Campanian, Pentz area, Butte County, northern

California.

Description: Shell medium (up to 83 mm height and

37 mm diameter, same sprcmnen)s ), fusiform to turreted.

Height to diameter ratio 2 to 2.7. Spire high, comprising

41 to 595% of total shell height. Pleural angle 33 to 42°.

Protoconch unknown. Teleacsadh whorls six to eight.

Spire whorls with shoulder angulate; last whorl with

periphery angulate. Ramp short to moderately long,

concave to rarely straight-sloped. Suture slightly un-

dulatory, weakly impressed. Collabral sculpture consist-

ing of many narrow ribs, closely to moderately widely

spaced; interspaces smooth. Collabral ribs slightly

opisthocline to opisthocyrt, usually extending fom

suture to suture. Collabral ribs present on base or

obsolete; if present, swollen and elongate. Spiral

sculpture consisting of several spiral ribs with variable

width and spacing, especially on last whorl. Spire whorls

with strongest spiral rib on whorl shoulder, several weak

or moderately strong spiral ribs occ: asionally near

anterior suture, and suture coincident with weak spiral

rib either bearing weak nodes or without nodes. Last

whorl sculpture with three to four widely spaced, strong

spiral ribs on periphery and one or two weaker spiral ribs

Page 138

THE NAUTILUS, Vol. 119, No. 4

or several spiral riblets on base. Intersections of collabral

and spiral ribs producing many nodes or strongly

projecting tubercles, either rounded (knob-like) to

spinose or narrowly elongate. Nodes and knobs most

pronounced on whorl shoulder, especially on last whorl.

Intersections also strong on anterior portion of last whorl

periphery. Aperture short but moderately wide, comma-

shaped; small, narrow arch (canal-like) present where

outer lip meets most posterior part of aperture.

Columellar lip smooth. Siphonal canal short and spout-

like or well developed, moderately short, and can be

twisted to left. Outer lip thin, markedly sinuous, interior

smooth. Growth lines sigmoidal between suture and

shoulder; antispiral sinus coincident with tuberculate

spiral rib.

Geologic Age: Coniacian to early late Campanian.

Etymology: Combination of the Latin bulla, mean-

ing knob, and the Latin mirifus, meaning to cause

wonder.

Discussion: Three species can be herein assigned to

this new genus. Two of these species, Bullamirifica

verruca and Bullamirifica elegans, are based on entirely

new material. The third species was originally tentativ ely

assigned by Dailey and Popenoe (1966) to Pse udoglau-

conia Douvillé, 1921. Dailey and Pope noe (1966) stated

that this particular species belongs in a new genus, but

they withheld their description mati better specimens

were obtained. Although representatives of the new

genus have sigmoidal growth lines (see Wenz, 1940: 764,

fig. 2214) similar to that of Psewdoglauconia, Bullamir-

ifica differs considerably from Pseudoglauconia by

having bucciniform rather than a_tapered/conical

shape, angulate rather than flat-sided whorls, and

tubercles on the sides of the whorls instead of only near

the suture, and the suture between the pe ultimate and

last whorl is not extremely deep and widely sunken.

Unfortunately, the aperture of Pseudoglauconia is not

known and none of the apertures on the available

specimens of Bullamirifica is complete. In spite of the

absence of knowledge about the aperture of Pseudo-

glauconia, Wenz ( (1940) believed this gastropod genus to

belong in the Cerithiidae.

The shape of the growth lines in Bullamirifica is

similar to that of Batillaria echinoides clavatulata

(Lamarck, 1804) from the middle Eocene (Lutetian) of

the Paris Basin, France. The siphonal canal of this

Eocene species is longer than normally found in

Batillaria Benson, 1842. Ponder and Warén (1988) and

Houbrick (1988) placed Batillaria in superfamily Cer-

ithioidea, family Batillariidae Thiele, 1929. The similarity

in growth-line shape between Bullamirifica and Batil-

laria suggests to us that the new genus might be

a cerithioid. The high turreted spire of Batillaria,

however, is quite unlike the lower, more paucispiral

spire of Bullamirifica.

Bullamirifica has the growth-line shape, shell shape,

sculpture, and twisted siphonal canal similar to that of

Pseudorapa Holzapfel, 1888, a monotypic genus from

the Vaals Greensand in the Netherlands and Germany.

The age of these strata was determined to be early

Campanian by Albers (1976). Wenz (1941: 1083,

fig. 3079) illustrated Pseudorapa. Bullamirifica differs

from it by having a less twisted siphonal canal and

more variable sculpture, including the possibilities of

having spiral ribs and < relatively narrow shell. In

addition, Pseudorapa bee an outer lip that is very

crenulate and a spiral band near the base of the last

whorl that produces a tooth-like projection on the outer

lip.

Bullamirifica verruca new species

(Figures 9-12)

Diagnosis: Small Bullamirifica with prominent round-

ed knobs, nine on shoulder of last whorl. Suture

coincident with noded spiral rib. Base of last whorl

without elongate collabral ridges. Siphonal canal short

and straight.

Description: Small (up to 34 mm estimated height

and 16.5mm_ diameter, same specimen), fusiform,

moderately slender. Height to diameter ratio approxi-

mately 2. Spire high, approximately 44% of shell height.

Pleural angle approximately 35 to 37°. Protoconch and

upper spire unknown. Teleoconch whorls approximately

six (estimated). Spire whorls with shoulder angulate, last

whorl with periphery angulate. Ramp short, concave

Suture slightly impressed, ‘possibly undulatory. Collabral

sculpture consisting of many ribs, widely spaced;

interspaces smooth. Collabral ribs slightly opisthocline

and extending from suture to suture. Collabral ribs

mainly prey valent on ramp. Spiral sculpture consisting of

several ribs with variable strength and spacing, especially

on last whorl. Intersections of collabral and spiral ribs

producing many strongly projecting rounded tuberculate

knobs or, less commonly, nodes: knobs and nodes most

prominent on shoulder and usually extending posteriorly

across ramp and become narrow ridges. Spire whorls

with strongest spiral rib on whorl shoulder, very faint

spiral riblets present between whorl shoulder and

anterior suture. Suture coincident with moderately weak

spiral rib bearing small nodes. Penultimate whorl with

spiral rib on shoulder bearing nine, moderately closely

spaced knobs. Last whorl with three spiral ribs on

periphery, strength of ribs progressively decreasing

anteriorly: strong rib on shoulder and bearing nine very

prominent tuberculate knobs: middle rib miodenately

strong and bearing more numerous nodes (about half

sized of those on shoulde sr) that tend to become smaller

and even obsolete adaxially; and most anterior periphery

rib weakest and weak nodes tending to be obsolete

adaxially. Base of last whorl usually with one or two

spiral ribs, both bearing nodes (best developed near

outer lip) and with strength of

spiral ribs decreasing anterior in direction: occasionally,

anterior region of base with only very weak spiral riblets.

or bearing no nodes,

R. L. Squires and L. R. Saul, 2005 Page 139

Figures 9-21. New cerithioid? gastropods. Specimens coated with ammonium chloride, 9-12. Bullamirifica verruca new genus and

species, LACMIP loc. 8133. 9. Paratype LACMIP 13327, apertural view, height 27 mm, diameter 19.2 mm, 10-11. Holotype LACMIP

13326, height 29.7 mm, diameter 17.5 mm. 10. Apertural view. 11. Abapertural view. 12. Paratype LACMIP 13327, basal view,

diameter 18.9 mm. 13-17. Bullamirifica elegans new genus and species. LACMIP loc. 24337. 13-14. Holotype LACMIP 13328,

height 66.9 mm, diameter 34.9 mm. 13. Apertural view. 14. Abapertural view. 15-16. Paratype LACMIP 13329, height 60.1mm

diameter 25.7 mm. 15. Apertural view. 16. Abapertural view. 17. Holotype LACMIP 13328, basal view, diameter 35.7 mm. 18-21.

Bullamirifica ainiktos (Dailey and Popenoe, 1966) new genus. 18-19. Hypotype LACMIP 13330, LACMIP loc. 10691, height 60 mm,

diameter 24.1 mm. 18. Apertural view. 19. Abapertural view. 20. Plasto-holotype LACMIP 40435, LACMIP loc. 24125, right-lateral

view, height 58.2 mm, diameter 29.1 mm. 21. Hyptotype LACMIP 13331, LACMIP loc. 24124, basal view, diameter 16.7 mm

Page 140

THE NAUTILUS, Vol. 119, No. 4

Aperture round with very small posterior “arch.”

Aperture elliptical, columellar lip smooth, outer lip thin;

siphonal canal short and spout-like.

Holotype: LACMIP 13326, incomplete specimen

with two whorls (upper spire missing), 29.7 mm height,

17.8 mm diameter.

Paratype: LACMIP 13327

Type Locality: LACMIP loc. $133.

Geologic Age: Coniacian.

Distribution: Redding Formation, Member IV, Oak

Run area, northern California.

Latin verrucus, wart.

Etymology:

Discussion: This new species is based on three

specimens. It differs from Bullamirifica elegans new

species below by smaller size and presence of rounded

rather than elongate knobs, fewer knobs on shoulder of

last whorl, no elongate collabral ridges on base of last

whorl, a noded spiral rib coincident with the suture, and

a straight siphonal canal. Bullamirifica verruca differs

from Bullamirifica ainiktos by being smaller with

a shorter spire and having rounded and much more

projecting nodes, many fewer nodes on shoulder of last

whorl, much less tendency for elongate collabral ridges

on ramp, and much weaker spiral in between shoulder

and anterior suture.

Bullamirifica verruca is very similar to the cerithioid

Tympanotonus aaa robustus | Dockery

(1993: 47, pl. 7, fig. 1) in the shape of the spire, strong

nodes on the spire, shape of the growth lines next to

the outer lip. Tympanotonus (T.)

Campanian age and from Mississippi, however, has no

siphonal canal.

Bullamirifica elegans new species

(Figures 13-17)

[2] Pseudoglauconia? aft. P. ainiktos Dailey and Pope-

noe. 1, 1993: pl. 2, fig. 11.

Diagnosis: Large Bullamirifica with prominent

opisthocline eollabval ridges on upper spire and base

of whorl. Shoulder of lect: whorl with 11 nodes. Suture

coincident with unnoded weak spiral rib. Siphonal canal

short but well developed and twisted to left.

Description: Medium large (up to 83 mm estimated

height and 37 mm diameter, same specimen), fusiform,

moderately wide, rarely slender. Height to diameter

ratio approximately 29. Spire high, approximate ly 40%

of shell height. Pleural angle approximately 33 to 42°,

rarely approximately 30°. Protoconch and uppermost

spire unknown. Te leoconch whorls approximately eight

(estimated). Whorls with angulate shoulder. Ramp short

and slightly concave. Coll: bral sculpture consisting of

narrow ribs:

many mode rately strong, inte rspaces

smooth. Collabral ribs exte nding from suture to suture.

\ oninehas. which is of

Collabral ribs sigmoidal between posterior suture and

shoulder, opisthocline between shoulder and anterior

suture. Collabral ribs somewhat swollen on base of last

whorl. Spiral sculpture consisting of several ribs with

variable strength and spacing. Intersections of collabral

and spiral aie. producing many swollen, axially elongate

nodes or, in some cases, rounded tuberculate inohie?

nodes and knobs most prominent on shoulder. Spire

whorls with strongest spiral rib on whorl shoulder and

occasionally two Tabderately strong, noded spiral ribs

between whorl shoulder and anterior suture. Suture

coincident with weak spiral riblet. Penultimate whorl

with spiral rib on shoulder bearing 11 widely spaced

nodes. Last whorl with three or fear spiral ribs on

periphery, strongest rib on shoulder and bearing 11

nodes. Spiral lie on periphery of last whorl usually show

strength progressively decreasing anteriorly, occasionally

rib eamediately anterior of shoulder weaker than other

ribs on periphery. Growth lines sigmoidal, with antispiral

sinus coincident with shoulder, Aperture _ elliptical,

columellar lip smooth, outer lip thin; siphonal canal

short but well developed and slightly twisted to left.

Holotype: LACMIP 13325; nearly complete specimen

of five whorls (upper spire missing), 66.9 mm height,

34.9 mm diameter.

Paratype: LACMIP 13329.

Type Locality: LACMIP loc. 24337.

Geologic Age: Early Campanian to possibly middle

Campanian.

Distribution: Early Campanian: Chico Formation,

Pentz Road member (informal), near Pentz, Butte

County, northern California; Possibly middle Campa-

nian: ‘Pigeon Point Formation, San Mateo County,

northern Cali fornia.

Etymology: Latin elegans,

choice.

meaning very fine or

Discussion: The new species is based on six speci-

mens from the Pentz area, and preservation is moder-

ately good on all of them. Nearly all the specimens are

moderately wide, but a few are somewhat slender, as the

specimen illustrated in Figures 15-16.

There might be one specimen of the new species from

the Pigeon Point Formation. This museum specimen,

which cannot be located, was figured by Elder and Saul

aie pl. 2, fig. 11). It was identif ied by them as

Pse sudoglauconia? sp. aff. P. ainiktos Dailey and Pope-

noe, aa it has a profile somewhat more like Bullamir-

ifica elegans and basal nodes like B. elegans. As will be

discussed later, there are specimens of B. ainiktos from

the Pigeon Point Formation. It seems likely that some of

the Bullamirifica specimens from the Pigeon Point

Formation represent transitional forms between B.

elegans and B. ainiktos.

Bullamirifica elegans differs from Bullamirifica ver-

ruca in having larger size, prominent collabral ribs

R. L. Squires and L. R. Saul, 2005

extending from suture to suture on the spire whorls,

swollen collabral ribs on the neck, more nodes on

shoulder of the penultimate and last whorls, and

siphonal canal longer and slightly twisted to the left. In

addition, B. elegans aac has strong unnoded

spiral ribs between suture and shoulder.

Bullamirifica elegans difters from Bullamirifica ainik-

tos by usually having a wider pleural angle, usually a less

elevated spire, wider and stronger nodes on shoulder,

fewer nodes on shoulder of last whorl, and base with

swollen collabral ribs instead of fine spiral ribs.

Bullamirifica elegans also has variability in morphology,

whereas B. ainiktos does not.

Bullamirifica elegans is very similar to the cerithioid

Fide cegia hagas (Tympanotonus) binodosus Dockery

(1993: 47, pl. 7, fig. 2) in the shape of the spire, strong

a »s on the spire, and shape the growth ae next to

the outer lip. Tympanotonus (T.) binodosus of Campa-

nian age and from Mississippi, lie has no siphonal

canal.

Bullamirifica ainiktos (Dailey and Popenoe, 1966) new

combination

(Figures 1S—25)

Pseudoglauconia? ainiktos Dailey and Popenoe, 1966:

21-22, pl. 6, figs. 3, 5, 6.

Pseudoglauconia? aff. P. ainiktos Dailey and Popenoe.—

Elder and Saul, 1993: pl. 2, fig. 1G:

Cimolithium miyakoense (Nagao, 1934). — Perrilliat-

Montoya, 1968: 20, pl. 4, fig. 2.

Vicarya (Shoshiroia) yabei Kamada, 1960.—

Montoya, 1968: 21, pl. 6, fig. 1

Perrilliat-

Diagnosis: Medium-size Bullamirifica with usually

device turreted whorls. Ramp on upper spire short,

with narrow collabral ribs crossed by weaker spiral ribs;

ramp on later whorls much longer and smooth and

concave. Spire whorls concave on posterior half but

angulate medially, with moderately strong tubercles.

Shoulder of last whorl with 14 nodes. Base of last whorl

with many, closely spaced spiral riblets. Siphonal canal

short and possibly sly straight.

Description: Medium, up to 67.5 mm height estimat-

ed and 25.4 mm diameter, same specimen. Shell usually

slender, turreted, occasionally wide, fusiform. Height to

diameter ratio approximately 2 2.7. Spire high, approxi-

mately 55% of total shell height. Pleural angle approx-

imately 33. to 37°. Protoconch unknown. Teleocouch

whorls approximately eight (estimated). Spire whorls

with shoulder angulate. Ramp usually short and concave,

occasionally long and straight-slope od. Suture obscured.

Collabral sculpture consisting of many spiral ribs closely

spaced and narrow. Coll bral ribs opisthocyrt, most

prominent on ramp. Spiral sculpture consisting of

several ribs, moderately weak and_ closely spaced.

Intersections of collabral and spiral ribs producing many

nodes, most prominent and somewhat spinose on

shoulder. Spire whorls with strongest spiral rib on whorl

shoulder and five moderately strong, non-noded spiral

ribs between shoulder and anterior suture; most anterior

one of these ribs coincident with suture. Penultimate

and last whorls with approximately 14 nodes on

shoulder. Last whorl with three spiral ribs on periphery,

progressively weaker anteriorly, and bearing con »5, Base

of last whorl be caring fine spiral riblets. Siphonal canal

short and twisted to left. Columellar lip smooth. Outer

lip sinuous; interior smooth.

Holotype: LACMIP 40435, incomplete specimen (tip

of spire and siphonal canal missing), 39 mm_ height,

29 mm diameter.

Type Locality: —LACMIP loc. 4125

Geologic Age: Middle Campanian to Late Campa-

nian.

