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AMERICAN

MALACOLOGICAL

BULLETIN

SMITHS?^

^tWO5?003

-^^aries

Journal of the American Malacological Society

hUp://www. malacological. org

i

VOLUME 23 December 27, 2007 number 1/2

The Publications of the American Malacological Union/Society.

EUGENE V. CO AN and ALAN R. RABAT 1

Taxonomic occurrences of gastropod spermatozeugmata and non-stylommatophoran

spermatophores updated. ROBERT ROBERTSON 11

A developmental perspective on evolutionary innovation in the radula of the predatory neogastropod family Muricidae.

GREGORY S. HERBERT, DIDIER MERLE, and CARLOS S. GALLARDO 17

Phylogenetic relationships of the columbellid taxa Cotoiiopsis and Cosrnioconcha

(Neogastropoda: Buccinoidea: Columbellidae). HELENA FORTUNATO 33

Family Pseudolividae (Caenogastropoda, Muricoidea): A polyphyletic taxon.

LUIZ RICARDO L. SIMONE 43

Gastropod mating systems: An introduction to the symposium. JANET L. LEONARD 79

Reproductive behavior of the dioecious tidal snail Cerithidea rliizophornrwn (Gastropoda: Potamididae).

MAYA TAKEUCHI, HARUMI OHTAKI, and KIYONORI TOMIYAMA 81

Causes of variation in sex ratio and modes of sex determination in the Mollusca — an overview.

YOICHIYUSA

continued on back cover

Cover photo: Radula of Chicopiiwntus laqueatus (Sowerby, 1841 ) from Herbert et al.

AMERICAN MALACOLOGICAL BULLETIN

Kenneth M. Brown, Editor-iii-Chief Department of Biological Sciences Louisiana State Lhiiversity Baton Rouge, Louisiana 70803

USA

BOARD OF EDITORS

Cynthia D. Trowbridge, Managing Editor

Department of Zoology

Oregon State University

Corvallis, Oregon 97331

USA

lattice Voltzow Department of Biology University of Scranton Scranton, Pennsylvania 18510-4625 USA

Robert H. Cowie

Center for Conservation Research and Training

University of Hawaii

3050 Maile Way, Gilmore 408

Honolulu, Hawaii 96822-2231

USA

Carole S. Hickman

University of California Berkeley

Department of Integrative Biology

3060 VLSB #3140

Berkeley, California 94720

USA

Timothy A. Pearce

Carnegie Museum of Natural History 4400 Forbes Avenue Pittsburgh, Pennsylvania 15213-4007 USA

Paula M. Mikkelsen Paleontological Research Institution 1259 Trumansburg Road Ithaca, New York 14850-1313 USA

Alan I. Kohn Department of Zoology Box 351800

University of Washington Seattle, Washington 98195 USA

Dianna Padilla

Department of Ecology and Evolution Stony Brook University Stony Brook, New York 11749-5245 USA

Roland C. Anderson The Seattle Aquarium 1483 Alaskan Way Seattle, Washington 98101 USA

The American Malacological Bulletin is the scientific journal of the American Malacological Society, an international society of professional, student, and amateur malacologists. Complete information about the Society and its publications can be found on the Society's website: http://www.malacological.org

AMERICAN MALACOLOGICAL SOCIETY MEMBERSHIP

MEMBERSHIP INFORMATION: Individuals are invited to com- plete the membership application available at the end of this issue.

SUBSCRIPTION INFORMATION: Institutional subscriptions are available at a cost of $65 plus postage for addresses outside the USA.

Further information on dues, postage fees (for members outside the U.S.), and payment options can be found on the Membership Application at the end of this issue.

ALL MEMBERSHIP APPLICATIONS, SUBSCRIPTION ORDERS, AND PAYMENTS should be sent to the Society Treasurer:

Dawn E. Dittman

Tunison Laboratory of Aquatic Science 3075 Gracie Rd.

Cortland, New York 13045-9357 USA

CHANGE OF ADDRESS INFORMATION should be sent to the Society Secretary:

Paul Callomon Department of Malacology

The Academy of Natural Sciences of Philadelphia 1900 Benjamin Franklin Parkway Philadelphia, Pennsylvania 19103-1195 USA

INFORMATION FOR CONTRIBUTIONS is available on-line and appears at the end of this issue.

MANUSCRIPT SUBMISSION, CLAIMS, AND PERMISSIONS TO

REPRINT JOURNAL MATERIAL should be sent to the Editor-in-Chief:

Kenneth M. Brown, Editor-in-Chief

Department of Biological Sciences

Louisiana State University

Baton Rouge, Louisiana 70803

USA

Voice: 225-578-1740 • Fax: 225-578-2597 E-mail: kmbrown@lsu.edu

AMERICAN MALACOLOGICAL BULLETIN 23(1/2) AMER. MALAC. BULL.

ISSN 0740-2783

Copyright © 2007 by the American Malacological Society

AMERICAN MALACOLOGICAL BULLETIN CONTENTS VOLUME 23 I NUMBER l/2

The Publications of the American Malacological Union/Society.

EUGENE V. GOAN and ALAN R. RABAT 1

Taxonomic occurrences of gastropod spermatozeugmata and non-stylommatophoran

spermatophores updated. ROBERT ROBERTSON 11

A developmental perspective on evolutionary innovation in the radula of the predatory neogastropod family Muricidae.

GREGORY S. HERBERT, DIDIER MERLE, and CARLOS S. GALLARDO 17

Phylogenetic relationships of the columbellid taxa Cotouopsis and Cosmiocoucha

(Neogastropoda: Buccinoidea: Columbellidae). HELENA EORTUNATO 33

Family Pseudolividae (Caenogastropoda, Muricoidea): A polyphyletic taxon.

LUIZ RICARDO L. SIMONE 43

Gastropod mating systems: An introduction to the symposium. JANET L. LEONARD 79

Reproductive behavior of the dioecious tidal snail Cerithidea rhizophoranun (Gastropoda: Potamididae).

MAYA TAKEUCHI, HARUMI OHTAKI, and KI YONORI TOMIYAMA 81

Causes of variation in sex ratio and modes of sex determination in the Mollusca — an overview.

YOICHIYUSA 89

Poecilogony and larval ecology in the gastropod genus Alderia. PATRICK J. KRUG 99

Food intake, growth, and reproduction as affected by day length and food availability in the pond snail Lymnaea stagnalis.

ANDRIES TER MAAT, COR ZONNEVELD, J. ARJAN G.M. DE VISSER, RENE F. JANSEN,

KORA MONTAGNE-WAJER, and JORIS M. KOENE 113

Phally polymorphism and reproductive biology in Arioliwax (Ariolimax) buttotii (Pilsbry and Vanatta, 1896) (Stylommatophora: Arionidae).

JANET L. LEONARD, JANE A. WESTFALL, and JOHN S. PEARSE 121

A review of mating behavior in slugs of the genus Deroceras (Pulmonata: Agriolimacidae).

HEIKEREISE 137

Reproductive biology and mating conflict in the simultaneously hermaphroditic land snail

Arianta arhustorum. BRUNO BAUR 157

A literature database on the mating behavior of stylommatophoran land snails and slugs.

ANGUS DAVISON and PETER MORDAN 173

The function of dart shooting in helicid snails. RONALD CHASE 183

Meeting Announcement 190

Melbourne Romaine Carriker: 25 February 1915 - 25 February 2007 An Appreciation.

CLEMENT L. COUNTS, III, ROBERT S. PREZANT, and J. EVAN WARD 191

Financial Report 195

Index to Vol. 23 197

Membership Form 201

Information for Contributors 203

1

\

i

1

I

j

Amer. Malac. Bull. 23; 1-10

The Publications of the American Malacological Union/Society

Eugene V. Coan^ and Alan R. Kabat^

‘ Santa Barbara Museum of Natural History, 2559 Puesta del Sol Road, Santa Barbara, California 93105, U.S.A., gene.coan@sierraclub.org

“ Museum of Comparative Zoology, Hai"vard University, Cambridge, Massachusetts 02138, U.S.A., alankabat@aol.com

Abstract: This paper documents the publications of the American Malacological Society, and its predecessor, the American Malacological Union, from 1931 through 2007. Information on the dates of publication is included, based primarily on library receipt records. Several publications were erroneously dated as to year, which has resulted in new taxa described in those publications, as well as abstracts and articles, being attributed to the wrong year.

Key words: AMS, AMU, publication dates, bibliography

The American Malacological Union (AMU), now the American Malacological Society (AMS), founcied in 1931, issued reports containing abstracts and papers presented at its annual meetings from its earliest years. At first these reports were published in The Nautilus. Shortly thereafter, reports in The Nautilus were supplemented by a printed report mailed to all members after the annual meeting that included a membership list. By 1941, The Nautilus no longer included meeting reports. The titles of the AMU’s reports changed several times over the years (Bieler and Kabat 1991 - present). During World War II, when no meetings were held, reports were issued to ensure communication and continuity.

In 1971, the AMU’s publication became known as the Bulletin. In 1983, the Bulletin became a peer-reviewed sci- entific journal, the American Malacological Bulletin, with an increasing number of original articles. By 1985, the American Malacological Bulletin no longer included reports on and abstracts of papers presented at the annual meeting, which were published only in the program and abstracts volume issued to meeting participants, with the organizer of each meeting generally responsible for its publication. By this time, the membership list was no longer included in the annual report, and was instead published as part of, or in connection with, the organization’s newsletter.

The American Malacological Union Newsletter began in 1968 and was issued irregularly. Its name changed to the AMU News in 1984, to American Malacological Newsletter in 1996, thence to American Malacological Society Newsletter in 1999, and finally to the Newsletter of the American Malaco- logical Society in 2003. The newsletter is now circulated chiefly electronically and is available on the AMS website: http://www.malacological.org/. In 2003, the membership list moved from the newsletter to become a separate, electroni- cally distributed publication.

The purpose of this paper is to provide the publication dates of AMU/AMS publications. In many cases, incorrect dates were printed on them, and some issues were misnum- bered. As a result, abstracts, papers, and even new taxa that appear in this series are sometimes misdated in subsequent literature.