Distribution: Middle Campanian: Pigeon Point For-

mation, near Pigeon Point, San Mateo County, northern

California. Middle late Campanian: Punta Baja Forma-

tion, ee sst of El Rosario, Baja California, Mexico.

Late Campanian: Jalama Formation, Santa Barbara

County, southern California.

Discussion: Mae fa of the new species is based

on 25 specimens: 1S from the Jalama Formation, five

from the Pigeon Point Formation, and two from the

Punta Baja Formation. The Jalama specimens have poor

to moderately good preservation, although the apices are

broken off and the siphonal canal missing. The Pigeon

Point specimens also have poor to uadley rately good

preservation, but the matrix is very hard to remove. “The

Punta Baja specimens have poor preservation.

The Pigeon Point Formation specimen illustrated by

Elder and Saul (1993: pl. 2, fig. 10) has a whorl profile

like that found on B. anita: but it has wider pleural

angle, similar to that found on B. elegans. Unfortunately

ihe specimen is missing. As mentioned earlier, it seems

likely that some of the Bullamirifica specimens from the

Pigeon Point Formation represent transitional forms

between B. elegans and B. ainiktos.

Bullamirifica ainiktos differs from Bullamirifica ver-

ruca by larger size and having less rounded and much

lower nodes more nodes on shoulder of last whorl,

much more tendency for elongate collabral ridges on

ramp, and much stronger spiral ribs between shoulder

and anterior suture.

Bullamirifica ainiktos differs from Bullamirifica ele-

gans by smaller size, having a narrower spire, usually

a more elevated spire, narrower and weaker nodes on

shoulder, more nodes on shoulder of last whorl, and base

with fine ribs instead of swollen collabral ribs.

Perrilliat-Montoya (1968) identified one of the Punta

Baja Formation specimens (Figures 23-25 herein) of B.

ainiktos as Vicarya (Shoshiroa) yabei and identified the

other specimen as Cimolithium miyakoense. With Wenz

(1940: 740-741, fig. 2145) as a basis for comparison, we

consider that Bullamirifica ainiktos differs considerably

from the former gastropod by having a non-conical she ll,

Page 142 THE NAUTILUS, Vol. 119, No. 4

Figures 22-34. New cerithioid? and trichotropid gastropods. Specimens coated with ammonium chloride, unless otherwise noted.

22-25. Bullamirifica ainiktos (Dailey and Popenoe, 1966) new genus. 22. Hypotype LACMIP 13332, USGS loc. M-SG601, height

35.5 mm, diameter 20 mm, 23-25. Plasto-hypotype IGM 1325, height S80 mm, diameter 36.9 mm. 23. Apertural view. 24.

Abapertural view. 25. Basal view. 26-29. Minytropis melilota new genus and species. 26. Paratype LACMIP 13334, LACMIP loc.

1077, apertural view, height 7 mm, diameter 2 mm, 27-29. Holotype LACMIP 13333, LACMIP loc. 10786, abapertural view,

height 62 mm, diameter 2.2 mm. 27. Abapertural view. 28. Oblique apertural view of tip; arrow indicates where protoconch ends

and teleoconch starts. 29. Protoconch, height 0.5 mm, diameter 0.8 mm; arrow indicates where protoconch ends and teleoconch

3336

starts; SEM photomicrograph (uncoated). 30-34. Paxitropis dicriota new genus and species. 30-31. Paratype LACMIP 13336,

LACMIP loc. 23643, height 12.7 mm, diameter 6.8 mm. 30. Apertural view. 31. Abapertural view. 32. Holotype LACMIP 13335,

LACMIP loc. 23639, apertural view, height 14.7 mm, diameter 6.6 mm. 33. Paratype LACMIP 13337, LACMIP loc, 23642,

apertural view, height 10.1 mm, diameter 4.4 mm. 34. Paratype LACMIP 13338, LACMIP loc, 24217, abapertural view, height

11 mm, diameter 4.3 mm

R. L. Squires and L. R. Saul, 2005

Page 143

more rows of nodes, and absence of a notch on the outer

lip near the suture and not having a thick, extensive

callus that covers part of the noded sculpture near the

suture on the last whorl. Using Kase (1954: 135-136, pl.

21, figs. 1-9; fig. 20) as a basis for comparison, we

consider that Bullamirific a ainiktos differs considerably

from the latter gastropod by having a much nae spire,

more strongly sinuous growth lines, and a well de-

veloped, twisted siphon: iT canal.

Superfamily Capuloidea Fleming, 1522

Family C apulidi ie Fleming, 1822

?Subtamily Trichotropinae Gray, 1850

Discussion: We include a subfamily name because

high-spired capulids, like those described below, are so

distinctive from cap-shaped capulids. The subfamily

allocation is tentative, pending much-needed taxonomic

work on capulids.

Genus Minytropis new genus

Type Species: = Minytropis melilota new species; Late

Cretaceous, Santonian, northern California.

Description: Shell very small, estimated maximum

height just over 10 mm. Shell thin, narrowly fusiform-

elongate. Protoconch flat-topped and smooth, approxi-

mately two whorls. Teleoconch whorls rounded. Sculp-

ture consisting of several strong spiral ribs with wide

interspaces both ribs and interspaces crossed by well

delineated prosocline | growth lines. Aperture ov al, pro-

duced anteriorly to short, with narrow siphonal canal.

Outer lip thin with no varix or inner denticulations.

Inner lip callused, narrow.

Geologic Age: Early and Late Santonian.

Etymology: Combination of the Greek minys, mean-

ing little, small, or short; and the Latin tropis, meaning

keel.

Discussion: The placement of Minytropis among the

trichotropids is suggested by the strongly prosocline and

well marked growth lines, strong spiral sculpture, and

the short, open siphonal canal. Minytropis resembles

Opposirius Iredale, 1931, which is known only from the

Recent of Australia (Wenz, 1940). Minytropis differs

from Opposirius by having a narrower aperture, a longer

siphonal canal, and a rounded rather than a bladelike

inner lip.

Finlay and Marwick (1937) reported that Opposirius is

more similar to Certhioderma Conrad, 1860, than to any

other trichotropid genus. In comparison to Certhioderma,

Minytropis is similar in having a flat-topped, smooth

protoconch and in having a short siphonal canal, but

Minytropis differs by having a much narrower shell,

longer and much narrower aperture, stronger spiral

sculpture. sculpture not recticulate on adult whorls, and

no hint of an umbilicus. Wenz (1940) reported Cerithio-

derma to have a geologic range of Late Cretaceous to

Oligocene, but Marwick (1965) reported that this genus is

also extant. Dockery (1993) reported Cerithioderma from

Campanian strata in Mississippi. High-spired trichotro-

pids very similar to Cerithioderma are best represented in

New Zealand Tertiary strata (Maxwell, 1992). According

to Maxwell (1992), Trichosirius Finlay, 1926, might be

a junior synonym of Cerithioderma. —

Minytropis melilota new species

(Figures 26-29)

Description: Small, up to 10.2 mm estimated height

and 3.5 mm diameter, same specimen, Shell narrowly

fusiform-elongate. Spire high, 60% (estimated) of shell

height. Pleural angle approximately 22°. Protoconch flat-

topped, with two hors , and smooth (0.5 mm height,

0.8 mm diameter). Teleoconch of approximately seven

to eight subangulate whorls, last whorl nearly one-third

total height. Suture deeply impressed, near ly coincident

with very weak rib. Sculpture nearly cancellate on

earliest teleoconch whorl, but four spiral riblets slightly

stronger than numerous collabral threads. Sculpture on

subsequent whorls progressing into five strong, flat-

topped ribs on middle spire, six ribs on lower spire, and

approximately eight to nine ribs on last whorl; most

posterior spiral rib weak to very weak on all these whorls.

Interspaces of nearly equal width on spire whorls but

becoming twice as wide as ribs on last whorl. On last

whorl, ribs on whorl face equally strong but on base, ribs

become much less prominent and more closely spaced.

Growth lines strongly prosocline, marked by fine

collabral threads strongest on early whorls where they

produce slight beading of spiral ribs. Growth lines less

prominent and closer- spaced on later whorls; growth

lines can be somewhat prominent on base of last whorl.

Aperture oval, produced anteriorly to short, narrow

siphonal canal. Outer lip thin, arcuate, crenulated by

ribs. Columellar lip narrow, weakly callused, and barely

forming a pseudo-umbilical chink between base of whorl

and indistinct fasciole.

Dimensions of Holotype: Nearly complete species

(siphonal canal mostly missing), 6.2 mm height, 2.2 mm

diameter.

Holotype: LACMIP 13333.

Type Locality: LACMIP loc. 10756.

Paratype: LACMIP 13334.

Geologic Age: Early and Late Santonian.

Distribution: Lower Santonian: Redding Formation,

Member V, Clover Creek, Shasta County, northern

California. Upper Santonian: Chico Formation, Musty

Buck Member, Chico Creek, Butte County, northern

California.

Etymology: Latin melitota, meaning sweet clover (in

reference to the occurrence of the new species near

Clover Creek).

Page 144

THE NAUTILUS, Vol. 119, No. 4

Discussion: De scription of this new species is based

on 29 specimens, most of which represented by small

fragments. The better preserved specimens are all from

the Redding Formation, however, none of these speci-

mens is as large as the larger ones from the Chico

Formation localities. Only two specimens, both from

LACMIP loc. 10786, have retained the protoconch. The

protoconch of one of these specimens is illustrated in

Figures 27-29. The other specimen’s protoconch is

mostly embedded in hard matrix. The flat- wlan

protoconch of M. meliota supports the position of this

gastropod among the Trichotropidae.

The new species can be distinguished from the suinlee

looking Opposirius idoneus ined le (1931: 210, pl. 2 29,

fig. 7: Wenz, 1940: fig. 2631), an extant species in

southwestern Australia, by having a narrower shell, one

more rib on the spire, two to four more ribs on the base

of the last whorl, a much longer siphonal canal, an

a rounded columella. Opposirius idoneus is the type

species of Opposirius Iredale, 1931.

Genus Paxitropis new genus

Type Species: Paxitropis dicriota; Late Cretaceous,

late Santonian to early Campanian, northern Califor-

nia.

Description: Shell medium small, narrowly pagodi-

form. Spire whorls with prominent keel medially, Suture

bordered posteriorly by relatively weak spiral rib

becoming keel-like on last whorl and, in concert with

aforementioned keel, producing bicarinate last whorl.

Shell base with several weak spiral riblets. Growth lines

prosocline. Columellar lip callused, abaxial margin well-

delineated. Siphonal canal short but distinct. Anompha-

lous.

Geologic Age: Late Santonian to early Campanian.

Etymology: Combination of the Latin pax, meaning

peace (in reference to the peaceful Pacific coast); and

the Latin tropis, meaning keel.

Discussion: Paxitropis is similar to Trichotropis Bro-

derip and Sowerby, 1829, a genus known (Sohl, 1960)

from the Late Cretaceous to Recent. The modern

distribution of Trichotropis is restricted to cireum-boreal

waters, in both the northern and southern oceans and, as

Sohl (1960) noted, this modern distribution is in sharp

contrast to its Cretaceous distribution in temperate

shallow-water deposits. Paxitropis differs from Tricho-

tropis by having a smaller size, much narrower shell

(including a narrower aperture), a siphonal canal, much

lower vz ariability in sculpture, and being anomphalous.

Paxitropis resembles the trichotropid Icuncula Ire-

2 , 1924, which is known from the Recent of the Indo-

-acific (Wenz, 1940). Paxitropis differs from Icuncula by

‘ wing no hint of an umbilicus, less prominent keels that

are unwavy, one less keel on the last whorl, and no axial

riblets on the ramp. The aperture of Paxitropis is not

entirely known.

The new genus has a whorl profile very similar to the

turrid Austrocarina Laseron, 1954, known only from the

Recent of New South Wales and Victoria, southeastern

Australia and Tasmania. Paxitropis, however, is not

a turrid because it does not have the turrid growth lines,

which tend to be opisthocline anterior to the notch and

across the whorl periphery curving toward prosocline

very near the siphonal area.

Paxitropis dicriota new species

(Figures 30-34)

Description: Shell small, up to height estimated

14.5 mm and 6.6 mm diameter. Shell pagodiform. Spire

high, approximately 60% of shell height. Pleural angle

approximately 30°. Protoconch iulcown, Teleseonth

approximately six strongly angulated whorls, with wide

ramp on middle spire w hors and on last whorl. Sculpture

dominated by prominent spiral rib, medially located on

shoulder and_ keel- -forming on later whorls. Early

teleoconch whorls with two spiral ribs anterior to carina.

On mature whorls, these additional ribs become mere

threads and equivalent in strength to growth lines, thus

producing microscopic caneellate surface, especially on

ramp. Suture impressed, nearly coincident with weakly

noded subsutural spiral rib located immediately posterior

to suture. Subsutural spiral rib relatively weak on spire

whorls but becoming keel-like on last whorl. Last whorl

with two sharp kee Is, one just above middle whorl height

and second one weaker and occasionally minutely noded,

emergent at anterior suture. Shell base ‘with several weak

spiral ribs. Growth lines prosocline, most prominent on

shell base, near outer lip. Aperture lanceolate and

produced anteriorly to short, but distinct siphonal canal.

Outer lip thin, angulate, crenulated by ribs. Columellar

lip callused, abasial margin well-de lineated. Columellar

lip narrow, barely forming pseudo-umbilical chink

between base of whorl and sadistinet fasciole.

Holotype: LACMIP 13335, 14.7 mm height, 6.6 mm

diameter.

Paratypes: LACMIP 13336, 13337, and 13338.

Type Locality: LACMIP loc. 23643.

Geologic Age:

Late Santonian to Early Campanian.

Distribution: Upper Santonian: Redding Formation,

Member area east of Redding, Shasta County,

northern California. Lower C vampanian: Chico Forma-

tion, Ten Mile Member, Chico Creek, Butte County,

northern California; Ladd Formation, upper Holz Shale

Member, Santa Ana Mountains, Orange County, south-

ern California.

Etymology: Combination of the Greek di, meaning

two, and the Greek criota, meaning ringed.

Discussion: Twenty incom ple ste specimens were avi ail-

able, and the six most ae te of these are the basis for

the above description. The only specimen from the

R. L. Squires and L. R. Saul, 2005

Page 145

Redding Formation is geologically the oldest specimen

(LAC MIP loc. 24217). The only specimen from the

upper Holz Shale is from LAC MIP loc. 21536.

The new species is very similar to the Pliocene to

Recent Tric yi bicarinata (Sowerby, 1825; Pitt and

Pitt, 1989: pl. 1, figs. 83-4; Egorov and Alexeyev, 199%:

25, figs. 22-23), which is a Pacific upper boreal Arctic

species, occurring in the waters off northern Japan,

Kamchatka, southern Chukchi Sea, and Queen Char-

lotte Islands in British Columbia (Egorov and Alexeyey,

1998). The new species di fers from T. bicarinata

by having a much narrower last whorl and a much

narrower aperture. These same differences distinguish

the new species from Trichotropis vokesae Pitt and Pitt,

1989, which is extremely similar to T. bicarinata.

Trichotropis vokesae is of Pliocene age and from

northwestern Ecuador, and the new species differs from

T. vokesae in the same ways that it differs from T.

bicarinata.

The new species can be distinguished from the similar

looking Icuncula torcularis (Tenison-Woods — 1879;

Wenz, 1940: fig. 2630), an extant species living off the

coasts of southeastern Australia and eee is having

less projecting and non-wavy keels, one less keel on the

last whorl, and an absence of axial riblets on the ramp.

Icuncula torcularis is the type species of Icuncula

Iredale, 1924.

The new species is remarkably similar to the extant

Austrocarina recta (Hedley, 1903; 1922: 223, fig. 3;

Powell, 1966: 34, pl. 3, fig. 2. 1969: 364, pl. 276; W ‘soa.

1994: 187) but dfers foun A. recta by having a shell

approximately 14 mm in height instead of 6 mm, a more

projecting and usually wider keel on the shoulder of the

adult whorls, several weak spiral ribs on base of last

whorl, several spiral ribs on earliest teleoconch whorls,

straight rather than a curved profile of the base,

columellar lip callused with abaxial margin well-de-

lineated, and no tendency to develop eélishral ribs on

earliest teleoconch whorls.

ACKNOWLEDGMENTS

The authors are especially grateful for the careful and

tireless collecting of Eric Goéhre of Oroville, California.

Without his help’ and willingness to donate the material,

the paper would have been “greatly reduced in its scope.

James H. McLean, Natural History Museum of Los

Angeles County, Malacology Division, provided valuable

comments about the id ienteatioh of the new trochid.

Carmen Perrilliat (IGM) graciously sent us excellent

casts of the Punta Baja Formation material. David Haasl

(UCMP) spent considerable effort unsuccessfully trying

to locate the hypotypes of Bullamirifica ainiktos from ea

Pigeon Point Formation. Jorge Vazquez, California State

University, Northridge, kindly took SEM photomicro-

graphs of the protoconch of Minytropis melilota. Steffen

Kiel (Smithsonian Institution) and an anonymous re-

viewer critiqued the manuscript.

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and Eocene neritid gastropods

gastropods from California.

APPENDIX 1

LOCALITIES CITED

Localities are LACMIP, unless otherwise noted. All

quadrangle maps listed below are U. S. Geological

Survey maps.

10786. Near crest of south slope of divide between Basin

Hollow and Clover creeks, at approximately southeast

corner of the northwest 1/4 of section 33, T. 32 N, R. 2

W. Lower Santonian. Redding Formation, Member \

(lower part). Coll: W. P. Popenoe and D. W. Scharf,

August 8, 1931.

10787. Near crest of north slope of divide between Basin

Hollow and Clover creeks, near northeast corner of

northwest 1/4 of section 33 and not more than 122 m

south of section line, T. 32 N, R. 2 W, Millville

Quadrang] e (15 minute, 1953), Shasta County, northern

California. Lower Santonian. Redding Formation, Mem-

Page 148

ber V (lower part). Coll: W. P. Popenoe and D. W.

Scharf, August 8, 1931.

12582. Turritellas and oysters along beach just south of

La Bocana Roja Formation outcrop, approximately 2 km

north of south tip on west side of Punta Baja, Mexico.

Middle upper Campanian. Punta Baja Formation. Coll:

L. R. Saul, October 12, 1990.

21536. Corona Quadrangle, Santa Ana Mountains,

Orange County, southern California. Lower Campanian.

Ladd Formation, upper Holz Shale Member. Coll.: T

Bear, 1940.

93639. East bank of Chico Creek in concretions in

massive, greenish-gray sandstone, 373 m south and

ane m west of northeast corner of section 23, T. 23 N,

R. 2 E, Paradise Quadrangle (15 minute, 1953), Butte

County, northern California. Lower eee Chico

Formation, Ten Mile Member. Coll.: _ and R. B.

Saul, August 20, 1952.

23642. On W bank of Chico Creek, west of big westward

projection of east lava cap and approximately N 27°W of

BM 1770, 122 m south and 632 m west of northeast

corner of section 26, T. 23 N, R. 2 E, Paradise

Quadrangle (15 minute, 1953), Butte County, northern

California. Lower Campanian. Chico Formation, Ten

Mile Member. Coll: L. R. and R. B. Saul, August 20,

1952.

23643. Concretionary sandstone on west side of Big

Chico Creek, 670 m south and 762 m west of merheaet

corner of section 26, T. 23 N, R. 2 E, Paradise

Quadrangle (7.5 minute, 1953), Butte County, northern

California. Lower Campanian. Chico Formation, Ten

Mile Member. Coll.: L. R. Saul and R. B. Saul, August,

1952.

24124. Hard, medium-grained, gray, arkosic sandstone,

158 m north of Jalama Creek, 3.3 km west and 0.6 km

THE NAUTILUS, Vol. 119, No. 4

north of southeast corner of the topographic sheet,

Lompoc Hills Quadrangle (7.5 minute, 1947). Upper

Campanian. Jalama Formation. Coll.: D. Dailey and J. R.

Dorrance, October, 1929.

24125. Hard, fine to medium-grained arkosic sandstone,

2.8 km west and 0.53 km aortli of southeast corner of

the topographic sheet, Lompoc Hills Quadrangle

(7.5 minute, 1947). Upper Campanian. Jalama Forma-

tion. Coll.: D. Dailey and J. R. Dorrance, October 29,

1929.

24917. Hard sandstone slabs in bed of Clover

Creek, 213 m north and 366 m west of southeast

corner of section 22, T. 32 N, R. 2 W, Millville

Quadrangle (15 minute, 1953), Shasta County, northern

California. Upper Santonian. Redding Formation,

Member VI. Coll: W. P. Popenoe and D. Dailey,

August 27, 1959.

24337. Along east side of Highway 70, 792 m north and

305 m west of southeast corner of section 36, T. 21 N, R.

3 E, Cherokee Quadrangle (7.5 mintue, 1949), Butte

County, northern California. Lower Campanian. Chico

Formation, Pentz Road member (informal). Coll.: E

Gohre.

UCMP B-3388. In south-trending arroyo which reaches

coast aoiit 2.8 km north of the tip of Punta Baja.

Marine invertebrates from massive, chert conglomerate

at confluence of this arroyo and northwest- trending

tributary about 122 m upstream from confluence. This

locality is about 488 m northwest of the Punta Baja road.

Middle upper Campanian. Punta Baja Formation. Coll:

F. H. Kilmer, date unknown.

USGS M-S8601. In lens in sandstone on beach 0.5 km

southeast of Bolsa Point, Pigeon Point Quadrangle, San

Mateo County, northern California. Middle Campanian.

Pigeon Point Formation. Coll.: W. P. Elder, 1989?

THE NAUTILUS 119(4):149-152, 2005

Page 149

Daffymitra lindae, a new genus and species of Volutomitridae

(Neogastropoda) from he Bellingshausen Abyssal Plain

M. G. Harasewy ch Yuri I. Kantor

Department of Invertebrate Zoology

National Museum of Natural History

Smithsonian Institution

Washington DC 20013-

7012 USA Moscow 117071

RUSSIA

Severtzov Institute

Russian Academy of Sciences

Leninski Prospect, 33

ABSTRACT

Daffymitra lindae, new genus, new species, is described from

the Bellingshausen Abyssal Plain off Ellsworthland, Antarctica.

Known only from its shell, this new taxon is included in the

family Volutomitridae, but differentiated from all known living

genera and species on the basis of its inflated shell shape, with

an attenuated anterior and distinctive siphonal canal, as well as

by the shape and disposition of its three recessed, obliquely

oriented columellar plaits. This new taxon represents the first

record of Volutomitridae from abyssal depths. The concholo-

gical similarity of Daffymitra to the Upper Cretaceous genus

Volutomorpha raises the possibility that Daffymitra may be

a surviving descendent of a lineage presumed extinct since the

end of the Cretaceous, and suggests that a reassessment of the

relationships between the various Cretaceous genera assigned

to Volutoderminae and the earliest Volutomitridae.

INTRODUCTION

The Volutomitridae is a small family of rachiglossan

oe a characterized by: a fusiform shell (reach-

ing 50 mm) with 2-5 columellar plaits, and a paucispiral,

mammilate protoconch; a radula with distinctive, wish-

bone-shaped rachidian teeth and small, needle-like

lateral teeth that are present in most species; a mid-

oesophagus that is long, convoluted, with a muscular

posterior end: a gland of Leiblein that is only partially

separated from the mid-oesophagus; and a_ single

accessory salivary gland (Ponder, 1972, 1998; Kantor

and Harasewych, 1992: Bouchet and Kantor, 2004). The

family ranges into the North Pacific, North Atlantic and

the Caribbean, but centers of diversity are concentrated

in the southern hemisphere, particularly in the waters

surrounding Antarctica, Australia, New Zealand, and

especially New Caledonia (Bouchet and Kantor, 2004),

Six genera and 50 species are currently recognized in the

Recent fauna (Bouchet and Kantor, 2004), with a single

genus (Paradmete) containing six species represented in

the Antarctic and Magellanic fauna (Numanami, 1996;

Bouchet and Kantor, 2004). Volutomitrids are known

from the sublittoral zone to depths of 1980 m (based on

dead specimens), but the majority of species are

confined to outer continental shelf and upper continen-

tal slope depths (Bouchet and Kantor, 2004: Appendix;

see Figure 7 herein),

In the course of our studies of Antarctic and sub-

Antarctic neogastropods sampled by the United States

Antarctic Program (USAP) and housed in the collections

of the National Museum of Natural History (USNM), we

encountered a single specimen of a volutomitrid that was

collected on fhe: Bellinghausen Abyssal Plain. This

specimen represents a new genus and species within

the family Volutomitridae, as well as the first record of

the family from abyssal depths. This new genus and new

species is described herein, and comparisons made with

other volutomitrid taxa.

SYSTEMATICS

Class Gastropoda Cuvier, 1797

Order Neogastropoda Wenz, 1935

Family Volutomitridae Gray, 1854

Genus Daffymitra new genus

Type Species: Daffymitra lindae new species, by

original designation.

Description: Shell medium-sized for the family (to

28 mm), with low, conical spire, broadly inflated body

whorl. Protoconch conical, large, 2.6 mm in diameter, of

2 5/8 smooth whorls. Teleocouch thin, of about 3

smoothly rounded whorls, with well demarcated, broad

siphonal canal. Axial sculpture of distinct, very narrow,

broadly spaced ribs that extend from suture to siphonal

canal. Spiral sculpture of narrow, cords, alternating in

prominence. Aperture high (74% of shell length) broadly

ovate. Outer lip thin, smooth. Columella with three

columellar folds (central strongest) and siphonal fold.

Shell white. Periostracum thin, olive brown. Operculum,

anatomy, radula, unknown.

Etymology: This genus is named after Daffy, a tor-

toiseshell longhair cat that belongs to the senior author's

wife Linda.

Page 150

THE NAUTILUS, Vol. 119, No. 4

Figures 1-6. Daffymitra lindae new species, holotype. 1. Apertural view of the shell. 2. Oblique view, shell is rotated slightly to

expose the columellar plaits. 3. Lateral view of the shell. 4. Dorsal view of the shell. 5. Apical and 6. lateral views of the protoconch.

Transition to teleoconch is marked by arrow.

Daffymitra lindae new species

(Figures 1-7)

Description: Shell (Figures 1—t) of moderate size (to

27.9 min), thin, fragile, with matte surface, inflated,

fusiform, tapering anteriorly, with conical spire. Proto-

oOo

o 7

conch (Figures 5-6) large, mammilate, 1970 um in

height, diameter increasing from 676 Um to 2570 Lm in

2.5/8 convex whorls. Protoconch-teleoconch transition

distinct (Figures 5, 6, arrow), marked by onset of weak

closely spaced prosocline ribs. Teleoconch of 3.1/8

strongly convex, ovate whorls with rounded shoulder.

Suture impressed Axial sculpture of thin, sharply

demarcated, weakly prosocline raised ribs on last

whorl, 34 on penultimate whorl. Ribs closely spaced on

first teleoconch whorl, becoming more widely spaced on

later whorls, but again closely spaced along final vs

whorl. Spiral sculpture of very low, narrow cords,

subequal in width, alternating in prominence, covering

entire shell surface, about 50 on final whorl, 10 on

penultimate whorl. Aperture large (0.74 shell length),

broadly oval, smooth, deflected from shell axis by 15°.

Outer lip very thin, weakly reflected, edge forming final

axial rib, with shallow anal sinus at suture. Columella

weakly sinuate, convex posteriorly, distinctly concave

medially, and again anterior to 3 obliquely oriented,

recessed folds (Figure 2). Central fold most pronounced,

anteriormost fold even more obliquely oriented than

central and posterior fold. Parietal callus, broad, very

thin. Siphonal canal broad, long, well delimited from

M. G. Harasewych and Y. I. Kantor, 2005

Page 15]

Number of Species

0 5 10 15 20 25

1000

1200

1400

1600

Depth (m)

1800

2000

2500 ae :

O Volutomitridae, all species

3000

det Paradmete only

4000

4500 ae Dattymitra lindae

5000

6000

Figure 7.

Volutomitridae, of the Antarctic genus Paradmete (data from

Bouchet and Kantor, 2004: Appendix), and of Daffymitra

lindae. A: continental shelf; B: upper continental slope; C:

lower continental slope; D: continental rise; E: abyssal plain, F:

hadal depths.

aperture. Shell color white. Periostracum thin, olive

brown, covering entire shell. Operculum, radula and

anatomy unknown.

Type Locality: —Bellingshausen Abyssal Plain, 61°27’ S,

94°5S'-95°22’ W. in 4419-4804 m [R/V ELTANIN

cruise 23, sta. 1621, 10 Apr. 1966].

Type Material: Holotype, USNM 1050443, shell

length 27.9 mm, final whorl length 23.4 mm, aperture

length 21.5 mm, shell width, 13.5 mm.

Distribution: Known from the type locality only.

Etymology: This species is named for the senior

author's wife, Linda Lee Harasewych.

Remarks: Despite the absence of anatomical and

radular data, this new species can be unambiguously

assigned to the family Volutomitridae on the basis of its

distinctive shell shape, sculpture, presence of the

diagnostic paucispiral mammilate protoconch, and weak

columellar folds.

Bathymetric distribution of the Recent species of

Seven genera are currently recognized within i

family Volutomitride ae (C ernohorsky, 1970; Bouchet and

Kantor, 2004), The large size, long, broad aperture, and

thin shell of this new species, as well as the presence of

three obliquely oriented and deeply recessed columellar

folds and a siphonal fold preclude its rae to

either the fossil genus Proximitra Finlay, 1927, or the

Recent genera Conomitra Conrad, 1865: hie

Angas, 1877; Peculator Iredale, 1924; or Magdalemitra

Kilburn, 1974.

Conchologically, the new species is closer to the

genera Volutomitra H. and A. Adams, 1853 and

Paradmete Strebel, 1908, which are considered to be

closely related (Powell, 1951: 165; Cernohorsky, 1970:

91). The 13 known species of Voliutomitra are widely

distributed in the World Ocean, ranging from South

Africa, Southern Australia, New Talend and to the

Bering Sea in the Pacific, and from Colombia to the

northern part of the Atlantic Ocean. Daffymitra lindae

differs from all known species of Volutomitra in having

a proportionally shorter spire and inflated rather than

fusiform shell, coarse spiral sculpture and sharp, narrow,

broadly spaced axial ribs, as well as columellar folds that

are weak, recessed within the aperture and obliquely

oriented rather than being strong, prominent, and nearly

perpendicular to the édhunvellar axis. The only species of

Volutomitra with pronounced axial sculpture, V. erebus

Bayer, 1971, from Colombia, has axial ribs that are

thicker, more rounded, orthocline, and more densely

spaced.

The genus Paradmete, contains six species, all

confined to Antarctic and sub-Antarctic waters. Daffy-

mitra lindae may easily be distinguished from Paradmete

fragillima (Watson, 1882), the type species, as well as

from P. briedensis Numanami, 1996, and P. arnaudi

Numanami, 1996, by its larger size, shorter spire,

inflated rather than narrowly fusiform shell, and

distinctive narrow, prosocline axial ribs, as well as by

its well demarcated siphonal canal. The Magellanic

Paradmete crymochara (Rochebrune and Mabille, 1885)

approaches Daffymitra lindae in size, but differs in its

elongate, fusiform shape, absence of a distinct siphonal

canal, and presence of four columellar folds. The

distinctive Paradmete percarinata Powell, 1951, can Hs

recognized by its prominent peripheral carina, sharply

shouldered shell and pronounced columellar folds that

are nearly perpendicular to the columellar axis. Most

similar to Daffymitra lindae is Paradmete curta (Strebel,

1908), which reaches a similar size, has a low spire, and

has axial ribs, which, however, are opisthocline rather

than prosocline. Daffymitra differs in lacking a strong

shoulder and in having an inflated shell shape with an

attenuated anterior and distinctive Si} shonal canal.

The shell of Daffymitra lindae aes a surprising

resemblance to some members of the Mesozoic genus

Volutomorpha, particularly V. mutabilis Wade, 1926 (see

Wade, ao pl. 37, fig. 10, pls. 40, figs. 6, 9; Sohl, 1964:

pl. 39, figs. 1, 2, 6). Volutomorpha was restricted to the

Upper Cretaceous faunas of the Gulf and Atlantic

Page 152

THE NAUTILUS, Vol. 119, No. 4

coastal plains (for a review, see Sohl, 1964: 252-254),

and was “the giant of Cretaceous gastropods” (Wade,

1926: 20) with shell lengths extrapolated to exceed

45 em. Pilsbry and Olson (1954: 19) included Voluto-

morpha in the Cretaceous subfamily Volutodermatinae,

which they placed in the family Volutidae together with

Volutomitrinae. More recently, Dzhalilov (1977: 93)

proposed a new subfamily Volutomorphinae, also within

Volutidae, while Bouchet et al. (2005: 255) considered

Volutomorphinae a synonym of Volutodermatinae,

which they transferred from Volutidae to the extinct

family Pholidotomidae.

While Daffymitra is easily distinguished from Voluto-

morpha by its far smaller size, thinner shell, absence of

thick axial ribs, and lack of a pronounced shoulder, this

conchological similarity raises the intriguing possibility

that Daffymitra is a “living fossil,” a surviving de-

scendent from a group presumed to have become extinct

at the end of the Cretaceous. Further research is clearly

required to reevaluate the relationships between the

various Cretaceous genera assigned to Volutoderminae

and the earliest Volutamitridae.,

DISCUSSION

The family Volutomitridae has a broad geographic range,

but has previously been known only from continental

shelf and continental slope faunas, while the genus

Paradmete has been reported only from shelf and upper

slope depths (Figure 7). The greatest diversities for both

the family and the genus occur at upper continental

slope depths.

Although Daffymitra lindae is represented by a single

empty shell. the fragility of the shell, the presence “of

periostracum, and the fact that it was collected below the

aragonite compensation depth indicate that the speci-

men could not have been dead for long, and that the

species inhabits the area in which this specimen was

collected. Thus, this taxon represents the first record of

Volutomitridae from abyssal depths. In a survey of

Antarctic oe Magellanic Buccinoidea, Harasewych and

Kantor (2004) 4) found that the abyssal buecinoidean fauna

of the aes has no genera in common with the

sublittoral or bathyal faunas, but that credible sister taxa

and likely origins for at least some of the abyssal genera

occur on the ‘adjacent continental slope. Based on shell

morphologies, the genera Volutomitra, Paradmete, and

Daffymitra appear to represent a lineage within

Volutomitridae distinct from the predominantly austral

genera Proximitra, Conomitra, Microvoluta, Peculator,

and Magdalemitra. The genus Paradmete, a member of

the upper slope fauna of Antarctica, is likely the sister

taxon of the abyssal genus Daffymitra.