Although the AMU/AMS and its publications have been documented in three histories (Murray 1999, Teskey 1964, 1982), three indices (Anonymous 1966, Counts 1988, Teskey 1975), and one listing (Anonymous 1981), none of these provided fully accurate or complete dates for the AMU/AMS publications. The Annual Report for 1967 (Report No. 34: 84) provided the dates of publication for the reports from 1961 through 1966; we are unaware of any other similar listings.

The reports appearing in The Nautilus from 1931 to 1941 are easily dated using the compilation of Coan and Harasewych (1993).

The individual reports and bulletins contain various kinds of dates. The most common is the date the included membership list was prepared, herein indicated by “Mem- bership List.” These cannot be taken as publication dates, although some other sources have used them, because the report was often printed and mailed weeks or months after the membership list was prepared. These dates, however, are included here for comparison and to indicate that the pub- lication could not have been issued any earlier.

Some reports are simply dated, without further indica- tion as to what this date means (herein “Stated Date”). Start- ing with volume 18 of the American Malacological Bulletin, the printed date, for the first time, gives both the month and day, and is intended and assumed to be the publication date, based on receipt of mailed copies within one or two weeks. In earlier years, however, these dates were demonstrably off by a year. In some cases, the printed dates are labeled as

2

AMERICAN MALACOLOGICAL BULLETIN 23 • 1/2 • 2007

being specifically a publication date or mailing date, essen- tially the same thing. In the mid-1990s, several issues did not have the year on the cover or on the table of contents, but the year was included on the spine; such data, of course, are lost when the journal issues are bound, as is the case for most libraries.

It is instructive to compare any printed dates with the dates the publication was received in libraries, most of which stamp the date of receipt on each copy upon arrival. How- ever, this is also not entirely conclusive, because there may be delays between receipt of the mail and date stamping. Moreover, some institutions with malacologists on their staffs did not separately subscribe to the AMU/AMS publi- cations, and those institutions instead obtained their copies through donations from these persons, often months or years after publication. It appears that for some years the reports were sent by airmail to Europe and by bulk mail within the U.S., accounting for The Natural History Mu- seum (London) having several of the earliest receipt dates.

We have examined the sets of AMU/ AMS publications in the Academy of Natural Sciences of Philadelphia (ANSP), Library of Congress (LOG), National Museum of Natural History (USNM), Harvard University Museum of Compara- tive Zoology (MCZ), California Academy of Sciences (CAS), University of Maryland (UMd), University of Washington Fisheries and Oceanography Library (UW), and The Natural History Museum, London (BMNH). Three colleagues pro- vided us with information on the sets at the Field Museum of Natural History (FMNH), the American Museum of Natural History (AMNH), and the Carnegie Museum of Natural History (CMNH). When these dates are consistently later than the published dates — sometimes much later — it is clear that the stated publication dates must be incorrect. If no date is given for a particular institution for a given year, this means that the institution ( 1 ) lacks the volume in ques- tion; (2) has the volume, but there is no date stamp; or (3) has the volume, but the date stamp is several weeks, months, years, or even decades later. In several cases, one of us (EVC) noted the date of receipt of his personal copies.

We have also included the three “Special Publications " which were issued in the 1980s as supplements to the Ameri- can Malacological Bulletin. However, as these were not in- cluded in the regular subscription, many libraries did not order them.

When we are certain or fairly certain about publication dates, these are indicated with a single date. In other cases, we present the evidence thus far available, which in many cases indicates that the Bulletins were issued the year follow- ing the meeting, or months, or even a year after the internal date.

For reference, the place and dates of the meetings through 1985 are given, along with the number of the meet-

ing. After the publications stopped covering the meetings, this information is not provided here, but is readily available online (Coan and Rabat 1996-present). Also indicated are the meetings of the Pacific Division of the AMU from its inception in 1948 until its separation as the Western Society of Malacologists in 1968.

The International Code of Zoological Nometiclature (ICZN 1999) emphasizes that editors and publishers should keep track of dates of publication in journals that may have new taxa or other nomenclatural acts:

“Recommendation 2 1C. Specification of date. An editor or publisher should state the date of publi- cation of a work, and of each component part of a serial publication, and of any work issued in parts.

In a volume made up of parts brought out sepa- rately, the day of publication of each part, and the exact pages, plates, maps, etc. that constitute it, should be specified” (p. 23).

This paper is designed to bring the AMU/AMS publications into retroactive compliance with the Code.

In the recent catalog of family and higher names in the Gastropoda, and elsewhere, the subfamily names Pseudo- melatominae Morrison and the (unavailable) Lophiotomi- nae and Crassispirinae Morrison are dated from 1 December 1965, the date printed on AMU Annual Reports for 1965 (Bouchet and Rocroi 2005: 57, 101, 144, 256), but this report was not actually published until 28 February 1966, as evi- denced in the list below. Four papers by Morrison in AMU publications are misdated in the bibliography of his works by Rosewater (1984), including this paper on the Turridae.

From the list below, it is clear that publication dates have been incorrectly stated, even in recent years. It is ob- vious that in spite of a “publication date” of 30 December 1976, on the cover of the report for 1976, a January 1977 publication/mailing date is more likely because no one re- ceived this issue until early February 1977. Fortunately, no new taxa appear to be included. Similarly, American Mala- cological Bidletin 3(1) is dated December 1984, but no insti- tutions received it until mid-Februai-y 1985, and a late Janu- ary or early February 1985 publication date is more likely. No new taxa were included in this issue. The issue AMB 8( 1 ) was dated August 1990, but it appears not be have been received anywhere before early October, so a publication date of mid- to late-September seems more probable. Again, no new taxa were included in this issue. However, AMB 17(1/2), indicated as having been published in December 2002, appears to have actually been issued in February 2003. This issue contains new taxa of marine and terrestrial gas- tropods, as well as a species of fossil rostroconch, and all of these should evidently be dated 2003 rather than 2002.

AMERICAN MALACOLOGICAL SOCIETY PUBLICATIONS

3

ANNUAL REPORTS AND BULLETIN

Notes

* Publication dates of The Nautilus from Coan and Harasewych (1993). n/a = not applicable.

* Issue numbers printed on some ot the Bulletins, but incorrectly numbered and not corresponding either to the number of the meeting or to the sequential number of separately issued bulletins.

The Nautilus; American Malacologkal Union [Report]

Meeting

Publication

Earliest Date of Publication

Later Receipt Dates

Comments

1 (Philadelphia,

The Nautilus 45{\): 1-5

13 July 1931

n/a

Publication date: Coan

Pennsylvania, 30 April-2 May 1931) n/a

“Members of the American

1 April 1932

n/a

and Harasewych (1993)

Date on cover

2 (Washington, D.C.,

Malacological Union,” 5 pages

The Nautilus 46(1): 1-3

23 July 1932

n/a

Publication date: Coan

26-28 May 1932)

“Mrs. Imogene C.

1932

n/a

and Harasewych (1993)

Undated, presumably

3 (Cambridge,

Robertson’s Rambling Notes”, 1 1 pages The Nautilus 47(1): 37-44

16 June 1933

n/a

issued in 1932 Publication date: Coan

Massachusetts, 25-27 May 1933)

4 (Stanford, California,

The Nautilus 48(2): 72

15 October 1934

n/a

and Harasewych (1993)

Publication date: Coan

25-28 June 1934)

Report, 12 pages

ANSP = 16 October 1934

n/a

and Harasewych (1993)

Stated date = 1 August

5 (Buffalo, New York,

The Nautilus 49(2): 62-63

8 November 1935

n/a

1934

Publication date: Coan

27-29 lune 1935)

Report, 10 pages, 1 plate

not known; possibly late

n/a

and Harasewych (1993)

[no data available]

6 (St. Petersburg, Florida,

The Nautilus 51(1): 33-36

1935 or 1936 14 July 1936

n/a

Publication date: Coan

21-24 April 1936)

The American

MCZ = 21 October 1936

n/a

and Idarasewych (1993)

Membership List = 1

7 (Ann Arbor, Michigan,

Malacological Union [Report], 14 pages The Nautilus 51(2): 68-71

22 October 1937

n/a

September 1936 Publication date: Coan

3-5 August 1937)

8 (Havana, Cuba, 1-6

The American

Malacological Union [Report], 16 pages The Nautilus 52(2): 66-72

MCZ = 26 February 1938 28 October 1938

n/a

and Harasewych (1993)

Membership List = 1 January 1938

Publication date: Coan

August 1938)

The American Malacological Union [Report], 20 pages, frontispiece

MCZ = 28 April 1939

and Harasewych (1993)

Membership List = 1 January 1939

4

AMERICAN MALACOLOGICAL BULLETIN 23 • 1/2 • 2007

Earliest Date of

Meeting

Publication

Publication

Later Receipt Dates

Comments

9 (Toronto, Ontario,

The Nautilus 53(1); 36;

21 July 1939; 20 October

n/a

Publication date; Coan

Canada, 20-23 June

The Nautilus 53(2);

1939

and Harasevvych

1939)

68-72

(1993)

The American

MCZ = 2 January 1940

Membership List =

Malacological Union [Report], 16 pages

November 1939

10 (Philadelphia,

The Nautilus 54(1); 35-37

23 July 1940

n/a

Publication date; Coan

Pennsylvania, 17-21 June

and Harasevvych

1940)

(1993)

The American

MCZ = 17 January 1941

Membership List =

Malacological Union [Report], 18 pages

December 1940

1 1 (Rockland and

The Nautilus 55(2); 70-72

24 October 1941

n/a

Publication date; Coan

Thomaston, Maine,

and Harasewych

26-29 August 1941 )

(1993)

The American

Probably 1942

Membership List =

Malacological Union [Report], 44 pages

January 1942

1942-1943 - No meetings

News Bulletin and Annual Report

Earliest Date of

Meeting

Publication

Publication

Later Receipt Dates

Comments

1944 - No meeting

News Bulletin and Annual

Probably 1944

Membership List =

Report, 24 pages, frontispiece

January 1944

1945 - No meeting

News Bulletin and Annual

MCZ = 26 November

Membership List =

Report, 21 pages, frontispiece

1945

October 1945

12 (Washington, D.C.,

News Bulletin and Annual

MCZ = 22 April 1947;

Membership List =

14-16 August 1946)

Report, 22 pages, frontispiece

ANSP = 22 April 1947

April 1947

13 (Pacific Grove,

News Bulletin and Annual

CMNH = 20 February

Membership List =

California, 18-21 June 1947)

Report, 32 pages

1948

December 1947

14 (Pittsburgh,

News Bulletin and Annual

MCZ = 9 April 1949

ANSP = 18 April

Membership List =

Pennsylvania, 25-27 August 1948); 1, Pacific Division (Los Angeles, California, 10-11 April 1948)

Report, 36 pages

1949

March 1949

15 (Coral Gables, Florida,

News Bulletin and Annual

MCZ = 1 February 1950

ANSP = 6 February

Membership List =

16-18 June 1949); 2, Pacific Division (Long Beach, California, 14-16

Report, 36 pages

1950

November 1949

June 1949)

16 (Chicago, Illinois, June

News Bulletin and Annual

MCZ = 26 January 1951

ANSP = 1 February

Membership List =

14-16, 1950); 3, Pacific Division (Santa Barbara, California, 7-9 April 1950)

Report, 39 pages

1951

December 1950

AMERICAN MALACOLOGICAL SOCIETY PUBLICATIONS

5

Earliest Date of

Meeting

Publication

Publication

Later Receipt Dates

Comments

17 (Buffalo, New York,

News Bulletin and Annual

28 lanuary 1952 [“date

ANSP = 10 March

Membership List =

22-24 August 1951); 4,

Report, 41 pages

based on annotation by

1952; MCZ = 17

December 1951

Pacific Division

Morrison on reprint in

March 1952

(Oakland, California,

MNHN” (Bouchet and

22-24 lune 1951)

Rocroi 2005: 333)]

Annual Report

Earliest Date of

Meeting

Publication

Publication

Later Receipt Dates

Comments

18 (Cambridge,

Annual Report, 44 + [i]

MCZ = 4 February 1953;

Memlaership List = 31

Massachusetts, 20-22

pages

ANSP = 4 February

December 1952

August 1952); 5, Pacific Division (Los Angeles, California, 20-22 lime

1953

[inside front cover]

1952)

19 (Lawrence, Kansas,

Annual Report, 51 pages

MCZ = 15 March 1954;

Stated Date /

25-27 lune 1953); 6,

ANSP =15 March 1954

Membership List = 31

Pacific Division (Pacific Grove, California, 12-13 lune 1953)

December 1953

20 (Durham, New

Annual Report, 44 pages

MCZ = 21 January 1955

ANSP = 26 January

Membership List = 31

Hampshire, 16-18 August 1954); 7, Pacific Division (Los Angeles, California, 18-19 June 1954)

1955

December 1954

21 (New York, New York,

Annual Reports 22*, 58

ANSP = 16 April 1956

MCZ = 1 7 April

Stated Date and

26-29 lune 1955); 8,

pages

1956

Membership List = 31

Pacific Division (Stanford University, California, 15-16 July 1955)

December 195

22 (San Diego, California,

Annual Reports 23*, 52

n/a

LOC = 23 February

Stated Date and

11-14 July 1956); 9,

pages

1958 [late]

Membership List, 31

Pacific Division (same)

December 1956

23 (Yale University, New

Annual Reports 24*, 56

UMd = 28 March 1958

Stated Date = 1 January

Haven, Connecticut, 16-19 July 1957); 10, Pacific Division (Santa Barbara, California, 30 May-1 June 1957)

pages

1958

24 (Ann Arbor, Michigan,

Annual Reports 25*, 73

UMd = 11 February 1959

Stated Date = 1 January

2-6 September 1958); 11, Pacific Division (Berkeley, California, 27-29 lune 1958)

pages

1959

25 (Philadelphia,

Annual Reports 26*, 79

UMd = 26 February 1960

Stated Date = 1 January

Pennsylvania, 30 June-3

pages

1960; meeting

July 1959); 12, Pacific

misnumbered as “26"

Division (Redlands, California, 9-12 July 1959)

on cover and title page

6

AMERICAN MALACOLOGICAL BULLETIN 23 • 1/2 • 2007

Meeting

Publication

Earliest Date of Publication

Later Receipt Dates

26 (Montreal, Quebec, Canada, 9-12 August 1960); 13, Pacific Division (Pacific Grove, California, 22-25 lune 1960)

Annual Reports 27*, 76 pages

EVC = February 1961

BMNH = 4 October 1961; LOC = 2 November 1961

27 (Washington, D.C.,

Annual Reports 28*, 81

12 December 1961

BMNH = 8 lanuary

19-23 lune 1961); 14, Pacific Division (Goleta, California, 28 Iune-1 I Lily 1961)

pages

(Report for 1967, 34: 84)

1962; LOC = 10 lanuary 1962

28 (St. Petersburg, Florida,

Annual Reports 29*, 68

21 December 1962

LOC = 23 lanuary

31 |uly-3 August 1962); 15, Pacific Division (Pacific Grove, California, 27-30 lune 1962)

pages

(Report for 1967, 34: 84)

1963; BMNH = 28 lanuary 1963

29 (Buffalo, New York,

Annual Reports 30*, 83

1 lanuary 1964 (Report

LOC = 23 lanuary

18-21 lune 1963); 16, Pacific Division (Goleta, California, 26-29 lune 1963)

pages

for 1967, 34: 84)

1964; BMNH = 14 February 1964; UMd = 14 February 1964

30 (New Orleans,

Annual Reports 31*, 102

14 December 1964

LOC = 31

Louisiana, 21-24 luly 1964); 17, Pacific Division (Pacific Grove, California, 18-21 lune 1964)

pages

(Report for 1967, 34: 84)

December 1964

31 (New York, New York,

Annual Reports 32*, 122

28 February 1966 (Report

LOC = 3 March

20-23 lune 1965); 18, Pacific Division (San Diego, California, 24-27 lune 1965)

pages

for 1967, 34: 84)

1966; UMd = 7 March 1966

32 (Chapel Hill, North

Annual Reports 33*, 120

28 February 1967 (Report

LOC = 8 March

Carolina, 22-27 August 1966); 19, Pacific Division (Seattle, Washington, 19-22 lune 1966)

pages

for 1967, 34: 84)

1967; UMd = 16 March 1967

33 (Ottawa, Ontario,

Annual Reports 34*, 119

Mailing Date: 20 March

BMNH = 2 May

Canada, 31 Iuly-6 August 1967); 20, Pacific Division (Pacific Grove, California, 28-30 lune 1967)

pages

1968

1968

34 (Corpus Christi, Texas,

Annual Reports 35*, 97

Mailing Date = 27

UMd = 20 lanuary

15-19 luly 1968); 21, Pacific Division (Pacific Grove, California, 19-22 lune 1968)

pages

December 1968

1969

35 (Marinette, Wisconsin,

Annual Reports 36*, 96

Mailing Date = 19

USNM = 13

21-23 luly 1969)

pages

December 1969

January 1970

Comments

Stated Date = 1 lanuary 1961; meeting misnumbered as “27” on cover and title page

Stated Date = 1 December 1961; meeting misnumbered as “28” on cover and title page

Stated Date = 1 December 1962; meeting misnumbered as “29” on cover and title page

Stated Date = 1 December 1963

Stated Date = 1 December 1964

Stated Date = 1 December 1965

Stated Date = 1 December 1966

Stated Date = 1 December 1967

Stated Date = 1 December 1968

Stated Date = 1 December 1969

AMERICAN MALACOLOGICAL SOCIETY PUBLICATIONS

7

Meeting

Publication

Earliest Date of Publication

Later Receipt Dates

Comments

36 (Key West, Florida,

Annual Reports 37*, 112

Mailing Date = 18

UMd = 8 March

Stated Date = 1

16-20 luly 1970)

pages

February 1971

1971; FMNH = 23 March 1971

December 1970

Bulletin

Meeting

Publication

Earliest Date of Publication

Later Receipt Dates

Comments

37 (Cocoa Beach, Florida, 15-19 July 1971)

Bulletiit, 70 pages

ANSP = 2 March 1972

FMNH = 8 March 1972

Stated Date = February 1972

38 (Galveston, Texas, 9-14 July 1972)

Bulletin, 64 pages

Publication Date = 23 March 1973

USNM = 30 March 1973

39 (Newark and Greenville, Delaware, 24-28 June 1973)

Bulletin, 69 pages

Publication Date = 22 May 1974

ANSP = 28 May 1974; UMd = 28 May 1974

40 (Springfield, Massachusetts, 3-7 August 1974)

Bulletin, 94 pages

Index, 1934-1974, [ii] -t 57 PP

Publication Date = May 1975

Probably issued with the preceding in 1975

ANSP = 6 June 1975; UMd = 10 June 1975

undated

41 (San Diego, California,

Bulletin, 94 pages -i- inside

Publication date = 30

UMd = 20 February

22-26 lime 1975)

back cover

January 1976

1976

42 (Columbus, Ohio, 2-6 August 1976)

Bulletin, 89 pages

Publication date = 30 December 1976

UW = 9 February 1977; FMNH = 14 February 1977

43 (Naples, Florida, 10-15 July 1977)

Bulletin, 1 18 pages

BMNH = 24 April 1978

USNM = 28 April 1978; UMd = 28 April 1978

Membership List = 15 October 1977

44 (Wilmington, North Carolina, 16-21 July 1978)

Bulletin, 86 pages

BMNH = 28 March 1979

USNM = 29 March 1979

Membership List = 15 October 1978

45 (Corpus Christi, Texas, 5-11 August 1979)

Bulletin, 86 pages

EVC = March 1980

USNM = 20 March 1980; BMNH = 21 March 1980

Membership List = 1 November 1979

46 (Louisville, Kentucky,

Bulletin, 94 pages

USNM = 19 March 1981;

LOC = 21 March

Membership List = 15

19-25 July 1980)

BMNH = 19 March 1981

1981

October 1980

47 (Ft. Lauderdale, Florida,

Bulletin, 76 pages

BMNH = 15 February

LOC = 23 February

Membership List = 20

19-25 July 1981)

1982

1982; ANSP = 23 February 1982; UMd = 23 February 1982

October 1981

48 (New Orleans,

American Malacological

LOC = 28 June 1983

USNM = 17 July

Stated Date = July 1983

Louisiana, 19-23 July 1982)

Bulletin 1: 136 pages

1983

[cover]; May 1983 [1st page and Counts (1988)]

49 (Seattle, Washington,

American Malacological

BMNH = 30 March 1984

ANSP = 3 April

Stated Date = February

7-13 August 1983)

Bulletin 2: 127 pages

1984; USNM = 3 April 1984

1984

50 (Norfolk, Virginia,

American Malacological

BMNH = 18 February

FMNH = 28

Stated Date = December

22-27 July 1984)

Bulletin 3(1): 1-133

1985

February 1985; USNM = 28 February 1985

1984

8

AMERICAN MALACOLOGICAL BULLETIN 23 • 1/2 • 2007

American Malacological Bulletin

Meeting

Publication

Earliest Date of Publication

Latest Date of Publication

Comments

n/a

American Malacological Bulletin 3(2): [ii] +135-272

BMNH = 17 July 1985

FMNH = 18 luly 1985; LOG = 18 July 1985; UMd = 18 July 1985

Stated Date =

June 1985

n/a

American Malacological Bulletin, Special Edition 1: 116 pages

(not known; presumably on or after July 1985)

LISNM = 7 August 1986 [late]; BMNH = 24 August 1987 [late]

Stated Date =

July 1985

51 (Kingston, Rhode Island, 28 July-2 August 1985)

American Malacological Bulletin 4(1): 1-147

CAS = 5 March 1986

UW = 14 March 1986

Stated Date = 1986

February

Note: the 1985 meeting was the last one to be reported upon in detail in the American Malacological Bulletin. For additional information about subsequent AMU/AMS meetings, see Coan and Rabat (1996-present).