ACKNOWLEDGMENTS

This research was supported by a grant from the NSF —

USAP United States Antarctic Program {Contract

Number OPP-9509761]. We are grateful to Bruce

Marshall for bringing to our attention the similarity of

Daffymitra and Volutomorpha.

LITERATURE CITED

Bouchet, P., J. Fryda, B. Hausdorf, W. F. Ponder, A. Valdés and

A. Warén. 2005, Part 2. Working Classification of the

Gastropoda [in] Bouchet, P. and J. P. Rocroi. Classification

and Nomenclator of Gastropod Families. Malacologia 47:

1-397.

Bouchet, P. and Y. I. Kantor. 2004. New Caledonia: the major

centre of biodiversity for volutomitrid mollusks (Mollusca:

Neogastropoda: Volutomitridae). Systematics and Bio-

diversity 1(4): 467-502.

Cernohorsky, W. O. 1970. Systematics of the families Mitridae

and Volutomitridae (Mollusca: Gastropoda). Bulletin of

the Auckland Institute and Museum 8: 1-190.

Dzhalilov, M. R. 1977. Cretaceous gastropods of Southeastern

Central Asia. Danek: Dushanbe, 1-202 pp.

Harasewych, M. G. and Y. I. Kantor. 2004. The Deep-Sea

Buccinoidea ( (Gastropoda: Neogastropoda) of the Scotia

Sea and Adjacent Abyssal Plains and Trenches. The

Nautilus 118: 1-42.

Kantor, Y. I. and M. G. Harasewych. 1992. Morphology of the

digestive system of Volutomitra alaskana Dall, 1902

(Gastropoda, Pectinibranchia, Volutomitridae), with notes

on the possible mechanism of feeding. Ruthenica 2:

45-53.

Numanami, H. 1996. Taxonomic study on Antarctic Gastro-

pods collected by Japanese Antarctic Research Expedi-

tions. Memoirs of National Institute of Polar Research

Series E (Biology and Medical Science) 39: 1-244.

Pilsbry, H. A. and A. A. Olsson. 1954. Systems of the Volutidae.

Bulletin of American Paleontology 35(152): 275-306, 4 pls.

Ponder, W. F. 1972. The morphology of some mitriform

gastropods with special reference to their alimentary and

reproductive systems (Neogastropoda). Malacologia 11:

295-342.

Ponder, W. F. 1998. Family Volutomitridae. pp. 842-843. In:

P. L. Beesley, G. J. Ross and A. Wells (eds) Mollusca: The

Southern Synthesis. Fauna of Australia. Vol. 5. CSIRO

Publishing: Melbourne, Part B viii, 565-1234 pp.

Powell, A. W. B. 1951. Antarctic and SubAntarctic Mollusca:

Pelecypoda and Gastropoda. Discovery Reports 26:

47-196, pls. 5-10.

Sohl, N. F. 1964. Neogastropoda, Opisthobranchia and Basom-

matophora from the Ripley, Owl Creek, and Prairie Bluff

Formations. United States Geological Survey cee

Paper 331-B: i-iv, 153-344, pls. 19-52, tables 1-2

Wade, B. 1926. The fauna of the Ripley Fonnation on Coon

Creek, Tennessee. United States Geological Survey Pro-

fessional Paper 137: 1-272, 72 pls., 2 figs. ,

THE NAUTILUS 119(4):153-156, 2005

Exilia alanbeui, a new species from the Neogene of central Chile:

the first record of Exilia (Gastropoda: Ptychatractidae) from

South America

Sven N. Nielsen

Freie Universitit Berlin

Institut fiir geologische Wissenschaften

Fachrichtung Paliontologie

Malteserstrasse 74-100

Haus D, 12249 Berlin

GERMANY

[email protected]

ABSTRACT

A new species of the ptychatractid genus Exilia, E. alanbeui

new species, is described from late Neogene sediments of the

Navidad Formation, central Chile. This new species represents

the first record, fossil or Recent, of the genus Exilia from the

South American continent. Given that Recent species of Exilia

are restricted to bathyal depths, the presence of representa-

tives of the genus in fine siltstones of the Navidad and Ranquil

formations lend support to a previous interpretation of bathyal

depths for these deposits.

INTRODUCTION

The genus Exilia Conrad, 1860, has been placed in the

family Turbinellidae by Maxwell (1988) and in the

subfamily Ptychatractinae within the Turbinellidae by

Kantor et al. (2001). The subfamily Ptychatractinae was

removed from the Turbinellidae and raised to family

level by Riedel (2000), as was suggested previously by

Kantor and Bouchet (1997). Riedel (2000) placed the

Turbinellidae together with other columellar plaits-

bearing groups in his suborder Volutina while the

Ptychatractidae were placed in the suborder Muricina

based mainly on radula type and absence of columellar

plaits. The most recent classification is that presented by

Bouchet and Rocroi (2005), placing Ptychatractidae

within Pseudolivoidea. The genus Evxilia has been

revised by Bentson (1940) and more recently by Kantor

et al. (2001). The latter authors Heyes nine

nominal genera, i.e., Mitraefusus Bellardi, Mesor-

hytis Meek, 1876, Surculina Dall, 1908, eh ee

Dall, 1918, Palaeorhaphis Stewart, 1927, Zexilia Finlay,

1926. Graphidula Stephenson, 1941, Benthovoluta

Kuroda and Habe, 1950, and Chathamidia Dell, 1956,

containing fossil and Recent species with Evxilia and

recognized a total of nine living species. Fossil species

were not formally treated by them but a number of

species were figured for comparison. Previously, the

Recent species of the synonymized genus Be mthovoluta

had been reviewed by Cernohorsky (1973) and Har-

asewych (1987). Species of Exilia are ee from the

Late Cretaceous (Coniacian) onward and fossil species

are known from all continents except Africa and South

America, “probably a reflection of insufficient studies of

Cretaceous and Tertiary mollusks in these regions”

(Kantor et al., 2001, p. 92). Some Cretaceous through

Miocene species come from shallow-water deposits

while deep-water occurrences are known since the late

Eocene (Kantor et al., 2001). Recent species of Exilia are

restricted to bathyal depths at tropical latitudes and in

the New Zealand region (Kantor et al., 2001).

GEOLOGY OF THE FOSSIL-BEARING LOCALITIES

The Navidad Formation (Darwin, 1846) and its equiva-

lents have recently been re-interpreted as late Miocene

bathyal basin deposits containing displaced intertidal to

outer shelf sediments (Finger et al., 2003; Nielsen et al.,

2003). Several fossiliferous sites have been sampled for

micro- and macrofossils. Of these, two localities from the

Navidad Formation (Figure 1B) and two from the

Ranquil Formation (Figure 1C) yielded specimens of

Exilia described here.

Locality PPP (Figure 1B) is a grey dee p-water

siltstone that today Reais the intertidal platform at

Punta Perro. Based on planktonic foraminifera this

locality has been dated as late Miocene (Tortonian) by

Tsuchi et al. (1990) and Ibaraki (1992), while new dating

by Finger et al. (2003) suggests an even younger age.

The fauna is of an unusual composition for the Navidad

Formation, containing otherwise unknown species of

Struthiochenopus (Zinsmeister and Griffin, 1995) and

Xenophora (Nielsen and DeVries, 2002). The specimens

coming from Pupuya (Figure 1B) were collected by

Page 154

THE NAUTILUS, Vol. 119, No. 4

j 70°

ex.

Santiago

(_— 4

«Matanzas

-\--

27~7""Navidad ‘8

Isla Coronel

Sta. Maria

. Lavapié

Pta. Perro

> ;

: -@®Curanilahue

Los Alamos

@ Caiiete

0 sf io 15820 25km

Figure 1. Fossil localities for Exilia alanbeui new species. in central Chile. A. Location of working areas. B. Localities of the

Navidad Formation, Platform Punta Perro (PPP), Pupuya. C. Localities of the Ranquil Formation, Punta El Fraile (FRM), northern

end of Caleta Ranquil (RQT), southwestern end of Caleta Ranquil.

V. Covacevich and D. Frassinetti a little south of the

village Pupuya and come from a grey siltstone for which

no ages are available. However, the faunal content (e.¢.,

Nielsen et al., 2004) justifies correlation with PPP. The

sediment at the type locality Punta El Fraile (FRM,

Figure 1C) consists of grey siltstone and underlies

a yellowish sandstone typical for the Ranquil Formation.

The limit between these two lithological units has been

observed to be a sharp, concordant one including some

pebbles at the boundary. The sediments at Caleta

Ranquil (ROT and MIB, Figure 1C) consist of a grey

siltstone similar to that present at Punta El Fraile. The

geological relation with other lithologies present at this

locality, like reddish sandstone with beds of glauconitic

sandstone and coarse light-grey sandstone, are not yet

clear. However, both localities of the Ranquil Formation

are similar to those of the Navidad Formation in

lithology, and faunal content and were also dated as late

Miocene by Finger et al. (2003).

MATERIALS AND METHODS

Specimens described in this study are deposited in the

collections of the Departamento de Paleontologia de

Invertebrados Museo Nacional de Historia Natural,

Santiago de Chile (SGO.PI) and Senckenberg Museum,

Frankfurt, Germany (SMF). Photographs were taken

using a Leicaflex SL2 or Olympus Camedia c730

camera. Images taken with the former camera were

scanned from [ford FP4 125 black and white 35 mm

negatives using an Acer ScanWit 2720S film scanner. All

images were processed with Adobe Photoshop 7.0.

SYSTEMATIC PALEONTOLOGY

Family Ptychatractidae Stimpson, 1865

Genus Exilia Conrad, 1860

Type Species: — Exilia pergracilis Conrad, 1860, by

monotypy: Midway Group, lower Eocene, Alabama, USA.

Exilia alanbeui new species

(Figures 2-14)

Description: Shell narrow-fusiform, with spire occu-

pying only little more than half of total height.

Protoconch unknown. Teleoconch of about seven

straight to slightly convex whorls, lacking pronounced

shoulder. Axial sculpture consists of rounded, slightly

S. N. Nielsen, 2005

Figures 2-14. Exilia alanbeui new species. 2-4. Holotype FRM 019 (height 28.35 mm). 5-6. Paratype FRM 012-3 (height

30.2 mm). 7-8. Paratype FRM 012-2

(height 18.03 mm). 12.

height 23.9 mm).

backwards bent, collabral ribs. Penultimate whorl bears

about 16 ribs, which become obsolete on base of whorl.

Spiral sculpture well and equally defined, consisting of

narrow, low, rounded, subequal cords, crossing axial

elements unchanged, separated by interspaces of about

equal width. Fine secondary spiral threads present in

some interspaces. On penultimate whorl 10-14 spiral

cords present, S—9 further cords present on base of last

whorl and about 30 weaker, closely spaced cords on

siphonal canal. Aperture narrow-elongate. Outer lip

thin. Siphonal canal narrow, long, straight. Inner lip

smooth. Columella smooth and almost straight, with

weak swelling at entrance of siphonal canal. -

Etymology: This species is named in honour of Alan

G. Beu (Institute of Geological and Nuclear Sciences,

New Zealand), who always helps when needed.

Type Material: Holotype SGO.PI.6371 (FRM, height

28.35 mm), paratypes SGO.PL.6372 (FRM, one speci-

men, height 10.5 mm), SGO.PI.6373 (FRM, 21 speci-

mens), SMF 327749 (FRM, 19 specimens).

Other Material Examined: SGO.PI.6374 (PPP,

height 23.9 mm), SGO.PI.6375 (PPP, one specimen),

SGO.PI.5303 ( (Pupuya, four specimens), SGO.PI.6376

(ROT, one specimen, height 9.25 mm), SGO.PI.6377

(MIB, one specimen, height 29.91 mm).

Type Locality: The grey siltstone at Punta El Fraile

(FRM). Arauco, Souther central Chile.

(height 15.46 mm). 9-10. Paratype FRM 012-1 (height 9.3 mm). 11. Paratype FRM 012-4

Paratype FRM 012-5 (height 17.62 mm). 13. Paratype FRM 012-6 (height 15.2 mm). 14. PPP 017 (PPP,

Occurrence: PPP, Pupuya, FRM, MIB, ROT; Navi-

dad and Ranquil formations, Late Miocene or Early

Pliocene, central Chile.

Discussion: Exilia alanbeui resembles the type

species, EF. pergracilis (see Bentson, 1940, pl 2,

fig. 25; Kantor et al, 2001, figs. 1 A-D), and

E. lincolnensis Weaver, 1916, but differs from those

North American Eocene species in having a_ less

convex whorl profile, almost straight instead of sinusoi-

dal axial sculpture and a more angulated whorl base.

It differs from the New Fede Miocene species

E. nodulifera (Marwick, 1931), E. leachi (Marwick,

1931) and E. wellmanni Maxwell, 1988 (see Maxwell

1988) in having stronger axial sculpture. Most species

of Exilia have more convex whorls and_ especially

the Recent species have weaker axial sculpture

(see Harasewych, 1987 and Kantor et al., 2001).

The geographically closest species, E. cortezi, has

wider whorls and different axial sculpture and _ is

therefore not considered to be closely related to

E. alanbeui. Exilia alanbeui is known only from

localities consisting of deep water siltstones which

have been dated as probably Messinian in age based

on foraminifera (Finger et al. 2003). Therefore,

this species is younger than the above mentioned

Miocene — species from New Zealand. Regarding

the similarity of the two species, E. alanbeui may prove

to be closely related to the New Zealand species

E. wellmanni.

Page 156

CONCLUSIONS

The occurrence of Exilia alanbeui in central Chile is the

first record of the genus from the South American

continent (except the Recent E. cortezi (Dall, 1908) from

the Galapagos Islands) and therefore closes a consider-

able biogeographic gap. Although shallow water species

of Exilia are known from Miocene deposits of Europe,

the occurrence of Exilia in siltstones of the Navidad and

Ranquil formations is here interpreted to support

microfossil data placing these deposits in a bathyal

environment (Finger et al., 2003).

Exilia alanbeui is another Chilean Miocene gastropod

species that shows many similarities to New Zealand

taxa. Trans-Pacific biogec ygraphic connections between

Chile and New Zealand during the Miocene have

already been shown for some vetigastropods ( (Nielsen

et al., 2004) and xenophorids (Nielsen and DeVries,

2002).

ACKNOWLEDGMENTS

I thank Klaus Bandel (Universitat Hamburg, Germany),

who helped in numerous ways and provide rd the

infrastructure to continue this work while I was in

Hamburg. Paulina Vasquez (TU Berlin, Germany) found

the figurec | specimen from PPP and was a cheerful help

during { field work. Daniel Frassinetti (SGO.PI, Chile)

provided access to collections under his care. Tom

DeVries (Burton, USA) improved the language of an

earlier draft. Constructive reviews by Philippe “Botichet

(Muséum National d’Histoire Naturelle Paris, France)

and an anonymous reviewer are gratefully acknowl-

edged. The material has been eéllecte d during field work

Aarne by the Deutsche Forschungsgemeinschaft

grant Ba 675/25 during the years 2000 to 2002.

LITERATURE CITED

Bentson, H. 1940. A systematic study of the fossil gastropod

Exilia. U niversity of California Publications 25: 199-238.

Bouchet, P. and J. P. Rocroi, 2005. Classification and nomen-

clator of ia rd families. Malacologia 47, in press.

Cernohorsky, W. O. 1973. The taxonomy of Benthovoluta

hilge moerh | von Martens) and allied turbinellid genera

(Mollusca: Volutacea). Records of the Auckland Institute

and Museum 10: 123-131.

Tsuchi, R., T

THE NAUTILUS, Vol. 119, No. 4

Darwin, C. 1846, Geological observations on South America.

Smith, Elder & Co., London, 279 pp.

Finger, k., A. Encinas, S. Nielsen and D. Peterson. 2003.

Microfaunal indications of Late Miocene deep-water

basins off the central coast of Chile. 10° Congreso

Geolégico Chileno, Concepcion, Chile. Abstract Volume

CD-ROM. 8 pp.

Harasewych, M. G. 1987. A revision of the genus Benthovoluta

with notes on the evolution of the subfamily Ptychatracti-

nae (Prosobranchia: Turbinellidae). The Nautilus 101:

166-181.

Ibaraki, M. 1992. Planktonic foraminifera from the Navidad

Formation, Chile: their geologic age and paleoceano-

graphic implications, pp. 91-95. In K. Ishizaki and

T. Saito (eds.), Centenary of Japanese Micropaleontology.

Terra Scientific Publishing Company, Tokyo.

Kantor, Y. I. and P. Bouchet. 1997. The Ptychatractinae: an

endemic deep-sea clade of the Turbinellidae? American

Malacological Union, 63"! Annual Meeting (Santa Bar-

bara, California), Abstracts: 37-38.

Kantor, Y. I., P. Bouchet and A. Oleinik. 2001. A revision of the

Recent species of Exilia, formerly Benthovoluta (Gastrop-

oda: Turbinellidae). Ruthenica 11: 81-136.