American Malacological Bulletin

PuJilication

Earliest Date of Publication

Latest Date of Publication

Comments

American Malacological Bulletin, Special Edition 2: 239 pages

(not Icnown; presumably on or after June 1986)

FMNH = 1 May 1987 [late]; BMNH = 24 August 1987 [late]

Stated Date = June 1986

American Malacological Bidletin

FMNH = 12 September 1986

AMNH = 15 September 1986

Stated Date = August 1986

4(2): 149-247

American Malacological Bidletin, Special Edition 3: 74 pages

(not known; presumably on or after October 1986)

USNM = 24 April 1987 [late]

Stated Date = October 1986

American Malacological Bulletin 5(1): 1-152

BMNH = 11 February 1987; AMNH = 11 February 1987; FMNH = 11 February 1987

ANSP = 12 February 1987

Stated Date = January 1987

American Malacological Bulletin

BMNH = 14 July 1987

UW = 15 July 1987

Stated Date = June 1987

5(2): 153-306

American Malacological Bulletin

BMNFI = 25 February 1988

FMNH = 1 March 1988

Stated Date = January 1988

6(1): 1-164

American Malacological Bulletin 6(2): 165-305

October 1988

USNM = 10 October 1988

Stated Date = October 1988 [as “July 1988” in Counts (1988); corrected to October in AMB 7(1): 89

American Malacological Bulletin 7(1): 1-91

BMNH = 24 May 1989; ANSP = 24 May 1989

USNM = 26 May 1989; UMd = 26 May 1 989

Stated Date = April 1989

American Malacological Bulletin

UW = 23 March 1990

FMNH = 26 March 1990

Stated Date = February 1990

7(2): 93-175

American Malacological Bulletin

UW = 10 October 1990

BMNH = 12 October 1990

Stated Date = August 1990

8(1): 1-96

American Malacological Bulletin 8(2): 97-182

BMNH = 31 May 1991; ANSP = 31 May 1991

UW = 3 June 1991; UMd = 3 June 1991

Stated Date = April 1991

American Malacological Bulletin 9(1): 1-[104]

BMNH = 30 December 1991

UW = 10 January 1992; ANSP = 11 January 1992

Issue 9(2): 219 gives “December 1991”

American Malacological Bulletin 9(2): 105-219

EVC = August 1992

BMNH = 3 September 1992; ANSP = 3 September 1992

Page 219 gives “August 1992”

American Malacological Bulletin 10(1): 1-[112]

USNM = 18 February 1993

ANSP = 19 February 1993

Issue 10(2): 295 gives “February 1993”

AMERICAN MALACOLOGICAL SOCIETY PUBEICATIONS

9

Publication

Earliest Date of Publication

Latest Date of Publication

Comments

American Malacological Bulletin 10(2); iv + 113-295

ANSP = 8 December 1993

USNM = 9 December 1993

Preface gives “July 1993”; page 295 gives “November 1993”

American Malacological Bulletin

USNM = 30 December 1994;

Page iii gives “December 1994,”

11(1): iii -1- l-[86] pages

ANSP = 30 December 1994

as does issue 11(2): 211

American Malacological Bulletin

ANSP = 8 September 1995

USNM = 11 September 1995

Page 211 gives “August 1995”

11(2): ii + 87-[212]

American Malacological Bulletin 12(1-2); ii -t 1-[156]

AMNH = 24 January 1997

ANSP = 28 January 1997

I^age 155 gives “September 1996”; issue 13(1/2); 154 gives “October 1996”; spine gives “1996”

American Malacological Bulletin 13(1/2): ii -t 1-[156]

AMNH = 10 January 1997

ANSP = 13 January 1997

Page 154 gives “December 1996”; spine gives “1996”

American Malacological Bulletin

ANSP = 5 January 1998

FMNH = 7 January 1998; LOC

Issue 14(2): 234 gives

14(1); i + 1-86

= 8 January 1998

“December 1997”

American Malacological Bulletin

LOC 25 January 1999

AMNH = 28 January 1999

Stated Date “December 1998”

14(2): ii -I- 87-234 pages

American Malacological Bidletin 15(1); iii -t 1-111

ANSP = 15 November 1999

AMNH = 16 November 1999

Stated Date “1999”; issue 15(2): 208 gives “October 1999”

American Malacological Bidletin 15(2); i -1- [113]-208

BMNH = 22 January 2001

AMNH = 29 [anuary 2001

Page 208 gives “December 2000”

American Malacological Bulletin 16(1/2): iii -1- 1-266 pages

EVC = 7 November 2001

Page 266 gives “September 2001”

American Malacological Bulletin

BMNH = 26 February 2003

ANSP = 3 March 2003; AMNH

Stated Date = “December

17(1/2): iii -h 1-165

= 4 March 2003

2002”; to be mailed January 2003 [e-mail, 3 Januaiy 2003 from A. Valdes to R. Bieler]

American Malacological Bulletin 18(1/2): i + 174 pages

7 May 2004

Publication date given on front cover

American Malacological Bulletin 19( 1/2); i -1- 150 pages

14 October 2004

Publication date given on front cover

American Malacological Bulletin 20( 1/2): ii -t- 165 pages

27 April 2005

Publication date given on front cover

American Malacological Bulletin 21( 1/2); i -H 131 pages

9 February 2006

Publication date given on front cover

American Malacological Bidletin 22( 1/2): V -1- 176 pages

26 March 2007

Publication date given on front cover

OTHER PUBLICATIONS

Anonymous. 1940. H. A. Pilshry: Scientific coiitribiitions made from 1882 to 1939. American Malacological Union. 63 pages. (This includes a short biography of Pilsbry, and may have been edited by his colleague, H. B. Baker].

How to Collect Shells

The predecessor to this stand-alone work was included in the re- port of the Eleventh Annual Meeting of the American Mala- cological Union as full-length papers by F. C. Baker, H. van der Schalie, W. (. Clench, B. R. Bales, T. Burch, (. S. Schwen- gel, and T. L. McGinty [with one additional page of discus- sion], 1942. Methods of collecting and preserving Mollusca. Symposium papers. American Malacological Union, Eleventh Annual Meeting, [Report]: 5-37.

Abbott, R. T., G. M. Moore, 1. S. Schwengel, and M. C. Teskey, eds. 1955 [March 31]. How to Collect Shells {A Symposium), 1st ed. American Malacological Union, Buffalo, New York, [iv] -I- 75 + [5] pp.

LaRocc]ue, A., ed. 1961. How to Collect Shells {A Symposium), 2nd ed. American Malacological Union, Marinette, Wisconsin, iv -I- 92 -H [5] pp.

Abbott, R. T., M. K. Jacobson and M. C. Teskey, eds. 1966. How to Collect Shells [A Symposium), 3rd ed. American Malacological Union, Marinette, Wisconsin, iv + 101 -I- [5] pp.

Jacobson, M. K., ed. 1974. How to Study and Collect Shells. {A Symposium), 4th ed. American Malacological Union, Wrights- ville Beach, North Carolina, [iv] -I- 107 pp.

Sherborn, C. D. and E. R. Sykes, 1906. [Reprint of] Museum Bolt- enianum by P. F. Roding, 1798, [iii] -I- viii -I- 199 pp. [subse-

10

AMERICAN MALACOLOGICAL BULLETIN 23 • 1/2 • 2007

quently reprinted by the American Malacological Union, 1986],

Sturm, C. F., T. A. Pearce, and A. Valdes, eds. 2006 [25 Inly], The Molliisks: A Guide to Their Study, Collection, and Preservation. Boca Raton, Florida: Universal Publishers, xiii + 445 pp.

ACKNOWLEDGMENTS

We thank R. Bieler (LMNH), P. Mikkelsen (AMNH), and T. A. Pearce (CMNH) for providing us with informa- tion about the dates of receipt of AMU/AMS publications in their institutional libraries. We also thank C. L. Sturm, lr.> for information on the editions of How to Collect Shells and R. E. Petit, J. Voltzow, and an anonymous reviewer for their helpful reviews of an earlier draft of this paper.

introduced by loseph P. E. Morrison (December 17, 1906 - December 2, 1983). The Nautilus 98(1): 1-9.

Teskey, M. C. 1964. Flistory of the American Malacological Union. The Shelletter of Shells and Their Neighbors [California] 23: 7.

Teskey, M. C. 1975. Index, 1934 through 1974. American Malaco- logical Union, 57 pp. [This index probably mailed with the Bulletin for 1975, but was not published as an issue or as a numbered supplement of that serial).

Teskey, M. C. 1982. Half-century of AMU. Bulletin of the American Malacological Union for 1981: [iii]-[v].

Accepted: 5 February 2007

LITERATURE CITED

Anonymous. 1966. Abstract index (1949-1965) [and] author index, abstracts, 1949-1965. American Malacological Union, Annual Reports for 1965: 105-122.

Anonymous. 1981. Miscellanea. Annual Reports and Bulletins of the American Malacological Lhiion. Malacological Review 14: 67-112.

Anonymous. 1989. [Correction]. American Malacological Bulletin 7(1): 89.

Bieler, R. and A. R. Rabat. 1991. Malacological lournals and News- letters, 1773-1990. The Nautilus 105(2): 39-61, periodically updated version available at: http://fml.fieldmuseum.org/ collections/search.cgi?dest=mjl

Bouchet, P. and l.-P. Rocroi. 2005. Classification and nomenclator of gastropod families. Malacologia 47(1-2): 1-397.

Coan, E. V. and M. G. Harasewych. 1993. Publication dates of The Nautilus. The Nautilus 106(4): 174-180.

Coan, E. V. and A. R. Rabat. 1996. Annotated catalog of malaco- logical meetings, including symposia and workshops in mala- cology. American Malacological Bulletin 13(1-2): 129-148, pe- riodically updated version available at: http://erato.acnatsci .org/ams/pdfs/symposia.pdf

Counts, C. L., 111. 1988. Index to the American Malacological Bul- letin: 1983 to 1988. Volumes 1 through 6, Special Edition numbers 1-3. American Malacological Bulletin 6(2): 219-305 [see also anonymous correction in 7(1): 89, 1989].

ICZN [International Commission on Zoological Nomenclature]. 1999. International Code of Zoological Nomenclature. London, England (International Trust for Zoological Nomenclature), xxix -r 306 pp.

Murray, H. D. 1999. History {evolution) of the American Malaco- logical Union (Society). Privately published, San Antonio, Texas. 15 pp. [distributed at the 1999 meeting of the American Malacological Society, Pittsburgh, Pennsylvania].

Rosewater, 1. 1984. A bibliography and list of taxa of Mollusca

Amer. Malac. Bull 23: 11-16

Taxonomic occurrences of gastropod spermatozeugmata and non-stylommatophoran spermatophores updated

Robert Robertson’*^

Department of Malacology, The Academy of Natural Sciences of Philadelphia, 1900 Benjamin Franklin Parkway,

Philadelphia, Pennsylvania 19103-1 195, U.S.A., hhandrrconch@aol.com

Abstract: Spermatozeugmata, not to be confused with spermatophores, that also transfer sperm, are compound structures (parasperms with attached eusperms) known only in certain “mesogastropods”: Loxonematoidea? (Abyssochrysidae), Littorinoidea (Littorinidae), Triphoroidea (Triphoridae and Cerithiopsidae), Tonnoidea (Ranellidae), Janthinoidea (Epitoniidae and Janthinidae), and doubtlully Cypraeoidea (Cypraeidae). This pattern of taxonomic occurrence does not match that of any other character known, their morphology is diverse, and it is concluded that the spermatozeugmata in these taxa are not all homologous and that, like spermatophores, they have evolved repeatedly. Littorinid spermatozeugmata have frequently been studied after fixation and shrinkage of the parasperms (“nurse cells”), during which the eusperms drop off. Spermatozeugmata are not a synapomorphy linking the Triphoroidea and Janthinoidea. Records of spermatophores (except in the “pulmonate” suborder Stylommatophora) since Robertson (1989) are updated.

Key words: sperm transfer, inferred homoplasy, euspermatozoa, paraspermatozoa, nurse cells

SPERMATOZEUGMATA

Spermatozeugmata (singular: spermatozeugma) are not to be confused with spermatophores (Robertson 1989) al- though both transfer sperm. Spermatozeugmata are of in- tracellular origin and consist of single paraspermatozoa (Hodgson 1997, Buckland-Nicks et al. 2000) with numerous euspermatozoa (fertile sperm) attached externally by their acrosomes. Parasperm are also called apyrene sperm and nurse cells. They are known only in “prosobranchs”, but do not all become spermatozeugmata. Spermatophores are se- creted extracellularly and contain the eusperms. Spermato- phores occur spioradically in many major groups within all gastropods, including many stylommatophorans. Sperma- tozeugmata are even more sporadic and are known only in certain “mesogastropods”:

Incertae sedis (Loxonematoidea?): Abyssochrysidae: Abyssochrysos (Healy 1989).

Littorinoidea: Littorinidae: Littoraria (Reinke 1911, as “Littorina”)y Littorina, Littoraria (Reinke 1912, latter as “Lif- torina”), Littorina (Ankel 1930: 599, 600; Linke 1933), Lit- toraria (Woodard 1942a, 1942b, Lenderking 1954, all as ‘'Lit- torina”, latter: nurse cell as “spermatophore”), Melarhaphe (Battaglia 1952, as “Littorina”), Littoraria (Marcus and Mar- cus 1963, as “Littorina”), Bembicium (Bedford 1965), Cen- chritis (as “Tectarius”), Tectarins (as “Echininus”), Littoraria (as “Littorina”), Nodilittorina (in part as “Littorina” (all Borkowski 1971), Littorina (Buckland-Nicks 1973, Buck-

Current address: 510 Homestead Ave., Haddonfield, New lersey 08033, U.S.A.

land-Nicks and Chia 1977), Nodilittorina (Jordan and Ramorino 1975, as “Littorina”), Littoraria (Reid 1986, Healy and Jamieson 1993, Buckland-Nicks et al. 2000).

Triphoroidea [including “Cerithiopsoidea”]: Triphori- dae: Triplwra (Houston 1985), Viriola (Healy 1987, 1990). Cerithiopsidae: Cerithiopsis (Fretter and Graham 1962, Houston 1985), Seda (Houston 1985, Healy 1990).

Cypraeoidea?: Cypraeidae?: Erronea? (Healy 1986a, as “Cypraea”).

Tonnoidea: Ranellidae: Fusitriton (Buckland-Nicks et al 1982).

Janthinoidea [“Epitonioidea”]: Epitoniidae: Epitoniiiin (Ankel 1926, 1938: 6-9, 1958, all as “Scala”, Fretter 1953, as “Clathrus”), Opalia (Bulnheim 1962a, 1962b), Epitoniuni (Nishiwaki 1964, Tochimoto 1967, Bulnheim 1968, Nishi- waki and Tochimoto 1 969), Opalia, Epitoniuni (Melone et al. 1978, 1980, latter as “Scala”), Epitoniuni (Robertson 1983a, 1983b, Collin 2000, as “Nituiiscala”). fanthinidae: Jantliina (Ankel 1926, 1930, Laursen 1953, Graham 1954, Wilson and Wilson 1956, as “lantliina”). Curiously, two epitoniid spe- cies have dimorphic spermatozeugmata (Nishiwaki and To- chimoto 1969).

The all-“mesogastropod” taxonomic pattern of occur- rence of spermatozeugmata is non-congruent with any other character known. Robertson ( 1985, 1989: table 1 ) and Collin (1997, 2000) reported the congruent occurrences of up to five non-homologoLis characters, suggesting that the taxa are related. I'he non-congruence of spermatozeugmata and their varied morphologies suggest that they are not homologous between the superfamilies listed above. The .same descriptive name in different taxa does not make a character homolo-

11

12

AMERICAN MALACOLOGICAL BULLETIN 23 • 1/2 • 2007

gous. The compound origin and nature of spermatozeug- mata could well have originated by homoplasy. The Tri- phoridae appear to have both spermatozeugmata and spermatophores, perhaps in different genera, but this needs confirmation.

The distinctive nurse cells of littorinids are believed to be parasperms. They are characteristic in being spherical to oblong, with or without projecting rods. In the Littoraria subgenus Pahistoriua the nurse cells are elongate or fusi- form, having a pseudotrich. Reid has not seen and reported attached eusperms since 1986, perhaps because since then he has consistently studied littorinid parasperms after fixation and shrinkage. As Reinke (1912), Woodard (1942b), Marcus and Marcus (1963), Jordan and Ramorino ( 1975), and Healy and Jamieson (1993) have observed, the acrosomes are at- tached weakly and the eusperms drop off easily. Perhaps all littorinids have these evanescent “spermatozeugmata”. A cypraeid [Erronea] appears to have dimorphic parasperms: vermitorm ones as well as spherical, littorinid-like “nurse cells.” Eusperms are semi-attached only to the latter, and these “could be considered a form of ‘spermatozeugmata’” (Healy 1986a).

Healy (1987) stated that the spermatozeugmata of Vi- riola (Triphoridae) are “very mobile”, but in 1990 he stated that they and those of Seila (Cerithiopsidae) are “capable of only slow movement.” Original observations on the behav- ior of living janthinoidean spermatozeugmata are included in Wilson and Wilson (1956), Nishiwaki and Tochimoto (1969), Melone et al. (1980), and Robertson (1983a). Uni- formly, there seems to be pseudocopulation. They are not “vigorously mobile,” swimming with “considerable speed” on their “relatively long journeys.”

Eretter (1953), Healy (1986b, 1987, 1990, 1994), and Niitzel (1998) believed that because of their spermatozeug- mata the Triphoridae and Cerithiopsidae are related to the Janthinoidea. They are similar in being large and containing numerous axonemes, but otherwise they are morphologi- cally different in the three groups. Niitzel (1998: 2) went so far as to suggest that spermatozeugmata are the “most con- vincing synapomorphy.” Overlooking Niitzel’s monograph, Collin (2000) reviewed most of the characters traditionally used to support the “Ptenoglossa” (including the Eulimidae but excluding the Architectonicidae, neither of which has known spermatozeugmata). There appear to be no other possible synapomorphies. Other literature on the Triphoroi- dea and Janthinoidea bears this out: Pruvot-Eol (1925, 1952), Johansson (1947, as “Scala”, 1953), Risbec (1953), Graham (1954), Marcus and Marcus (1963), Houston (1985), Houbrick ( 1987), and Collin (2004). “Ptenoglossan” radulae are diverse morphologically. If there is only one supposed “synapomorphy” linking two superfamilies, the validity of it may be questioned. Thus, I agree with Ponder

(1998) that it is improbable that Triphoroidea and Janthi- noidea are closely related.