Maxwell, P. A. 1988. Late Miocene deep-water Mollusca from

the Stillwater Mudstone at Greymouth, Westland, New

Zealand: paleoecology and systematics. NZ Geological

Survey Paleontological Bulletin 55: 120 pp.

Nielsen, S. N. and T. J. DeVries. 2002. Tertiary Xenophoridae

(Gastropoda) of western South America. The Nautilus

116: ee

Nielsen, S. N., T. J. DeVries, A. Encinas, K. L. Finger and D.

Peterson, sie Towards an understanding of the age of

the Navidad Formation. 10° Congreso Ge ol6gico Chileno,

Concepcion, Chile. Abstract Volume CD-ROM. 7 pp.

Nielsen, S. N., D. Frassinetti and K. Bandel. 2004. Miocene

Vetigastropoda and Neritimorpha (Mollusca, Gastropoda)

of Central Chile. Journal of South American Earth

Sciences 17: 73-88.

Riedel, F. 2000. Ursprung und Evolution der “héheren”

Caenogastr ypoda. Berliner Geowissenschaftliche Abhand-

lungen E32. 240 Pp

. Shuto, T. Takayama, I. Koizumi, A. Fujiyoshi, R.

Nomura, M. Ibaraki, H. Duque-C, R. Tirado-S, M.

Aldana-A, E. Villavicencio-R and R. Martinez-P. 1990.

Trans-Pacific correlation of Neogene geologic events.

Reports of Andean Studies, Shizuhoka University, Special

vol. 3: 1-7.

Zinsmeister, W, J. and M. Griffin, 1995, Late Cretaceous and

Tertiary aporrhaid etl es from the southern rim of

the Pacific Ocean. Journal of Paleontology 69: 692-702.

THE NAUTILUS 119(4):157-163, 2005

Page 157

Spawn of Amphissa sp. and Cosmioconcha sp. (Caenogastropoda:

Columbellidae) from the Colombian Caribbean

Patricia Miloslavich

Ana Karinna Carbonini

Departamento de Estudios Ambientales

Universidad Sim6n Bolivar

P.O. Box S9000

Caracas LOSO

VENEZUELA

[email protected]

Bogota D.C.

COLOMBIA

Juan Manuel Diaz

Instituto Alexander von Humboldt

Néstor E. Ardila

Instituto de Investigaciones Marinas

INVEMAR

Santa Marta, A.A. 1016

COLOMBIA

ABSTRACT

We describe the egg capsules and embryos of Amphissa sp. and

Cosmioconcha sp. collected in the Caribbean Sea off Colombia

between 20-160 m depth. Amphissa sp. had one layer of 24

egg capsules attached to the shell.

completely covered by several layers of egg capsules; only the

outer layer capsules contained embryos, the rest had the

escape aperture open. Egg capsules of both species were

translucent, dome-shaped, with an oval escape aperture at the

center of the dome top. The surface was smooth, with no ridges

nor sutures, and attached to the shell by an oval basal

membrane that was surrounded by a thin, irregular flange. Egg

capsules of Amphissa sp. measured 2 mm in diameter and

Cosmioconcha sp. was

contained 6 embryos/capsule at the gastrula stage; egg capsules

of Cosmioconcha sp. measured 1 mm in diameter and

contained 25 embryos/capsule at all stages of development

(eggs, trochophores, and veliger larvae). Gastrulae of Amphissa

sp. measured 750-900 {tm in length and development in this

species is probably direct. Uncleaved eggs of Cosmioconcha sp.

measured 150 tm in diameter, the ve liger had an operculum,

a transparent shell measuring 188-219 um in length, and

a small velum; hatching occurs as planktonic larvae. No nurse

eggs were observed in the two species but late cannibalism

among sibling embryos may occur in Cosmioconcha sp.

INTRODUCTION

The family Columbellidae is very diverse in number of

species in the Southern Caribbean. In this region, a total

of 13 genera (Columbella Lamarck, 1799; Rhombinella

Radwin, 1968; Amphissa H. and A. Adams, 1853;

Zafrona Tredale, 1916; Anachis H. and A. Adams,

1853: Nassarina Dall, 1889: Mitrella Risso, 1826:

Cosmioconcha Dall, 1913; Aesopus Gould, 1860; Deci-

pifus Olsson and McGinty, 1958: Mazatlania Dall, 1900;

Strombina Morch, 1852: Nitidella Swainson, 1840, and

Pyrene Réding, 1798) and more than 30 species have

been recorded (cf. Radwin, 1877 a, b; 1978; Costa,

2005): however, columbellid classification, as well as the

identity and interpretation of the characters that support

this classification, is not well resolved (DeMaintenon,

1999). The shells of these specie s are usually small and

fusiform, within a size range of 5 to 24 mm and the outer

lip usually bears denticles. The egg capsules and

reproduction of several species have been described

worldwide: Thorson (1940) summarized the different

forms of egg capsules found in species from the Iranian

Gulf, Knudsen (1950, 1995) described the egg capsules

and development of species of tropical West Africa and

the Azores, Amio (1957, 1963) described the eggs, larvae

and embryology of Japanese species, DAsaro (1970)

described the egg capsules of Panamanian Pacific

species, Marcus and Marcus (1962) described the

reproductive biology of several species from Brazil and

Fortunato et al. (1998) characterized the reproduction of

Bifurcium bicanaliferum (Sowerby, 1832) from the

Pacific coast of Panama. In the Caribbean region,

Bandel (1974) described the egg capsules of 10 species

of Columbellidae from Santa Marta, Colombia; Pench-

aszadeh et al. (1983) characterized the reproduction of

Mazatlania consentini Philippi, 1836 (as M. aciculata)

(Lamarck, 1822) and Cipriani and Penchaszadeh (1993)

that of Strombina francesae |. Gibson—Smith, 1974, and

Strombina pumilio (Reeve, 1859), all three from

Venezuela. Despite these studies, egg capsule morphol-

ogy and developmental mode is still inknow for most of

the recognized species.

Bandel (1974) distinguished 6 morphological groups

of columbellid egg capsules according to the material he

observed in Colombia and the literature. These are: (1)

shallow domes on a broad, irregularly rounded disk of

adhesion only extending on a narrow rim beyond the

capsule walls as in Mitrella ocellata (Gmelin, 1791)

reported as Mitrella argus (d@Orbigny, 1542), (2) cone

shaped capsule, i walls rise from the round adhesion

disk and end in a projecting edge forming a collar

around the pane aperture, which is concave as in

Costoanachis sparsa (Reeve, 1859), (3) cupola shaped

with a suture dividing the capsule in 2 halves, oval base

and the basal membrane extends in an irregular rim

beyond the capsule walls, the escape aperture is

asymmetrically located as in Columbella mercatoria

(Linnaeus, 1758), (4) flask shaped and oval or round at

the base, radial symmetry, the round adhesion disk is

smooth and transparent, the capsule is sculptured with

ridges as in Nitidella nitida (Lamarck, 1822), (5) brick

Page 158

THE NAUTILUS, Vol. 119, No. 4

76°0'00"W 74°0'00"W 72°0'00"W

Punta Gallinasg="85

Zz Ke Zz

8 = S}

f=) = (=)

w *_/N

2 2

° i)

S S|

2 =

E-159..~

Puerto®

Escondido

Z COLOMBIA 5025 0 50 1002

S km Ss

S 76°0'00"W 74°0'00"W 72°0'00"W

Figure 1.

Colombian Caribbean.

Map of the collecting localities in the

shaped egg capsules, escape aperture is absent as in

Anachis sp. and (6) oval shaped standing on a peduncle

as in Columbella tryngas Pils. A common spawning

feature among these species was that they attached the

egg capsules io hard substrates such as rocks, stems and

algae (Sargassum), aquarium walls and hydroid stalks.

deter spawning strategy found in the family involves

attaching the egg capsules to the shells of * conspecific

living adults, such is the case of Mazatlania consentini,

Strombina pumilio, S. francesae and Bifurcium bicana-

liferum (Penchaszadeh et al., 1983; Cipriani and

Penchaszadeh, 1993; Fortunato et al., 1998). There is

no report in the literature for the Columbellid family of

females using the shells of other living species as

a substrate for egg laying.

In this paper, we describe the egg capsules and

embryos of Amphissa sp. (Amphissa H. and Adams,

1853) and Cosmioconcha sp. (Cosmioconcha Dall, 1913)

two apparently yet undescribed species from the Colom-

bian Caribbean continental shelf. We also report on the

number of egg capsules spawned by individual females

and some reproductive parameters such as number and

size of embryos contained in the egg capsules.

MATERIALS AND METHODS

One specimen of Amphissa sp. and three specimens of

Cosmioconcha sp. were collected during 2001 by the

Colombian ship B/I ANcon during an expedition to the

Colombian Caribbean shelf (Figure 1). The specimen of

Amphissa sp. was collected at Puerto Escondido, station

E-159 at 158 m in muddy bottom (9°17' N, 76°26’ W),.

The three specimens of Cosmioconcha sp. were

collected at Punta Gallinas, station E-85 at 22 m depth

in muddy-sand bottom (12°27' N, 71°41’ W). Samples

were obtained by trawling for 10 minutes with a trawling

net type V measuring 12.7 m in total length and with

a mesh size of 10 mm (Marinovich Trawl Co, Inc, USA).

Samples were washed and separated in a 2 mm mesh

seine, observed alive and preserved in ethanol 70%. The

material of both species is deposited at the Museo de

Historia Natural Marina de Colombia (MHNMC),

INVEMAR, catalog numbers INV MOL3746 for Am-

phissa sp. and INV MOL5396 for Cosmioconcha sp.

The following reproductive aspects were studied: (1)

number and size of egg capsules attached to each specimen,

(2) number and size of embryos within each capsule and

(3) observation of the different stages of development.

RESULTS

The specimen of Amphissa sp. (Figure 2) measured

12.5 mm and5 mm in shell length and width respectively;

the protoconch of this species measured 480 um in length

and had two whorls. A seemingly related species is

A. acuminata (Smith, 1915) ont the southern West

Atlantic. Specimens of Cosmioconcha sp. (Figure 3)

measured between 11 and 12 mm in shell length and

between 4.5 and 5 mm in shell width: the protoconch

measured 570 um in length and had 3—3'% whorls. The

taxonomic status of this species has to be confirmed.

A total of 24 egg capsules were attached to the shell of

Amphissa sp. (Figure 4); these were arranged in one

layer and covered most of the shell. The three specimens

of Cosmioconcha sp. were completely covered by egg

capsules in several layers that added 3 mm to the size ‘of

the shell (from 5 to 8 mm) (Figure 5). The egg capsules

of the internal layers had no embryos aie ‘had their

escape apertures open; the closed ae containing

embryos were limited to the external layer (Table 1).

The egg capsules of Amphissa sp. (Figure 6) were

translucent, dome-shaped, with an oval escape aperture

located on the center of the dome top. The capsule

surface was smooth with no ridges nor sutures. They

were attached to the shell by an oval basal membrane

that was surrounded by a thin, irregular flange. Egg

capsules measured around 2mm in diameter and

contained around 6 gastrulae embryos in each capsule.

The egg capsules of ‘Cosioroicha sp. (Figure 7) were

very similar to those of Amphissa sp. but smaller,

measuring around 1mm in diameter and containing

around 25 embryos in each capsule at the three

developmental stages examined: eggs, trochophore, and

veliger larvae (T Table 2). No nurse eggs were observed.

The open egg capsules were filled with debris composed

of organic matter and muddy sediment. Between the egg

capsules, within the empty spaces, several specimens of

the genus Turbonilla were found.

All egg capsules of Amphissa sp. contained embryos in

the gastrulae stage in a number of six per capsule. The

gastrulae were yellow, elongated, measuring 750-

90 0 um in length and 600-660 tm in width (T. Table 3).

Each ege capsule of Cosmioconcha sp. contained

embryos in the same stage of development; however,

three different stages were identified in the spawn:

uncleaved eggs, trochophore and veliger larvae (Table 3)

P. Miloslavich et al., 2005 Page 159

Figures 2-5. Shells and attached egg capsules. 2. Adult shell of Amphissa sp. and detail of protoconch. Scale bar = 2 mm, detail =

200 um. 3. Adult shell of Cosmioconcha sp and detail of protoconch Scale bar = 2 mm, detail = 100 tm. 4. Adult shell of Amphissa

sp. covered by attached egg capsules. Scale bar = 2 mm. 5. Adult shell of Cosmioconcha sp. covered by attached egg capsules. Scale

bar = 2 mm. All photos taken from preserved material.

Empty embryonic shells were observed in a few capsules

Table 1. Cosmioconcha sp. Number of layers and summary of with embryos at the veliger stage. The uncleaved eggs

the attached egg capsules for each of the three specimens ;

measured 150 um in diameter and the trochophore

collected (ND = not determined HM |

measured 165-212 um in leneth. The veliger had an

operculum and was characterized by a transparent,

Specimen . ; ; oO nad xen

fragile shell measuring 188-219 um in length and 156

i 5] ; ; 194 um in width. The velum was small, measuring 144-

Teal ie i meee ee ee tee 494 219 um across from one lobe to the other and 31-94 um

otal number of attached capsules 537 775 A é : a ; ;

Total number of capsules in external laver ND 217 157 in height; the cilia of this velum were very small

Number of open empty capsules , 447 623 355 measuring 6-13 pm

Number of closed empty capsules 3 0) |

Number of closed capsules containing S6 152 108 WATT

ated : DISCUSSION

Sumber ope ‘ 5 0 é o ( .

Number of open c ipsules containing | 0 ) Amphissa sp aetna heen previously reported in the

mat

mbDryvos i 7

Colombian Caribbean; the sediment at the locality

Page 160

THE NAUTILUS, Vol. 119, No. 4

Figures 6-7.

500° ttm,

6. Egg capsule of

7. Ege capsule of

500 um. All photos taken from preserved material.

Amphissa sp. Scale bar =

Cosmioconcha sp. Scale bar

where the spe cimen was collected is muddy, suggesting

a broader habitat distribution for the s The shell

of this — resembles somewhat that of A. acuminata

spe cies.

(Smith, 1915), a southwestern Atlantic pe (Costa,

2005), but is not as slender and the whorls are

more rounded: the teleoconch consists of six rounded

whorls and the sculpture of uniform spiral lirae and

fine axial ribs, forming small rounded nodules at the

intersections; aperture narrow, suboval; outer lip a lit-

tle thickened exteriorly, with about five slender, short

lirae within. For a detailed description of A. acuminata,

see Simone and Leme (2001) and Costa (2005). The

living species apparently most closely related to Cosmio-

eonieha sp. is Costoanachis helenae Costa, 1983, whose

distribution range is apparently restricted to the

tropical sector of the Brazilian coast, from Amapa to

Rio de Janeiro, in depths ranging from 8 to 100 m (Rios,

1994; Costa, 2005). We compared the egg capsules of

Cosmioconcha sp. to those of C. helenae collected off

Boipeba, an island on coast of the Brazilian state of

Bahia (deposited at the Museu Nacional do Rio Janeiro,

MNRJ 1029). The specimen measured 11.9 mm_ in

length and 5.2 mm in width. Several spawning events

had occurred on this shell since there were two layers of

egg capsules, the external with about 30 and the internal

with about 15 capsules, a number considerably low-

er than the number of capsules found in our species

Table 2). The capsules were

very similar in shape to those of Cosmioconcha sp. but

their size was large rr; the “y me sasured 1.4 mm in length

and 1.3mm in width, and the aperture measur red

0.6 mm in length and 0.5 mm in width (n= 10 capsules

measured),

of Cosmioconcha (see

Cosmioconcha sp. was previously reported from the

Colombian Caribbean as Anachis cf. fraudans Jung,

1969, a very closely related form from the Miocene-

Pliocene of Trinidad (Diaz and Puyana, 1994). It also

resembles C. helenae (Costa, 1983) in shell size and

form, as well as in the number and form of embryonic

whorls. However, the latter has a more slender spire, the

upper half of its last whorl is sculptured with 11-18 axial

ribs and its shell color is pale brown with white spiral

bands (Costa, 2005). The last whorl of Cosmioconcha sp.

lacks axial ribs and the shell is pale yellow under the dark

brown periostracum. Costoanachis helenae has so far

been only recorded from the central Brazilian coast,

from Amapa to Rio de Janeiro (Rios, 1994; Costa, 2005).

The re productive strategy of spi awning or atte aching the

egg capsules to the adult shell of conspecifics is

comparable to the spawning of other Caribbean species

1983; Cipriani

However, it was not reported

found in Venezuela, (Penchaszadeh et al.,

and Penchaszadeh, 1993).

Table 2. Summary of the egg c apsule characteristics of Amphissa sp. and Cosmioconcha sp. Values represent mean + SD, numbers

in parenthesis indicate range (n = number of egg capsules measured)

Species Capsule Capsule Capsule Aperture Aperture Embryos

Shell length (mm length (mm) width (mm) height (mm) length (mm) width (mm) per capsule

\mphissa sp 22+0.1 2.0+0.01 0.8+0.2 0.90+0.04 O0.74£0.07 5.9+1.2 (4-8)

2.5 (2.0-2.4) (1.9-2.1) (0.6-1.0) (0.7S—0.93) (0.60—0.90) n=l)

n=18 n=18 n=18 n=1] n=11 (Gastrulae)

Cosmioconcha sp LI5*+0.15 LOL+#0.12 0.56£0.06 ).35+0.05 0.27+0.03 25.0+2.9 (19-31)

1-12 (O.75-1.5) (0.75-1.47) (O.24—0.50) (0.27-0.54) (0.02—0.02) n=23

n=63 n=63 n=62 n=62 n=62 (All stages)

P. Miloslavich et al., 2005

Page 16]

—

Table 3. Size of embryos of Amphissa sp. and Cosmioconcha sp. during intracapsular development. Values represent mean + SD

and are reported in Um, numbers in parenthesis indicate range (n

Species Egg Gastrula

Trochophore

= number of embryos measured).

Veliger Protoconch (in adult)

S40 474 (750-900)

n=6

Cosmioconcha sp. 150+0 —

n=16

Amphissa sp. —_—

189+19 (165-213)

n=4 n=7

500-600

200+ 14 (188-219) 570

for any of the 10 columbellid species studied by Bandel

(1974) in the Santa Marta region, which atti wched the egg

capsules to other hard substrates such as rocks, Sar assum

plants, stalks of hydroids and, when in captivity, to the

glass walls of the aquarium. There are no records in the

literature of columbellids spawning on the shells of other

species, a strategy that has been ‘observed in the sandy

beach gastropod Olivancillaria deshayesiana Ducros,

1857, from Argentina, which spawns the egg capsules on

the adult shells of Buccinanops monilifer Kiener, 1834,

and Buccinanops duartei Klappenbach, 1961, either

directly on the adult shell or on top of the egg capsules

of these species, which are also attached to the shell of

conspecific individuals (Borzone, 1985). The fact that

several lavers of egg capsules are found, as well as different

stages of embrvonic dev elopment and empty egg capsules,

on Cosmioconcha sp. suggests that several different events

of oviposition have occurred using this specimen as

substrate. Penchaszadeh et al. (1983) reported that

females of Mazatlania consentini (as M. aciculata) attach

their egg capsules to the shells of other individuals of the

population with shells larger than 9 mm, apparently

exclusively males, as shoven by histological sections of

the gonads of bearing individuals ( (Penchaszadeh, 1981).

Fortunato et al. (1998) stated that the egg masses of

Bifurcium bicanaliferum usually have several layers as

well, whereas those of the Venezuelan Caribbean

Strombina pumilio, S. francesae, and M. consentini have

only one layer of egg capsules attached to the shell as we

observed in Amphissa sp. Regarding this remarkable

difference in the number of egg capsules that a single

animal is bearing, we hypothesize that the egg capsules of

species that bear few of them on the shell arranged on

a single layer are easily detachable since they are usually

individually attached, as is the case in M. aciculata. In this

species, up to 20 egg capsules can be observed at the same

time on one single adult shell; these have embryos at

different stages of development. which means that not all

egg capst ules are spawned simultaneously. Once the

veliger larvae hatch, the empty capsule usually falls off

the shell, sometimes leaving only the basal membrane

attached to the shell, and new capsules are spawned

(Penchaszadeh, 1981). On the other hand, egg capsules of

species with several layers are more resistant since they are

attached not only to the adult shell by the basal membrane,

but also form a very compact structure of egg capsules

attached between them by lateral membranes (B.

bicanaliferum, Cosmioconcha sp.). As pointed out by

Pechenik (1986), gastropod egg capsules are structurally

and chemically complex, the different layers are composed

in general among and other components by protein and

carbohy drates (Bayne, 1968; Miloslavich, 1996: Rawlings,

1999). Hunt (1966) also reported that since the c -apsule

walls are extremely resistant to chemical treatment, the

protein and carbohy drate components should be strongly

linked in a glycoprotein complex and thus stabilized by

sclerotization. In this way, the bond between two egg

capsules is much stronger than the bond between an eg

capsule and a shell.

The description of the egg capsule of Amphissa sp. is

similar to the description i -d by Bandel (1974) for

Zafrona pulchella (cited as Anac his pulchella) in its

dome shape. However, ne. oe of Amphissa sp. lacks

the division into lower and upper parts separated from

each other by one or a few concentric rings, it does not

have a suture that continues the axis of the escape

aperture and the membrane is not striped nor wrinkled.

On the contrary, the surface is very smooth and only very

thin imperceptible lines are visible. A comparable

feature among both species is that the number of

embryos per ae was very similar and that no nurse

eggs nor cannibalism seem to occur.

The egg capsules of Cosmioconcha sp. also fit in the

first category of egg capsule morphology described by

Bandel (1974), a simple dome attached by a rounded

base. It is very similar to the capsule of Amphissa sp. but

half its size in length, width and height. This capsule

shape and general morphology are also comparable to

the two species of the Strombina group from the

Caribbean described by Cipriani and Penchaszadeh

(1993), which have about the same size than the capsules

of Amphissa sp., and to B. canaliferum trom the Pacific

coast of Panama described by Fortunato et al. (1998),

which has about the same size than the capsule of

Cosmioconcha sp.

No nurse eggs seem to be present in the two species

studied. In comparison to other columbellid gastropods,

the reproductive strategy of Amphissa sp. is similar to

that of S. francesae and S. ounaliss large egg capsules

(more than 2 mm in length), large eggs (more than

500 um) and few embryos (about 4 to 8) that hatch as

large crawling juveniles (larger than 0.9 mm). We did

not observe the hatching stage of Amphissa sp., but the

large size of the gastrula (about $40 um) indicates that it

Page 162

comes from a large egg which would most probably lead

to direct dev elopment as in the two species of Strombina

studied by Cipriani and Penchaszadeh (1993). On the

other hand, the reproductive strategy of Cosmioconcha

sp. is similar to that of Bifurcium bic -analife rum, Anachis

sparsa, and Cosmioconcha sertulariarum (d’Orbigny,

1839): small egg capsules (about 1 mm in length), small

eggs (about 140-200 ttm) and many embryos (more than

20) that hatch as small veliger larvae (about 300 Um)

(Fortunato et al., 1998: Marcus and Marcus, 1962). Late

cannibalism among sibling embryos inferred from the

presence of empty shells "inside a few egg capsules of

Cosmioconcha sp. has also been reported in B.

bicanaliferum by Fortunato et al. (1998). Although late

cannibalism might take place in the egg capsules of

Cosmioconcha sp., it is very unlikely that this limited

extraembryonic food source ‘will allow the larva to grow

from 200 ttm to 600 f1m, which is the protoconch size

found in the adult shell. It is most likely that

Cosmioconcha sp. will hatch as a veliger larva spending

some time in the plankton as is also the case of B.

bicanaliferum, which grows from 1.5 whorls to 2.5

whorls in the planktonic period (Fortunato et al., 1998).

In the case of Amphissa sp., we can conclude from the

size of its protoconch that this species must hatch as

a crawling juvenile since the gastrula stage is already as

large as the protoconch, a_ situation similar to what

happens with the two Strombina species from the

Venezuelan Caribbean studied by Cipriani and Pench-

aszadeh (1993).

We hope that in a near future, and thanks to the

collection efforts carried out by INVEMAR in the

southern Caribbean, more specimens will be collected.

This will not only allow for the proper description of

these species th rough use of traditional characters such

as those from aaule shell, anatomy, and radular

morphology, but may also help increase the number of

observations on egg capsules and the development of

other embryonic stages.

ACKNOWLEDGMENTS

This work was partially supported by a Decanato de

Investigacion y Des sarrollo, Universidad Simon Bolivar

grant to the Grupo de Ciencias Marinas, by a FONACIT

(S1-2001-000764) grant to the first author. Material

collecting was possible through the “Macrofauna I and

I” projects funded by COLCIENCIAS and INVEMAR.

We are especially indebted to Paulo Marcio Costa,

Departamento de Invertebrados, Malacologia, Museu

Nacional, Rio de Janeiro, and Marta DeMaintenon,

University of Hawaii at Hilo for helping with the

problematic taxonomic status of our material. Paulo

Marcio Costa also provided us with one reproductive

specimen of Costoananchis helenae from Brazil for

comparative purposes. We also wish to thank Pablo

Penchaszadeh, Facultad de Ciencias Exactas y Natur-

ales, Universidad de Buenos Aires, for our fruitful

THE NAUTILUS, Vol. 119, No. 4

discussions in the subject and José L. Garcia Rondon,

Artis Publishers, for his invaluable help with the

photographs.

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lusca: Gastropoda) de una playa arenosa expuesta del sur

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THE NAUTILUS 119(4):164—168, 2005

Page 164

Pterorytis pacanana new species (Gastropoda: Muricidae):

circumstantial evidence for late Pliocene El Nifio events in

southern Peru

Thomas J. DeVries!

Burke Museum of Natural History and Culture

University of Washington

Seattle, WA 98195 USA

ABSTRACT

Pterorytis pacanana new species was discovered in upper

Pliocene bioclastic sandstone near Chala, southern Peru.

Although it was found farther south than other Pterorytis

species from the eastern equatorial Pacific Ocean, P. pacanana

most resembles P. roxaneae Petuch, 1994, from the Pliocene

Pinecrest beds of Florida. Its presence among endemic

late Pliocene cool-water mollusks from high-energy shoreface

paleoenvironments is thermally anomalous in the same sense

as the rare occurrence of other species from northern Peru

and Ecuador in upper Pliocene and Pleistocene strata

from southern Peru. By analogy with modern thermally

anomalous mollusks that appear episodic lly off the coast of

southern Peru and Chile, the Pliocene and Pleistocene

examples are circumstantial evidence for the occasional south-

ward incursion of warm equatorial waters during former E]

Nino events.

INTRODUCTION

Pliocene ee in southern Peru are noted for a variety

of muricid gastropods, most belonging to genera still

represented ‘by extant species (e.g., Ac anthina Fischer

von Waldheim, 1807; Chorus Gray, 1847; Concholepas

Lamarck, 1801; Crassilabrum Jousseaume, 1880; Mur-

egina Vermeij, 1998; Stramonita Schumacher, 1517;

Xanthochorus Fischer, 1854), but some from genera

entirely or locally extinct (e.g., Herminespina DeVries

and Vermeij, 1997; Trophon, Montfort, IS10) (DeVries,

1995, 1997, 2000, 2003, in press a, in press b: DeVries

and Vermeij, 1997).

muricids are not exceedingly difficult to find.

Specimens of most Pliocene

A well-preserved muricid from Pliocene bioclastic

sandstone south of Chala is, literally, the rare exception.

None have been found other than a single shell plucked

from a roadeut of the Panamerican Highway that

overlooks Playa Huacllaco. The lamellar fimbriate

varices of the Huacllaco specimen invite comparison

with ocenebrines from Ecuador, the southeastern

Mailing address: P.O. Box 13061, Burton, WA 98013 USA

United States, and the northwestern Pacific Ocean. Its

lone appearance in a collection of endemic cool-water

muricids is another example of a thermally anomalous

molluscan species encountered in upper Pliocene and

Pleistocene beds of southern Peru (Muizon and

DeVries, 1985; DeVries, 1986; Ortlieb et al., 1990). A

reasonable hypothesis is that these equatorial species

were introduced to higher austral latitudes by warm-

water incursions during the Pliocene and Pleistocene

comparable to modern incursions that carry Panamic

molluscan larvae poleward during El Nifio events

(DeVries, 1988; Arntz and Tarazona, 1990; Paredes et

al., 1998).

GEOLOGY

Bioclastic sandstone and gravel and balanid coquina

overlie igneous basement in roadcuts along a series of

sweeping curves in the Panamerican Highw ay where it

descends towards the beach at Play a Huacllaco

(Figure 1). The sediments represent high- energy fore-

shore and _ intertidal paleoenvironments that once

flanked steep cliffs. The section, previously published

fe)

Playa

Huacllaco

Figure I.

(DV 1628).

Type locality of Plerorytis pacanana new species

T. J. DeVries, 2005

Page 165

Figures 2-8. Pterorytis species. 2, 3, 5. salve pacanana new species. Upper Pliocene. Holotype, UWBM 97772, length =

30. 9 mm, width = 19.6 mm. 2. Apertural view.

3. Abapertural view. 5. Oblique view of spire. 4, 6-8. Prer rorytis roxaneae Petuch,

1994. Upper Pliocene. Holotype, Florida Museum of Natural History, Gainesville, UF 66254, length = =

3.3 mm, width = 21.5 min.

4. Lateral view. 6. Apertural view. 7. Abapertural view. 8. Oblique view of spire.

by DeVries (2003), consists of four stratigraphic units.

The ocenebrine specimen was found at the base of Unit

IH, just below strata where rounded clasts of blackened

andesite first appear in great numbers and below beds

where specimens of Concholepas and Acanthina acquire

their modern form (DeVries, 2000, 2003).

The age of the Huacllaco beds is bracketed by basal

beds with specimens of Concholepas nodosa Hupé,

1854, Acanthina triangularis DeVries, 2003, and Her-

minespina mirabilis (Mericke. 1896), which collectively

indicate an early late Pliocene age (DeVries and

Frassinetti, 2003), and the uppermost and oldest of

several marine terraces, whose elevation and largely

extant taxa suggest a latest Pliocene age (Muizon and

DeVries, 1985).

MATERIALS AND METHODS

The specimen from Peru described in this study was

found by the author. Dimensions affected by breakage

are enclosed by parentheses. The holotype is deposited

at the University of Washington’s Burke Museum of

Natural History and Culture in Seattle, Washington

(UWBM).

SYSTEMATICS

Family Muricidae Rafinesque, 1S15

Subfamily Ocenebrinae Cossmann, 1903

Genus Pterorytis Conrad, 1862

Subgenus Pterorytis sensu stricto

monotypy.

Pterort ae yaeaane new species

(Figures 2, 3, 5)

Diagnosis: Shell small; texture waxy, shell thin. Five

lamellar, fimbriate varices; intervarical nodes absent.

Three spiral cords; shoulder spiral cord strong. Sutural

platform horizontal. Labral tooth absent.

Description: Shell 30.9 mm long (first teleoconch

whorls missing), quadrate in profile, very thin, with

waxy texture. Spire estimated to be about 40 percent of

shell length. Siphonal canal about 20-25 percent of shell

length. Protoconch and earliest teleoconch whorls

missing. Upper sides of whorls planar, vertical; base of

body whorl sharply constricted. Shoulder orthogonally

Page 166

THE NAUTILUS, Vol. 119, No. 4

angulate, sutures deeply impressed; sutural platform

hareonte! to slightly concave. Three last whorls with five

lamellar varices "extending from fasciolar ridge to suture,

each varix joining across suture with varix of previous

whorl. Lamellae broader basally, narrower adapically,

extended adapically at shoulder but not spinose;

fimbriate on adapertural face; erect or recurved

adaperturally except apertural lamellae, which is weakly

recurved ae rturally. Intervarical nodes absent. Spiral

sculpture of prominent but ill-defined rounded primary

cord at shoulder angulation and two additional broad low

spiral cords anteriorly, almost obsolete. Additional broad

low secondary cords barely visible adjacent to adaper-

tural face of varices. Lamellar fimbria slightly extended

at intersection with primary spiral cords. Aperture with

inverted tear-drop shape. Parietal rib, anal sulcus absent;

parietal area unexcavated. Columella smooth, inner lip

very weakly concave, adherent anteriorly. Outer lip

without dentition on inner edge. Labral tooth absent.

Siphonal canal open, slightly recurved to right. Siphonal

fasciole strongly arched, without rostrae. Peewde: umbi-

licus narrow, extending adapically beyond siphonal

canal.

Holotype: UWBM 97772, DV 1628-5, lower Upper

Pliocene, length (30.9) mm, width 19.6 mm.

Type Locality: —Roadcut along the Panamerican High-

way, 10 km south of Chala, on a winding descent from

a 200 m elevation terrace towards Playa Huacllaco,

locality DV 1628, north side of first sweeping outside

curve from base of outcrop, south side of road, near

outcrop of igneous basement rock (Figure 1), 15°52" S,

74°10’ W (Chala 1:100,000 quadrangle).

Occurrence: Upper Pliocene, southern Peru.

Etymology: ‘Paca, Quechua for ‘high, and ‘nan,

Quechua for ‘path, referring to the horizontal to

concave sutural platform bordered above and below by

vertical walls of the whorls.

Remarks: Pterorytis or Ocinebrellus Jousseaume,

1880, is the most appropriate genus in which to place

the Huacllaco specimen, which has five varices on at

least the last three whorls, consistent with the three to

nine varices observed on fossil Atlantic species of

Pterorytis (Emerson, 1959; Vermeij, 2001), the four to

five varices on the modern eastern Pacific Pterorytis

hamatus (Hinds, 1844) (Emerson, 1985), and three to

twelve varices on specimens of Ocinebrellus (Amano and

Vermeij, 199Sa). The ocenebrine genera Ceratostoma

Herrmannsen, 1846, Pteropurpura Jousseaume, 1580,

and Mic rorhytis Emerson, 1959, in contrast, have three

prominent lamellar varices on all or at least the last two

whorls (Vermeij and Vokes, 1997; Amano and Vermeij,

1998a, 199Sb). Specimens of Ceratostoma, Pteropur-

pura, Microrhytis, and some Ocenebra Gray, 1547, also

have intervarical nodes, which are absent on the

Huacllaco specimen and specimens of Pterorytis and

Ocinebrellus.