SPERMATOPHORES

Records in non-stylommatophorans since Robertson (1989):

Neritoidea: Neritiliidae: Pisulina (Kano and Kase 2002), Neritilia (Kano et al 2001, Kano and Kase 2003). Neritidae: Clithon, Neritina [as “Neriptewn” and “Vittina”] (Starmiihl- ner 1970), Neritina, Septaria (Starmiihlner 1974), Clithon, Neritina, Septaria (Starmiihlner 1976), Clithon, Neritina (Starmiihlner 1983, 1984), Nerita, Neritilia, Neritina, Pu- perita (Starmiihlner 1988), Septaria (Haynes and Wawra 1989), Nerita, Neritina (Houston 1990), Theliostyla (Zehra and Perveen 1991), Nerita, Piiperita, Clithon, Neritina, Sep- taria, Neritilia (Starmiihlner 1993), Nerita (Sasaki 1998), Septaria (Haynes 2001), Septaria, Neritina (Haynes 2005). Phenacolepadidae: Cinnalepeta? (Sasaki 1998).

Campaniloidea: Plesiotrochidae: Plesiotrochiis (Hou- brick 1990). [Placement: Healy 1993].

Cerithioidea: Cerithiidae: Bittium? Bittiolnin (Houbrick 1993), Cerithinni (Houbrick 1992). Dialidae?: Diala? (Pon- der 1991). Melanopsidae (as “Thiaridae; Melanopsinae”): Fauniis (Houbrick 1991b). Paludomidae (including Thiari- dae, s. /. ): Lavigeria (Michel 1995), Tanganyicia (West 1997, Strong and Glaubrecht 2002), Tiphobia, Paramelania, Lim- notrochus, Chytra, Tanganyicia, Stanleya, Mysorelloides, Rey- nwndia, Lavigeria, Spekia, Stornisia (Glaubrecht and Strong 2004). Pleuroceridae: Elimia (Jones and Branson 1964, as “Mudalia”), Semisulcospira (Nakano and Nishiwaki 1989). Potamididae: Terebralia (Houbrick 1991a). Scaliolidae: Finella (Ponder 1994). Thiaridae, s. /.: Vinnndu (Michel 2004). Thiaridae, s. s.: Thiara (Glaubrecht and Strong 2004). ®

Turritellidae: Tiirritella (Kennedy 1995).

Pterotracheoidea: Atlantidae: Atlanta (Jamieson and Newman 1989). Carinariidae: Pterosoma (Lalli and Gilmer 1989). The “spermatophores” in Atlantidae reported by Tesch (reference in Robertson 1989) are actually the egg cases of a pleustonic insect, Halobates (Seapy 1996).

Vermetoidea: Vermetidae: Dendroponia, Serpidorbis, Vernietns (Calvo and Templado 2005).

Pyramidelloidea: Pyramidellidae: Pyramidella? (Ponder 1987), Boonea (Wise 2001), Fargoa (Robertson 1996), lolaea (Hori and Kuroda 2001 ), Odostoniella (Schander et al. 1999), Parthenina (Hori and Kuroda 2002).

Cavolinioidea: Cavoliniidae: Diacria (Lalli and Gilmer 1989). Limacinidae: Limacina (Lalli and Gilmer 1989).

|1

Clionoidea?: Pneumodermatidae?: Criicibranchaea? ’ (Lalli and Gilmer 1989).

GASTROPOD SPERMATOZEUGMATA AND SPERMATOPHORES

13

Hedylopsoidea: Parhedylidae: PoutoJjedylc, Unela (Poizat 1989).

Aeolidioidea: Aeolidiidae: Aeolidiella (Haase and Karls- son 2000).

Spermatophore presence can he difficult to ascertain, and a “spermatophore hursa” in a pallial oviduct may not always receive one (despite my assumption to the contrary in 1989: 362, A7). In Robertson (1989) I reported spermato- phores in the cerithioidean Litiopidae, hut Houhrick ob- served only bursae in them. Glaubrecht and Strong (2004) inferred the spermatophore-forming organ in male paludo- mid cerithioideans.

ACKNOWLEDGEMENTS

This study would not have been possible without access to the magnificent Ewell Sale Stewart Library of The Acad- emy of Natural Sciences of Philadelphia.

LITERATURE CITED

Ankel, W. E. 1926. Spermiozeugmenbildung durch atypische (apy- rene) und typische Spermien bei Scala und Jmithiiia. Verliaiid- hmgen der Deutschen Zoologischen Geselhchaft, 31 Jahresver- saminlung zu Kiel, Zoologischer Aiizeiger Supplemendmud 2: 193-202.

Ankel, W. E. 1930. Die atypische Spermatogenese von Janthina (Prosobranchia, Ptenoglossa). Zeitschrift fiir Zellforschiing und Mikroskopische Anatomie 11: 491-608.

Ankel, W. E. 1938. Beobachtungen an Prosobranchiern der schwedischen Westkiiste. Arkiv for Zoologi 30A(9): 1-27.

Ankel, W. E. 1958. Beobachtungen und Uberlegungen zur Mor- phogenese der atypischen Spermien von Scala clathnis L. Zoologischer Anzeiger 160: 261-276.

Battaglia, B. 1952. Ricerche sulla spermatogenesi atipica dei gaster- opodi prosobranchi. II.- Le cellule nutrici nella spermatogen- esi di Littorina neritoides L. (Gasteropodo Prosobranco). Bolkttino di Zoologia 19: 195-201.

Bedford, L. 1965. The histology and anatomy of the reproductive system of the littoral gastropod Bembiciwn nanum (Lamarck) (Earn. Littorinidae). Proceedings of the Linnean Society of New South Wales 90: 95-105.

Borkowski, T. V. 1971. Reproduction and reproductive periodici- ties of south Floridian Littorinidae (Gastropoda: Prosobran- chia). Bulletin of Marine Science 21: 826-840.

Buckland-Nicks, J. A. 1973. The fine structure of the spermatozoon of Littorina (Gastropoda: Prosobranchia), with special refer- ence to sperm motility. Zeitschrift fiir Zellforschiing und Mikroskopische Anatomie 144: 11-29. [Not seen]

Buckland-Nicks, LA. and F.-S. Chia. 1977. On the nurse cell and the spermatozeugma in Littorina sitkana. Cell and Tissue Re- search 179: 347-356. [Not seen]

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Accepted: 14 May 2007

Amer. Maine. Bull. 23: 17-32

A developmental perspective on evolutionary innovation in the radula of the predatory neogastropod family Muricidae"^

Gregory S. Herbert\ Didier Merle^, and Carlos S. Gallardo^

' Department of Geology, University of South Florida, Tampa, Florida, U.S.A., gherbert@cas.usf.edu

^ Unite de Paleontologie, Departement Histoire de la Terre, Museum national d’Histoire naturelle, UMR 5143 CNRS, 8, rue Buffon, 75005 Paris, France, dmerle@mnhn.fr

^ Institute de Zoologia, Universidad Austral de Chile, Casilla 567, Valdivia, Chile, cgallard@uach.cl

Abstract: The neogastropod family Muricidae includes a diverse set ot radular bauplane, including a beaked, three-dimensional form, a flattened-pentacusped form, and a third “dagger” type in which the central rachidian cusp is massive and elongate. Examination of the radular ontogenies of representatives of five muricid subfamilies reveals that several species undergo changes in radular form during ontogeny on a scale comparable to the evolutionary differences between higher taxa. The species Concholepas concholepas (Bruguiere, 1789) (Rapaninae) and Trophoii geversianiis (Pallas, 1774) (Trophoninae) begin ontogeny with a tliree-dimensional rachidian characteristic of the Ocenebrinae or Muricopsinae but end with the dagger rachidian typical of their respective subfamilies. Young individuals of Vitularia salebrosa (King and Broderip, 1832) (Muricopsinae?) also have a three-dimensional rachidian but shift to a double-dagger morphology by adulthood. Chicoreus (Phyllonotiis) pomiim (Gmelin, 1791) (Muricinae) has a typical flattened muricine rachidian as an adult but possesses a “buccinoid”-like rachidian just after hatching. Urosalpinx cinerea (Say, 1822) (Ocenebrinae), was unique among the species examined in exhibiting no ontogenetic changes in radular form. The occurrence of two radular bauplane within the same individual snail during ontogeny suggests great potential for rapid, convergent evolution of adult features through simple changes in developmental timing. A three-dimensional rachidian, for example, could be retained into adulthood through paedomorphosis in any lineage possessing the three-dimensional-to-dagger ontogeny. Systematic assignments of muricids based solely on radular features should be reexamined.

Key words: muricid, rachidian, bauplan, ontogeny, heterochrony

The radulae of a number of gastropod species undergo small to large-scale changes in the number, type, and struc- tural complexity of teeth between the pre-metamorphic lar- val stage, when the radula first forms, and maturation (ref- erences in Fujioka 1984a, Page and Willan 1988, Nybakken 1990, Waren 1990). A controversial but long-standing idea is that such changes in development play a dominant role in the evolutionary origins of new characters, or innovations, and, hence, in the origin ot higher taxa (reviewed in Gould 1977). Rather than requiring widespread alteration of struc- tural genes controlling morphology, innovation may derive simply through small-scale changes in regulatory genes con- trolling the rate and/or timing of development, i.e., het- erochrony. Size changes associated with heterochronic evo- lution may also provide a catalyst for extensive innovation throughout the organism. Changes in skeletal structure, for example, often evolve to compensate for the detrimental by-products of developmental miniaturization (see refer- ences in Hanken 1985, Hanken and Wake 1993).

With few exceptions (e.g., Guralnick and Lindberg

1999), however, molluscan biologists have yet to investigate radular evolution from the perspective of development. In the present study, we document radular ontogenies in sev- eral taxa belonging to the predatory gastropod family Mu- ricidae and examine the possibility that the evolution of developmental timing has played a central role in the re- peated origins of subfamily-level radular bauplane within the Muricidae. Phylogenetic studies are necessary to test the hypothesis that heterochronic mechanisms have been in- volved in the evolution of any particular structure, but on- togenetic analyses presented herein are a necessary first step.

BACKGROUND

Radular bauplane in the Muricidae

Two nomenclatural schemes have been utilized in the past to describe the basic structural types (referred to throughout this paper as “bauplane”) of the muricid rachid- ian teeth and to delineate the muricid subfamilies. The first

* From the symposium “Relationships of the Neogastropoda” presented at the meeting of the American Malacological Society, held 31 iuly- 4 August 2004 at Sanibel Island, Florida.