The absence of a closed siphonal canal does not argue

against assignment of the single Huacllaco specimen to

Pterorytis or Ocinebrellus, since juvenile and some adult

specimens of the two genera may have an open siphonal

canal. The Peruvian specimen, itself partly abraded,

might be a juvenile specimen or have a broken siphonal

canal.

Distinguishing Pterorytis from Ocinebrellus for place-

ment of the "Huacllace specimen is problematic.

Ocinebrellus has four primary spiral cords on the swollen

portion of the body whorl, posterior to the labral tooth, if

present (Vermeij and Vokes, 1997; Amano and Vermeij,

1998a). Pterorytis and the Huacllaco specimen have

three primary spiral cords, with an additional weak

primary spiral between the shoulder and suture. On

smoother specimens of both genera the number of

primary spiral cords can be diff sulk to enumerate, and

on strongly sculptured specimens the distinction be-

tween primary and secondary spiral cords is unclear if

the ontogeny is unknown.

Other features are equally unsatisfactory for distin-

guishing the two genera. The reflection of lamellar

varices is not reliable, since the lamellae may be

adaperturally reflected, or not, in Ocinebrellus (Amano

and Vermeij, 1998a), abaperturally reflected or erect in

Pterorytis (Vermeij and Vokes, 1997), and erect or

reflected in either direction on the Huacllaco specimen.

The angularity of the shoulder also fails as a distinguish-

ing character. Most specimens of Ocinebrellus have an

angulate shoulder and a horizontal sutural platform (as is

ee case for the Huacllaco specimen), but some have

rounded shoulders. Most specimens of Pterorytis have

broad, planar, steeply sloping sutural platforms, but

some have narrower, less inclined sutural platforms

more like those of typical Ocinebrellus and the

Huacllaco specimen.

The presence or absence of a labral tooth is not

diagnostic. Some ocenebrines acquire a labral tooth only

in adulthood; the Huacllaco specimen might be a juve-

nile. Species of Pterorytis that normally have a tooth (P.

umbrifer Conrad, 1832; P. fluviana Dall, 1890) have

specimens lacking a tooth. Specimens of P. roxaneae

Petuch, 1994, never have a labral tooth (Vermeij and

Vokes, 1997). Species of Ocinebrellus (sensu Amano and

Vermeij, 199Sa) that normally lack a labral tooth have

specimens with a tooth. (In the more restrictive

classification of Houart and Sirenko (2003), species of

Ocinebrellus do not have a labral tooth.)

Taking into account this taxonomic ambiguity, the

Huacllaco specimen is assigned to Pterorytis. a cies of

Pterorytis (P. umbrifer, P. fluv iana, P. roxaneae) typically

have smooth-shelled variants such as the spe cimen from

Huacllaco. Most specimens of Pterorytis, like the

Huacllaco specimen, feature a prominent primary spiral

cord at the shoulder, but nonetheless usually lac k spines

or angulations typically seen in specimens of Ocineb-

rellus.

Pliocene Ecuadorian specimens assigned to Ocineb-

rellus by Vokes (1988) are smooth, have a prominent

T. J. DeVries, 2005

Page 167

shoulder spiral bordering an inclined sutural platform,

and have varices that are only weakly angulate; they may

be referred to Pterorytis ecuadoria (Olsson, 1964). They

have a more fusiform profile and greater number of

varices than the Huacllaco specimen.

The specimen of Pterorytis pacanana most closely

resembles specimens of the Pliocene Pterorytis (Pteror-

ytis) roxaneae Petuch, 1994, (Figures 4, 6-S) from

the Pliocene Pinecrest beds of Florida (Vermeij and

Vokes, 1997). Specimens of both species are thin with

a waxy texture, have reduced spiral sculpture, and lack

a labral tooth. Pinecrest specimens differ from the

Huacllaco specimen in having four varices, not five,

varical lamellae that are buenas adapically, not abapi-

cally, and a weak peripheral spiral cord bordering

a sloping sutural platform, rather than the strongly

defined horizontal sutural seeee of P. pacanana.

Contrary to Vermeij and Vokes (1997), the holotype of P.

roxaneae is neither excessively worn nor lacking the

labral varix.

DISCUSSION

Pterorytis pacanana is the third or fourth species of

Pterorytis recognized in the eastern Pacific Ocean, after

P. ecuadoria (Pliocene, Ecuador), P. hamatus (Recent,

northern Peru [Alamo and Valdivieso, 1997], a species

with a protoconch unlike that of Ocinebrellus or any

other ocenebrine [R. Houart, personal communication,

2005]), and an unnamed Recent ocenebrine from

northern Peru (Radwin and D’Attilio, 1976). These taxa

define an eastern tropical Pacific complement to a clade

of Pterorytis species from the southeastern United States

whose oldest members date to the late Miocene.

Ocinebrellus, which may be endemic to the northwest-

em Pacific (Amano and Vermeij, 1998a; Houart and

Sirenko, 2003), extends back to the Early or Middle

Miocene. Ocinebrellus seems morphologically more

similar to Pterorytis than other ocenebrine clades, but

the trail of fossil species that might lead from Japanese

Ocinebrellus to Panama and the Caribbean and beyond

to Peruvian and Floridian Pterorytis has yet to be

discovered (Amano and Vermeij, 1998a).

The specimen of Pterorytis pacanana from 16°S is

remarkable for its singular occurrence and equatorial

affinity. Associated taxa “(Table 1) are entirely endemic or

cool-water species that became prevalent after a pro-

vincial mid-Pliocene extinction that coincided with

a global cooling event (Dowsett et al., 1996; DeVries,

2001). The rare appearance of Pterorytis in southern

Peru resembles that of a mangrove bivalve, Anadara cf.

A. grandis (Broderip and Sow erby, 1829), whose speci-

mens are found in small numbers in uppermost Pliocene

beds at 15°30’ S with cool-water species (Muizon and

DeVries, 1985: DeVries, 1986). In a more recent

example, specimens of Chione broggi (Pilsbry and

Olsson. 1943) and Cerithium stercusmuscarum Valenci-

ennes, 1833, both living today only as far south as the

Table 1. Molluscan species associated with Pterorytis paca-

nana, new species, which was found at the base of Unit II of

the Upper Pliocene beds above Playa Huacllaco, Peru

= Extinct).

Unit ILIV (younger)

Acanthina unicornis (Bruguiére, 1789)

Cancellaria buccinoides Sowerby, 1832

Choromytilus chorus (Molina, 1782)

Chorus grandis (Philippi, 1887) / C. giganteus (Lesson,

1546)

Concholepas camerata DeVries, 2000

Concholepas concholepas (Bruguiére, 1789)

Crassilabrum crassilabrum (Sowerby, 1834)

Crepidula dilatata (Lamarck, 1822)

Eurhomalea lenticularis (Sowerby, 1835)

Glycymeris ovata (Broderip, 1843)

Mesodesma donacium (Lamarck, 1818)

Mulinia edulis (King, 1831)

Oliva peruviana Lamarck, 1811

Piscoacritia new species

Prisogaster niger (Wood, 1828)

Sinum cymba (Menke, 1828)

Xanthochorus cassidiformis (Blainville, 1832)

Unit I+II (older)

Acanthina triangularis DeVries, 2003

Acmaeids

* Chlamys cf. C. vidali (Philippi, 1887)

Choromytilus chorus (Molina, 1782)

Chorus grandis (Philippi, 1887)

Concholepas camerata DeVries, 2000

Concholepas nodosa (Méricke, 1896)

Fissurella spp.

Herminespina mirabilis (Moricke, 1896)

Lithophaga sp.

Piscoacritia collapsa DeVries and Hess, 2004

Stramonita new species

Tegula (Chlorostoma) new species

Xanthochorus buxeus (Broderip, 1833)

Xanthochorus new species

Sechura coastline of northern Peru (5°S) (Alamo and

Valdivieso, 1997), are occasionally found in upper

Pleistocene terrace deposits near San Juan de Marcona

(15°20'S), Saeaco (15 ae and Ilo (17°40'S) (DeVries,

1986, 1988: Ortlieb et al., 1990). Ortlieb et al. (1990)

proposed that the late Pleistocene thermally anomalous

species were introduced southward from equatorial

latitudes during El Niho events. Several such immigra-

tions of equatorial mollusks have been documented

during modern El Ninos events (Arntz and Tarazona,

1990: Paredes et al., 1998). The rare Pliocene equatorial

species in southern Peru were probably introduced in

the same manner.

ACKNOWLEDGMENTS

I would like to thank Brandur Karlsson of Reykjavik,

Iceland, for his assistance in the field and Greg Herbert

(University of South Florida) for helpful discussions on

Page 168

ocenebrine taxonomy. J. Vermeij and R. Houart

provided helpful tee in their reviews of the

manuscript.

LITERATURE CITED

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moluscos marinos del Pert. Instituto del Mar del Peru,

Callao, 183 pp.

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of the genus Ocinebrellus Sea Re Muricidae) in

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Amano, K. and G. J. Vermeij. 1998b. Origin and biogeographic

history of Ceratostoma (Gastropoda: Muricidae). Venus

57: » 209— 993.

Arntz, W. A. Fr J. Tarazona. 1990. Effects of El Nifio 1982-

1983 on benthos, fish, and fisheries off the South

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ecological consequences of the 1982-1983 El Nino-

Southern Oscillation. Elsevier: Amsterdam, pp. 323-360.

DeVries, T. J. 1986. The geology si paleontology of tablazos

in northwest Peru. Doctoral dissertation, The Ohio State

University: Columbus, 964 pp.

DeVries, T. J. 1988. A review of geological evidence for ancient

El Nino activity in Peru. Journal of Geophysical Research

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DeVries, T. J. 1995. Cone tholepas Lamarck, 1801 (Neogastro-

poda: Muricoidea): A Neogene genus native to South

America. The Veliger 38: 254-297.

DeVries, T. J. 1997. A review of the genus Chorus Gray, 1847

(Gastropode i: Muricidae) from western South America.

Tulane Studies in Geology and Paleontology 30; 125-147.

DeVries, T. J. 2000. Two new Neogene species and the

evolution of labral teeth in — oe Lamarck, 1801

(Neogastropoda: Muricoidea). The Veliger 43: 43-50,

DeVries, T. J. 2001. Contr isting patterns of Pliocene and

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North and South America. Geological Society of America,

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DeVries, T. J. 2003. Ac cantina Fischer von Waldheim, 1807

(Gastropoda: Muricidae), an ocenebrine genus endemic to

South America. The see 46; 332-350.

DeVries, T. J. a. (In press). The Late Cenozoic history of

Xanthochorus Fischer, 1884 (Gastropoda: Muricidae) in

western South America. The Veliger.

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Peru: Seven new species and a biogeographic summary.

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DeVries, T. J. and C. D. Frassinetti. 2003. Range extensions

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mollusks Peay in Peru. Boletin del Museo de Historia

Natural, Chile 52; 141-157.

DeVries, T. J. and G. J. Vermeij. 1997. Herminespina: New

genus of Neogene muricid gastropod from Peru and

Chile. Journal of Paleontology 71: 610-615.

Dowsett, H., J. Barron and R. Poore. 1996. Middle Pliocene

sea surface temperatures: a global reconstruction. Marine

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West American species assigned to the Neogene paciphile

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and F, Cardozo. 1998. Presencia de moluscos tropicales

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Biologia (Universidad Nacional Mayor De San Marcos)

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World. Stantord University Press, Stanford, 254 pp.

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origins and fates of gastropods with a labral tooth:

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the western Atlantic region. Part XIE - the subfamily

Ocenebrinae (in part). Tulane Studies in Geology and

Paleontology 29: 69-115.

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in Geology and Paleontology 21: 1-50.

THE NAUTILUS 119(4):169-173, 2005

Page 169

A new species of Falsimargarita (Gastropoda: Vetigastropoda:

Trochidae) from the South Atlantic Ocean

Eliézer de Carvalho Rios

Museu Oceanografico

Fundagao Universidade de Rio Grande

Cx. Postal 379

96200-970 Rio Grande

BRAZIL

Sao Paulo

Cx. Postal 42494

BRAZIL

[email protected]

04299-970 Sao Paulo

F F e l

Luiz Ricardo L. Simone

Museu de Zoologia da Universidade de

ABSTRACT

A new trochid species, Falsimargarita stephaniae, is described

from about 1200 m depth off the Malvinas (Falkland) Islands,

South Atlantic Ocean. The new species is distinguished from

the most similar congeneric ones by its excee -dingly large spiral

cords located only along the periphery of its chouldeved whorls.

A re-hydrated specimen allowed for the description of some

details of the anatomy of the new species, including head-foot,

buccal mass, and radula.

INTRODUCTION

Representatives of the trochid genus Falsimargarita

Powell, 1951, can be distinguished by shell diarapters

such as external iridescence, well-defined spiral whorls,

strong spiral sculpture, opened umbilicus, and thin shell

wall. The genus encompasses five species occurring in

the cold or freezing deep waters off Antarctica and “the

Magellanic region of South America. The taxon was

more recently revised by Dell (1990), who outlined the

diagnostic characters of the genus and described two

species. :

The analysis of a specimens collected by a boat deep-

fishing for king crab and tuna rev ealed the presence of

the new species. This paper is part of a larger project of

revision of western Atlantic molluscan species, which at

the moment is focused on the study of deep-sea trochids.

MATERIALS AND METHODS

A single specimen with dry soft parts was available for

study. Dry soft parts were carefully removed and re-

hy drated in physiological solution and 3—4 drops of KOH

20% for 3 h, then transferred to 70% ETOH. Only the

* Author for correspondence

head-foot was adequately extracted. The dissection was

performed with the specimen immersed under the

fixative, in a stereomicroscope. All drawings were done

under camera lucida. The radula was removed and

cleaned in a boiling solution of KOH for 1 h, then

cleaned by sonication in water. The examination was

done under a Zeiss electron microscope at the

Laboratorio de Microscopia Eletrénica of Museu de

Zoologia da Universidade de Sao Paulo.

Institutional abbreviations used in this article are:

MORG, Museu Oceanografico da Fundagao Universi-

dade de Rio Grande, Rio Grande, Brazil; MZSP, Museu

de Zoologia da Universidade de Sao Paulo, Sao Paulo,

Brazil.

SYSTEMATICS

Genus Falsimargarita Powell, 1951

Type species: Margarites gemma Smith, 1915; by

original designation, Antarctica.

Falsimargarita stephaniae new species

(Figures 1-12)

Diagnosis: Shell with broad spire, 5 prominent large

and tall spiral cords restrict to periphery; umbilicus

protected by strong plate.

Description: SHELL of medium size (16.6 mm), tro-

choid to turbiform, whitish, iridescent-gray to pale-

reddish; wall relatively thin, light. Protoconch of one

smooth, glossy whorl. Separation protoconch-teleoconch

poorly detned! Spire with 3.5 teleoconch whorls; each

whorl highly convex, relatively high and shouldered;

superior half weakly descendent, sculj stured by 6-7 low

and narrow spiral and numerous axial lee »5, both e qually

predominating; inferior half abruptly descendent, sculp-

SE SSE

THE NAUTILUS, Vol. 119, No. 4

Figures 1-5.

larger diameter 16.6 mm). 5. Operculum, outer view

tured by five strong and promiment spiral cords, two

adapical ancl one abapical cords clearly larger latter cord

coinciding with suture; smooth area bearing only growth

lines present between cords of abapical halt ol whorl

Body whorl about twice spire width, sculptured with five

strong spiral cords that continue unchanged from spire;

five spiral cords restricted to periphery area of body

vhorl: base s« ulptured with 15 spiral lines successively

and gradually broader and more spaced toward umbi-

licus. Body whorl well separated from umbilicus by

viral cord almost ral low Carlilla i mbilicus open

Falsimargarita stephaniae new species. 1-4. Shell of holotype, apertural, abapertural, apical

deep, surface with s

from

expansion of t

umbilicus \p

slightly deflected o

adapical half marke

body whorl ho cal

thin, with

sculpture

Heap-roor: Total

whorl. Head protru

head-foot volume

small pr

Snout with about “4 of

and umbilical views

imple srowth lines: a strong plate-

he inner lip separates this latter

erture rounded, ample. Inner lip

1 abapical half, somewhat thick;

cd only by thin glazed area on

lus present Outer lip rounded,

ojections corresponding to spiral

ength about “> length of last shell

ded, occupying about 3 of total

foot size,

E. C. Rios and L. R. L. Simone, 2005

age 17]

Figures 6-11. Falsimargarita stephaniae new species. 6. Buccal mass, ventral view, ventral wall opened longitudinally along

median line and deflected to expose jaws. Scale bar = 1 mm. 7-8. Left jaw plate, outer and inner views. Scale bar = 0.5 mm. 9-11.

Radula. 9. General view; 10. Detail of central and lateral teeth. Scale bars = 100 ttm. 11. Detail of lateral teeth, arrow indicating

fifth lateral tooth. Scale bar = 50 um. Abbreviations: jw, jaw; mo, mouth; od, odontophore; ra, radula; rs, radular sac

cylindrical, broad; distal surface flattened, fully covered

with small papillae: each papilla cylindrical, tip rounded:

mouth central. Tentacles long (about twice snout

length), narrow, tip rounded. Ommatophore with about

¥ of tentacle length and approximately with same width;

located just posterior to tentacles: eyes dark, on

ommatophore tips. Foot occupying about 74 of head-

foot volume. Mesopodium constituting most of foot,

outline somewhat triangular; sole flat, simple; anterior

furrow of pedal glands bordered by thick margins,

restricted anteriorly, not protruding beyond lateral

edges. Epipodium divided into two apparently symmet-

rical lateral flaps. covering entire dorsal surface of

mesopodium, from snout base to opercular pad; bearing

eight pairs of long epipodial tentacles projecting about

twice longer than epipodial width, each tentacle pro-

truding on ventral but not on dorsal epipodial surface;

each epipodial tentacle bearing papillae at ventral

surface of basal region; papillae increasing in number

and size toward middle tentacles; some epipodial

tentacular papillae bifid at tip. Opercular pad with

edges as continuation of epipodium. Columellar muscle

thick, encompassing a half whorl

OpercuLuM: Circular, horny, multispiral: nucleus

central; occupying entire shell aperture

Dicestive System: Buccal mass somewhat larger than

snout internal space. Buccal cavity having a pair of very

large, dark-brown jaw plates, outline somewhat ellipt

are. 179

Page 172

THE NAUTILUS, Vol. 119, No. 4

Falsimargarita stephaniae new species. Head-

foot, ventral to slightly lateral right view, epipodium deflected

upward. Scale bar = 1 mm. Abbreviations: ep, epipodium; et,

epipodial tentacle; fs, foot (mesopodium) sole; he, head; mo,

mouth; om, ommatophore; pg, anterior furrow of pedal glands;

sn, snout; te, cephalic tentacle.

Figure 12.

both jaws occupying most of dorsal and lateral surfaces

of buccal cavity; posterior and lateral regions of jaws low,

medial and anterior regions taller, with “projected edges;

series of small cusps present along anterior and medial

edges, each cusp pointed and well separated from each

other. Odontophore about half projected into buccal

cavity. Radular ribbon about three times odontophore

length; about half of radular ribbon projected beyond

posterior end of odontophore.

RADULA:

radular ribbon width, triangular, narrowing somewhat

abruptly, strongly curved over its own base; cutting edge

sharply pointed, margins with very slender, elongated

cusps. First to fourth lateral teeth long, slender,

narrowing gradually, curved inward; distal half bearing

edges with slender, elongated cusps; tip sharply pointed.

Rachidian and four more central lateral teeth thin,

flexible. Fiftieth lateral tooth thick, hook-like, curved

inward; base broad, thick; distal region arched, re-

sembling a thick scythe. Marginal teeth slender, tall,

about 20 pairs per row, slightly broader toward medial

region; base ruler-like, weakly curved inward; distal half

sharpening gradually, with several slender, elongated

cusps along edges about 25 pairs per tooth; tip sharply

pointed.

Holotype: MORG 49650 (shell and operculum),

MZSP 46559, diameter: 16.6 mm; height: 14.6 mm;

includes re-hydrated soft parts and radula.

Type Locality: Argentina, off Islas Malvinas (45°S

58°W), 1200 m depth (fishing boat col., x/2004, Helen

Racz leg.).

Distribution: Known only from type locality.

Etymology: The

collector's mother, Ms.

Latinized specific epithet honors the

Teodora Stefania.

Rachidian tooth encompassing about 4 of

DISCUSSION

Although it has been recognized that definitions of

generic boundaries in the Trochidae merits further

revision, we are reasonably certain of its generic

allocation of the new species, because the species

possesses the conchological attributes reported in the

Introduction for Falsimargarita.

There are two other genera also occurring in the

South Atlantic Ocean that also exhibit iridescent shells,

a character associated with the presence of thin outer

shell layers. One of these genera is Margarella Thiele,

1893 (see Zelaya, 2004): tie new species cannot be

allocated into. this genus because of its larger size,

presence of thin shell wall, and absence of a : parietal

callus. The other genus is Gaza Watson, 1879; Falsi-

margarita stephaniae can not be included in Gaza given

its ‘aller shell, more rounded spiral whorls, and absence

of a flap covering the umbilicus.

Additional comparisons and discussion about Falsi-

margarita is provided by Dell (1990: 93). At first glance,

the new species could also be assigned to the Indo-

Pacific genus Otukaia Ikebe, 1942; however the new

species has lower profile, more elaborate sculpture, and

a more widely open umbilicus.

Falsimargarita stephaniae differs from the remaining

congeneric species F. iris (Smith, 1915), F. gemma

(Smith, 1915), F. thielei (Hedley, 1916), F. georgiana

Dell, 1990 and F. benticola Dell, 1990) by having

shouldered whorls, and by the strength of the spiral

folds. The other species have a rounded whorl profile

and a uniformity of spiral sculpture. Only F. thielei

possesses differentiable spiral cords resembling those of

F. stephaniae; however, F. stephaniae additionally differs

from F. thielei by having a larger number of those

outstandingly large spiral “eons at the shell periphery

and a taller spire.

The bathymetry is also a distinctive among Falsi-

margarita species. Falsimargarita gemma, F. iris, and

F. thielei occur in depths to 400 m. Falsimargarita

benthicola and F. georgiana are found in deeper waters

around 3000 m, ‘while "F. stephaniae occurs at interme-

diary depths, around 1200 m.

Until the discovery of the new species, the only

Falsimargarita known to occur in latitudes north of 50°S

was F. iris, which reaches 35°S (Rosenberg, 2004).

Falsimargarita stephaniae is the second species recorded

for these latitudes.

ACKNOWLEDGMENTS

We thank Helen Racz for the donation of the studied

specimen; José H. Leal, The Bailey-Matthews Shell

Museum, Sanibel, Florida, for literature; Lara Guimar-

aes, MZSP, for help with SEM; Diego Zelaya and one

anonymous referee for comments on the matsp

This study is We artially developed with financial help from

the State of Sao Paulo through grants from Pape sp

E. C. Rios and L. R. L. Simone, 2005 Page 173

(Fundagao de Amparo a Pesquisa do Estado de Sao Powell, A. W. B. 1951. Antarctic and Subantarctic Mollusca: Pelecy-

Paulo). processes 04/00309-2, 04/02333-S. poda and Gastropoda. Discovery Reports 26: 47-196 + pls. 5-10

Rosenberg, G, 2004. Malacolog version 3.3.3, Western Atlantic

Gastropod Database. http://data.acnatsci.org/wasp/

LITERATURE CITED index.php (accessed on 07/11/2005).

; Smith, E. A. 1915. Mollusca. Part I. Gastropoda Prosobranchia,

Dell, R. K. 1990. Antarctic Mollusca, with special reference to Scaphopoda, and Pelecypoda. British Antarctic (“Terra

the fauna of the Ross Sea. Royal Society of New Zealand Nova”) Expedition, 1910. Natural History Report. Zoology

Bulletin 27: i-iv, + 1-311. 2: 61-112, pls. 1-2.

Hedley, C. 1916. Australia—Antarctic expedition 1911-1914. Zelaya, D. G. 2004. The genus Margarella Thiele, 1893

C — Zoology and Botany. Government Printer. Adelaide (Gastropoda: Trochidae) in the southwestern Atlantic

4(1): 1-80 + pls. 1-9 Ocean. The Nautilus 118; 112-120.

Notice

BIVALVIA 2006 - INTERNATIONAL CONGRESS ON BIVALVIA, BELLATERRA (BARCELONA), SPAIN, 22—27 JULY 2006

The congress, to be held at the Universitat Autonoma de Barcelona, calls together neontologists and palaeontologists with research

interests in bivalve mollusks. Plenary talks include population genetics, evolution of ontogeny, evolutionary paleontology,

biomineralization, and freshwater conservation biology, but contributions need not be restricted to these topics. In addition, there

will be a planning session for a new bivalve treatise

Two one-day excursions—one each on recent and fossil bivalves—will be organized.

Interested parties are asked to register and submit abstracts via the congress webpage http://bivalvia2006.uab.es.

Further inquiries may be directed to Niko Malchus ([email protected]). Please include in the subject line “Bivalvia 2006”.

THE NAUTILUS 119(4):174, 2005 Page 174

Notice

2006 MEETINGS OF THE AMERICAN MALACOLOGICAL SOCIETY AND THE WESTERN SOCIETY OF MALACOL-

OGISTS, SEATTLE, WASHINGTON, 29 JULY—35 AUGUST 2006

The 72" amual meeting of the American Malacological Society and the 39th annual meeting of the Western Society of Malacologists

will be held jointly in Seattle from 29 July—3 August 2006 under the coordination of AMS and WSM co-president Dr. Roland C.

Anderson. The meeting's main venue will be the University of Washington. Reasonably priced housing will be available at the

University dormitories and the University Inn Motel. The opening night reception will be held at the Burke Museum, located on

campus, and the closing banquet will be at UW's University Club, also on campus. Thursday, 3 August, will be devoted to field trips.

The meeting will include three symposia: one on cephalopod behavior organized by Jennifer Mather of the University of Lethbridge,

one on chitons organized by Douglas Eernisse of the California State University at Fullerton, and one on opisthobranchs organized

by Sandra Millen of the University of British Columbia, Canada.

There will be a sale of malacological reprints to benefit the student fund of WSM and the traditional spirited auction of books and

molluscan memorabilia (no shells) that will benefit the student funds of both organizations. Several notable items of cephalopod art

have already been donated as well as a copy of R. T. Abbott's 24 edition of American Seashells. Bring some of your reprints, books,

and molluscan art to benefit this very worthy cause! Reprints and auction items can be sent to Roland Anderson at the address below.

For further information please contact AMS and WSM president:

Dr. Roland C. Anderson

1483 Alaskan Way

Seattle, WA 98101 USA

[email protected]

THE@MNAUTILUS

Volume 119

2005

AUTHOR INDEX

ARDDIUAS NUE svcidvetare ns aseeaaneeeheepie saat Ooteins eRonanaee es 157 INTEDSEN: 'S. (No aswriewe tecnica saat sna akalaa eas ennaeieaneue ge yee 153

BARWICK RG orig oc gaqiGes ace einenaeeascencsts eneensaastdasete 45 INQRBIS. Ws cos Moen sec cua cme Gneruieushaeusicndecotan odeniges 83

CARBONINIg Al IK: geeieiedjacsoassdane vores aes tams ot arteadene 157 RIOSS Es. (Crass cep badges oerohee ebehee ban metas decks dade duce vs 169

CARPENTERS D3. icinntasdncentst aun adawas acduaoede weananinlseaes 105 PASTORING OG. s..sicisideshinin avaesatertana ated eeadnareedartaiagdeor 55

GARRANZA, (As, 61gscne tacticant cebcee Stas ca ditbenoeang Ted ide ond SS ROE, Kea Jie sates cedh ca des ntse att ecwcaetedt one wees deaeeleuesasa ]

DEV RIES) Ts Pv deasecccedeageiae dace sonmegemeethesdeara 164 ROEAN,, Bis seis vier crasscasbaledeawsaies sadselstials obeaualiedes ee L109

GESUBREGBT, Mi. ois.ccaceannnes a acuuielichinns gedeaainadeasmovasints 15 ROTH, Disseshesat saute dele eeneatance tantane se chad Kone bes ened tests 116

IDIAZS! Iles Mic sta%ememnesicdoe seme arouse nwcsiag tee twacceh tees 157 SAUD Mie Ri sor avinsdcteics Swe do ONE ee ea ewe e spake een sedate 133

FIARASEWYGH, (Mi (Gide. vice cccscadineamugnonieesaetasasc 149 DAVARESE: 05 Po wesash ganansecaaeeaians camsieniia panda viedajeutemad ete 1]

FIARTRIBEDY Pe IO) osiseccsat ctahunt chide aber betta tite aoerien teed ] SIMONE: ia. i, Us, Jace ctcatevehtdeesd neh ecadesgiame se oacseres 169

ITUARTEMC 5, cco ictcagioteemannenontenieae oem eaatunyiaeees teeta 93 SLAPCINSRYS. [sd veeciros cei terete dealers ln Ga-as Mahasls sta ginctsteraueislet 27

KANTOR, Xo We saw rtevatanesd fesitinaa waa ndsaiaatetee badass 149 SMBIGLION Ce caciats aissaied'e ba /anacigle’d aitiniederuiila aintde gtuene Pees edce 109

INOMEER:, ij-c tsa cecensne cusiined etakeerien rede tuniaatenenas 15, 90 SOWIE SAI Uae A statate cscs wna egeardetomn kev neki nad vctase ear ohe soa tees 133

MARIOTIING 4Pas gaiacines ater staceaa ceovandices weananbanne sane 109 STRONG (E.G aiden uate Meranctqianihea enneagad sop waned weiss 119

NITLOSEAVIOH:. PS 2c 2.60.8 oActcacat ect etepeuecaniutaemeeaaees 157 MATES? Aso fc Roscch ictus Seta ea ctee eo netenereyetesane seas 45, 91

MINTON:, Re s.o2o2cce ccna sdensieedecunteieoudatevertereaeeyet exe ll VILVENS Os.s eeancuPeae eteiar + teaendeawed adareracyneras des 50

NEW TAXA PROPOSED IN VOLUME 119 (2005)

GASTROPODA

Akera julieae Valdés and Barwick, 2005, new species (Akeridae) .. 2.2... eee 44

Bullamirifica Squires and Saul, 2005, new genus (Indeterminate family)... ..........0...005 baa aedeeite aad pei oe Se Bs 37

Bullamirifica elegans Squires and Saul, 2005, new species (Indeterminate HAT Y i. asco cavis Bane inns che dceye aore id deh, Sk Saji ettee as 140

Bullamirifica verruca Squires and Saul, 2005, new species (Indeterminate family) Rod barhak Bedaes Seaga os 4 bee ene eS 138

Calliotropis pulvinaris Vilvens, 2005, new em (Trochidaé).. icc shoe ave seeded aon FoR be ee dea eh Pa Eee eee bee 50

Coralliophila trigoi Mariottini, Smriglio and Rolan, 2005, new species (Muricidae)... 2... 0 110

Daffymitra Harasewych, 2005, new genus (Volutomitridae).. 2... eee eee 149

Daffymitra lindae Harasewych, 2005, new species (Volutomitridae) .. 0... ee 150

Exilia alanbeui Nielsen, 2005. new species (Ptychatractidae)s 3 sc05 cee ev aks ee Oa dee ba PAR a PRT oe wees OE 154

Falsimargarita stephaniae Rios and Simone, mae new species (Trochidae). 20h. sate ganda Vane tak wa de dae ee san end 169

Minytropis Squires and Saul, 2005, new genus (Capulidae). 2... ee 143

Minytropis melilota Squires and Saul, 2005, new species (Capulidae). . 0... ee 143

Nerita orovillensis Squires and Saul, 2005, new species (Neritidae) ... 0... eee 136

Paryphantopsis abstrusa Slapcinsky, 2005, new species (Charopidae)... 2... 0. eee 29

Paryphantopsis koragae Slapcinsky, 2005, new species (Charopidae) .. 2... 0. ee 3

Paryphantopsis lebasii Slapcinsky, 2005, new species (Charopidae) ... 2... eee 33

Paryphantopsis matawanensis Slapcinsky, 2005, (Charopidae) ... 0... 0. ee ee 35

Paryphantopsis ubwamensis Slapcinsky, 2005, new species (Charopidae) . 2... ee 37

Paryphantopsis yawii Slapcinsky, 2005, new eee 610170120) 0) (6 1c) APA eR ee rc 39

Paxitropis Squires and Saul, 2005, new genus (Capulidae). 2.2... ee 144

Paxitropis dicriota Squires and Saul, 2005, new species (Capulidae).. inci ic0e aiwadaad si ade nee edas Ge ow ea ... 144

Pterorytis pacanana DeVries, 2005, new species (Muricidae). 2... ee 165

Tegula jeanae Squires and Saul, 2005, new species (Trochidae)... 2.2.0. eee 135

Trophon parodizi Pastorino, 2005, new species (Muricidae)... 1.0.2... eee ee 76

BIVALVIA

Hamiota Roe and Hartfield, 2005, new genus (Unionidae) .. 2.2... ee mceine ene

Pisidium chicha Ituarte, 2005, new species (Sphaeriidae) . 2.2... ee ‘8 : .. 100

Pisidium ocloya Ituarte, 2005, new species (Sphaeriidae).... 2.0.2 ee Pe, ope earaeoet .. 97

Pisidium omaguaca Ituarte, 2005, new species (Sphaeriidae). 2... 0.0.0 pd oe eeces ous . 94

REVIEWERS FOR VOLUME 119

Philippe Bouchet Steffen Kiel Luiz Ricardo L. Simone

Arthur E. Bogan Sadao Kosuge Geerat J. Vermeij

Robert H. Cowie Bruce A. Marshall Emily H. Vokes

Kevin S. Cummings Paula M. Mikkelsen Richard C. Willan

Marta J. deMaintenon Russ Minton James D. Williams

Emilio F. Garcia Guido Pastorino Andrzej Wiktor

Daniel L. Graf Maria del Carmen Perrilliat John B. Wise

M. G. Harasewych Richard E. Petit

Diego Zelaya

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