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system, established by Arakawa (1962, 1964, 1965) and Wu (1965, 1968, 1973), groups muricids according to the num- ber of cusps on the rachidian tooth. These workers subdi- vided muricid radulae into two classes — a complex “pen- tacusped” rachidian having a central cusp, two lateral cusps, and two marginal cusps, and a simplified “tricusped” rachid- ian having only the central and two lateral cusps (Fig. 1). Fujioka (1985a) later added a third class for taxa having a “monocusped” rachidian (i.e., only a central cusp) and rec- ognized a number of intermediate classes as well. As noted by Kool (1987), however, this system suffered from the rather arbitrary manner in which it was applied and is no longer used. For example, some authors counted only major cusps but others counted both cusps and denticles. The vari- able size, position, and number of cusps and denticles in different muricids make this system impossible to apply un- ambiguously (Kool 1987).

An alternative system, built upon in this paper, was developed by Yokes (1971), Radwin and D’Attilio (1971, 1976), and D’Attilio (1980, 1991) and is based largely upon variation in the morphology of the most prominent struc- ture on the rachidian — the central cusp — rather than the number of cusps on the tooth. In the “flattened” type, the rachidian is broad, with all the cusps lying in the same plane and the central cusp being slightly longer than either of the

laterals (Fig. 2M). This type occurs in almost all species presently assigned to the subfamilies Muricinae, Typhinae, Tripterotyphinae, and Haustrinae, as well as in many species currently assigned to the Trophoninae and the Miirexsid- Muricopsis genus group of the Muricopsinae (Radwin and D’Attilio 1971, 1976, Yokes 1971) (Figs. 2A-L). Because the Muricinae predate all other muricid subfamilies by at least 20 million years (see Yokes 1971, 1990, 1992, 1994, Garvie 1991, 1992, Marko and Yermeij 1999, Merle 1999, Yermeij and Carlson 2000), and because the anatomical condition of the Muricinae is likely primitive within the Muricidae (Ha- rasewych 1984), the flattened rachidian type of muricines and other muricids is presumably the plesiomorphic condi- tion for the family.

In a second type, which Yokes (1971) referred to as “three-dimensional” or “3-D,” the rachidian base is narrow (Fig. 3J) and rectangular, with a short, beak-like central cusp that projects up to 90 degrees away from the rachidian base and up to 45 degrees away from either lateral cusp (Fig. 31). Yokes (1971) and Radwin and D’Attilio (1971) have used terms such as “triangular harrow,” “cowl-like,” and “fang- like” to describe this type as well. A 3-D rachidian type characterizes Yokes’ (1971) Murexiella genus group of the Muricopsinae (= Favartia/Pygmaepterys subclade of Merle and Houart 2003), some species of Murkopsis (see Radwin

Figure 1. Pentacusped, tricusped, and monocusped rachidian bauplane of Arakawa (1962, 1964, 1965), Wu (1965, 1968, 1973), and Fujioka ( 1985a). A. The pentacusped rachidian characterized by the rapanine Neothais harpa (Conrad, 1837), locality: Maui, Hawaiian Islands, scale bar = 50 pm. B. The tricusped rachidian characterized by the ergalataxine Crania crassiilnata (Hedley, 1915), locality: Gulf of Carpentaria, northern Australia, scale bar = 50 pm. C. The monocusped rachidian characterized by the rapanine Dmpella data Blainville, 1832, Kauai, Hawaiian Islands, scale bar = 20 pm. D-F, penta-, tri-, and monocusped rachidia, modified from Fujioka (1985a, fig. 8).

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Figure 2. The “flattened” rachidian bauplan. A-B. The muricine Aspella indentata Carpenter, 1857, locality; Mira Mar, north of Manzanillo, Mexico, scale bar = 50 pm. C-D. The muricine Miirex brevispina var. macgilUvrayi Dohrn, 1862, locality: Papau New Guinea, scale bar = 100 pm. E-F. The “trophonine” Xymetwpsis bucdneiis (Lamarck, 1816), locality; Tierra del Fuego, Argentina, scale bar = 50 pm. G-H. The haustrine Haiistrmn haustoriiim (Gmelin, 1791), locality; New Zealand, scale bar = 100 pm. 1-1. The muricopsine Miircxsul octagoinis Quoy and Gaimard, 1833, locality: New Zealand, scale bar = 50 pm. K-L. The typhine Typhisahi graiidis (A. Adams, 1855), locality: Golfo tie Tehuantepec, Mexico, scale bar = 50 pm. M. The muricine rachidian bauplan, modified from Vokes (1971, fig. 2a).

and D’Attilio 1976), and the Ocenebra-Ocinebrina genus group of the Ocenebrinae {sensu Vermeij and Vokes 1997). Other muricid taxa that possess a beak-like central cusp include the putative-muricopsines Vitularia Swainson, 1840; Acanthotrophon Hertlein and Strong, 1951; and Bizetiella Radwin and D’Attilio, 1972; and the muricine Chicopinnatiis laqueatus (Sowerby, 1841) (Figs. 3A-H).

In addition to these two general categories of rachidia,

we recognize a third — a “dagger” rachidian — for muricids having a flattened rachidian but modified with a more mas- sive and consicierably more elongate central cusp and much weaker lateral and marginal cusps (Fig. 41). Fujioka’s (1985a) “monocusped” rachidian (Figs. 1C, IF) is an ex- treme form of this third type. Muricids possessing a dagger- type rachidian include most rapanine and ergalataxine spe- cies as well as the “I7nn's-like” ocenebrines (c.g., Nticella

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Florida, scale bar = 50 |am. C-D. The muricopsine Caribiella alveata (Kiener, 1842), locality. Discovery Bay, Jamaica, scale bar = 20 pm. E-F. The muricopsine Acanthotrophon sorenseni Hertlein and Strong, 1951, locality: Gulf of California, Mexico, scale bar = 20 pm. G-H. The muricine Chicopmnatiis laqueatus (Sowerby, 1841), locality: Orote Point, Guam, Oceania, scale bar = 20 pm. I-J. The three-dimensional rachidian bauplan, modified from Yokes (1971, fig. 2b).

Roding, 1798; Acanthimi Fischer de Waldheim, 1807; etc., see Vermeij and Yokes 1997), Trophon geversianus (Pallas, 1774) (type species of the type genus of the Trophoninae), and several prohlematic South American genera {e.g.. Chorus Gray, 1847; Xanthochoriis Fischer, 1884) (Figs. 4A-H).

Researchers have long relied upon these bauplane to assign species to subfamilies and reconstruct muricid phy- logeny based on the assumption that the radula is a conser- vative character complex relative to features of the shell and operculum (Yokes 1964, 1971, Radwin and D’Attilio 1971, 1976, fiouart 1992; see additional references in Kool 1987). Yokes (1971), for example, regarded the subfamily Ocene- brinae as phylogenetically nested within the Muricopsinae on the basis of both groups sharing a 3-D rachidian, despite the fact that other morphological features, such as those of the early shell whorls and operculum, support different re- lationships (Yokes 1971, Kool 1993a, Merle 1999, Yermeij and Carlson 2000). Using the same logic, Radwin and D’Attilio (1976) assigned the genus Vitiilaria to the Muri- copsinae, although shell and opercular characters are diffi-

cult to reconcile with this classification (see Yokes 1967, 1977, 1986). Most recently, Bouchet and Houart (1996) re- assigned the muricine Chicoreus gubbi (Reeve, 1849) to the Ocenebrinae as a new genus Chicocenebra Bouchet and Houart, 1996 based solely on its having a 3-D rachidian, even though its classification as a member of the Muricinae had not been questioned previously when only the shell morphology of this species was known.

Previous studies of radular development and evolution in muricids

If bauplan-level transformations in the muricid radula can occur as a result of small-scale changes in developmental timing, then radular features may be less conservative and, thus, less phylogenetically informative, than is currently thought. At present, the best answer to this question conies from a series of seminal papers by Fujioka (1982, 1984a, 1984b, 1985a, 1985b) on the radulae of the subfamilies Rapaninae and Ergalataxinae. Fujioka found that the lateral cusps, marginal cusps, and intermediate and marginal den-

DEVELOPMENT AND EVOLUTION OF THE MURICID RADULA

21

Figure 4. The dagger rachidian bauplan. A-B. The ergalataxine Cronia (Cronia) avellana (Reevem, 1846), locality: western Australia, scale bar = 100 pm. C-D. The rapanine Agnewia tritoniformis (Blainville, 1833), locality: Manly, New South Wales, Australia, scale bar = 50 pm. E-F. The ocenebrine Nucella ostrina Gould, 1852, locality: Monterey, California, scale bar = 20 pm. G-H. The ergalataxine? Xaiithochoriis cassidiformis Blainville, 1824, locality: Metri Bay, Chile, scale bar = 50 pm. I. The ergalataxine rachidian bauplan, modified from Fujioka (1985a, fig. 8).

tides in many species of these two subfamilies become pro- gressively atrophied during ontogeny, while the central cusp becomes longer and its base becomes wider.

Rapanines, however, begin ontogeny with a pen- tacusped rachidian and typically end at the pentacupsed or tricusped stage, whereas many of the ergalataxines studied begin ontogeny at the tricusped stage and end with a tri- cusped or monocusped rachidian. Under the assumption that new characters, such as atrophication, are only added to the end of ontogeny, Fujioka reasoned that the less atrophied pentacusped rachidia of rapanines is the relatively primitive condition and that the ergalataxine condition evolved hy peramorphic heterochrony (extended atrophication) of this ancestral ontogeny. Phylogenies published recently for the Rapaninae generally support this evolutionary scenario with ergalataxines depicted as a nested clade within the Rapaninae {i.e., Kool 1993a, Vermeij and Carlson 2000, but see Tan 2003).

More recently, DiSalvo ( 1988) and DiSalvo and Carriker (1994) documented ontogenetic changes in the rachidian tooth morphology of the rapanine muricid Coticholepas con- cholepas (Bruguiere, 1789). This species was found to change from a 3-D rachidian in early post-metamorphic animals to a dagger-type rachidian in small sub-adults. Although these workers did not comment on the evolutionary significance of their observations or document the exact size at which this transition occurs, it suggests to us that the 3-D and dagger rachidia in adults could potentially evolve rapidly from one to the other through heterochronic processes.

Focus of the present study

A 3-D rachidian was reported in none of the many juvenile rapanine species studied hy Fujioka, which makes its more recently reported occurrence in the rapanine Coticho- lepas suspect. Thus, the initial goal of the present study was to test the results of DiSalvo and Carriker by collecting new

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early post-larval specimens for Concholepas concholepas, verifying their taxonomic identity, and re-examining their early stage radulae using SEM.

The second focus of this paper was to examine the on- togenies of muricids outside of the Rapaninae and Ergal- ataxinae. With the exception of Houart’s (1992) illustration of the radular ontogeny of the muricine Chicoreus (Triplex) torrefactiis (Sowerby, 1841), there have been no attempts to document ontogenetic series in non-rapanine or non- ergalataxine muricids. Our present study of new and previ- ously collected material allows us to examine for the first time the ontogenies of species representing the Trophoni- nae, Ocenebrinae, a second species of the Muricinae, and a putative member of the Muricopsinae.

The third focus of this study was to investigate whether large-scale morphological shifts occur during the earliest on- togenetic stages of development, i.e., between larval meta- morphosis and juveniles around 10 mm shell length. Eujioka examined no radulae from early post-metamorphic stage individuals to small juveniles 3 mm in shell length. Eor more than half the species Eujioka studied, he examined no juve- niles smaller than 10 mm in length, and many of his smallest “juveniles” were greater than 25 mm in shell length. DiSal- vo’s work, in contrast, suggested that changes during earliest ontogeny [i.e., less than 10 mm shell length) may be sub- stantial (DiSalvo 1988, DiSalvo and Carriker 1994) and pos- sibly be related to changes in mode of predation and feeding behavior that can occur within this size range (see discussion section). This study documents the endpoints of the entire ontogenetic sequences beginning with rachidia in earliest post-metamorphic individuals, material permitting.

MATERIALS AND METHODS Radula preparation

Radulae were recovered from late juvenile and adult alcohol-preserved and dried specimens by dissolving dis- sected proboscis tissues in a concentrated solution of potas- sium hydroxide (KOH) for 1-3 days. Radular ribbons, visible with the naked eye, were collected with forceps, rinsed in a series of hot distilled-water washes, fixed to aluminum tabs with double-sided conductive tape, air dried, and gold coated (40 nm thickness) for scanning electron microscopy.

Radulae of early juvenile snails were recovered from dried and alcohol preserved specimens by gently crushing the larval shells in a shallow petri dish filled with a concen- trated solution of KOH. After two hours, the dish was heated to 90°C to reverse any precipitation of KOH crystals that might have formed on the radular ribbon. Because they were too small to be collected with forceps, radulae were removed from the KOH solution using a dropper partially filled with

warm distilled water. This was done to dilute the KOH so- lution collected with the radula and prevent later precipita- tion of KOH crystals on the radular ribbon during drying. The dilute KOH solution with the radula was then trans- ported by dropper through two rinses of hot distilled water in separate dishes for additional dilution. After two rinses, cleansed radulae were transferred in distilled water by drop- per to an aluminum tab coated with double-sided conduc- tive tape and air dried. Aluminum tabs with radulae were gold coated (40 nm thickness). All radulae were examined with an ISI DS-130 scanning electron microscope at the University of California, Davis’s Eacility for Advanced Instrumentation.

Radula Terminology

Throughout the remainder of this paper, we use the standard terminology illustrated by Radwin and D’Attilio (1976) and Kool (1987, 1993a) to refer to parts of the ra- chidian radular tooth.

RESULTS

Subfamily RAPANINAE

Concholepas concholepas (Bruguiere, 1789)

(Figs. 5A-F)

Material examined

Nineteen juvenile to sub-adult specimens of Concho- lepas concholepas ranging in shell length from 11 to 30 mm I were collected in March 2001 from Mehuin, 70 km north of Valdivia, Chile, and placed in 50% ethanol for later dissec- ' tion. An additional nine early post-metamorphic individuals ranging from 1.7 to 3.9 mm were captured between Septem- i ber 1999 and lanuary 2000 and between January 2000 and August 2000 from artificial collection plates installed at Las Cruces (coast of Santiago Province), Chile, and stored in 50% ethanol. ^

Ontogeny

In early post-metamorphic juveniles with shell lengths ranging from 1.7 to 3.9 mm, rachidian widths are 20 pm and have a 3-D rachidian morphology (Figs. 5A-D). Most have a ; single marginal denticle, although the largest individual in ; this class had two in one marginal area and one in the other ■ (Fig. 5A). The rachidian base end-point is marked by a |’ prominent marginal cusp that runs parallel to the rachidian base. Behind this marginal cusp is a shorter marginal cusp that is poorly developed and bud-like. The two cusps com- bined loosely resemble the double marginal cusps of the 1 Ocenebrinae. :■

In young animals with shell lengths between 11 and 13 j

DEVELOPMENT AND EVOLUTION OF THE MURICID RADULA

23

Figure 5. Radular ontogeny of the rapanine muricid Concholepas coiiciwlepas. A-D. Front and lateral views of rachidia of early post- metamorphic individuals ranging from 1.7 to 3.9 mm in shell length, locality: Las Cruces, Chile, scale bars = 10 pm. E-F. Front and lateral views of rachidia of a small juvenile with shell length of 15 mm, locality: Mehuin, Chile, scale bar = 50 pm.

mm, the rachidian tooth increases to approx. 100 pm in width and assumes the dagger-type morphology (Figs. 5E- F). The lateral cusps are turned outwardly at their distal ends, marginal cusp number increases to two or three, and outer lateral denticles begin to appear. The rachidian base

begins to develop a small, rounded lateral extension at both ends, which may be homologous with the hud-like second marginal cusp observed in the early post-metamorphic ju- veniles. The radular ontogeny of Concholepas is essentially stabilized at the dagger-type rachidian by a shell length of 1 1

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mm with little modification afterwards. Large adults may reach shell lengths upwards of 125 mm. Radulae of larger adult animals are figured elsewhere (Kool 1987, 1993b, Di- Salvo 1988) for comparison.

Remarks

DiSalvo (1988) and DiSalvo and Carriker (1994) illus- trated the rachidian tooth morphology of the pediveliger stage (1.6- 1.9 mm shell length) of the rapanine Concholepas coticholepas from neustonic pediveliger larvae reared from egg capsules captured at sea and hatched in the lab. The present study confirms their data showing this early stage to have a 3-D nrchidian. Possession of a 3-D-type rachidian during any stage of ontogeny in a rapanine is unusual be- cause previous studies of radular ontogeny of rapanines and a nested subclade within the Rapaninae, the Ergalataxinae, have reported only the flat rachidian type at any stage of development (Fujioka 1984a, 1984b, 1985a). However, given the size range of juvenile rapanines examined by Fujioka, it is possible that this stage of ontogeny was overlooked.

The transition from 3-D to dagger-type rachidian oc- curs after metamorphosis (DiSalvo, pers. comm.) and be- tween shell lengths of 4 and 11 mm (this study).

Subfamily TROPHONINAE

Trophon geversianus (Pallas, 1774)

(Figs. 6A-D)

Material examined

Research material for Trophon geversianus (Pallas, 1774) was obtained from dried and alcohol-preserved specimens in the personal collection of E. H. Yokes (Tulane University). This material included ten dried pre-hatched juveniles, all less than 2 mm in shell length, removed from a single egg capsule collected at a beach at Rio Grande, Tierra del Euego, Argentina. Although not yet hatched, the animals appear to have undergone metamorphosis, as indicated by the initia- tion of teleoconch (adult) sculpture. Also sampled were twenty alcohol-preserved adult specimens (all > 30mm) from Bahia El Pescador, south of Puerto Piramides, in the northeastern part of Golfo Nuevo, Argentina. This collection was also from the collection of E. H. Yokes.

Ontogeny

In post-metamorphic, pre-hatched juveniles of this spe- cies, the rachidian tooth is approx. 10 pm in width and has a 3-D morphology (Figs. 6A-B). The intermediate denticle is long and has an attachment site on the rachidian base inde- pendent of the lateral cusp but closer to the lateral than the central cusp. The basal region is rectangular and deep, with a strong marginal cusp and a second bud-like cusp closer to the radular ribbon.

Rachidian teeth in adults sampled are approx. 200 pm in width and exhibit a dagger- type morphology (Figs. 6C-D). The intermediate denticle is shorter than in early ontogeny (only one-fifth the height of the lateral cusp) and fused with the lateral cusp instead of having a separate attachment site on the rachidian base. The outer edges of the lateral cusps possess small serrations or outer lateral denticles. The basal end-point is rectangular but shallow and marked by a single marginal cusp. The second bud-like cusp of early ontogeny is obsolete or nearly so.

Remarks

Several authors have published line drawings and scan- ning electron micrographs of the radulae of adult Trophon geversianus, including Radwin and D’Attilio (1976), Kool ( 1993a), and Pastorino (2002). The present study is the first to document the morphology of the rachidia in early post- metamorphic individuals.

Subfamily OCENEBRINAE

Urosalpinx cinerea (Say, 1822)

(Eigs. 7A-D)

Material examined

Specimens of the ocenebrine muricid Urosalpinx cinerea were obtained in May 2001 from intertidal barnacle, mussel, and bryozoan-encrusted rocks from a jetty in San Erancisco Bay in Burlingame, California, USA. This species is native to the western Atlantic, but was introduced to the eastern Pa- cific nearly a century ago (Radwin and D’Attilio 1976). Twenty large adults (20-30 mm), including both males and females, were collected at the Burlingame locality and pre- served immediately in 75% ethanol for later dissection. An- other 10 individuals were placed in a single aquarium with recirculating seawater and provided with barnacles for food. Within days, adult snails deposited egg capsules on tank walls. After approx, six weeks, several hundred hatchlings (1. 5-2.0 mm shell length) emerged from the capsules as crawl-away juveniles and began drilling barnacles provided and cannibalizing one another by drilling. Approximately 50 hatchlings were harvested immediately and preserved in 75% ethanol.

Ontogeny

Animals ~2.0 mm in shell length possess rachidia 10 pm in width with a 3-D morphology (Pigs. 7A-B). Adult rachidia are larger (approx. 100 pm) but nearly identical in shape (Pigs. 7C-D).

Remarks

Radwin and D’Attilio (1976) and Kool (1993b) illus- trated the adult radula of Urosalphix cinerea, but there have

DEVELOPMENT AND EVOLUTION OE THE MURICID RADULA

25

Figure 6. Radular ontogeny of the trophonine muricid Twphon geversiauiis. A-B. Front and lateral views of rachidia ot early post- metamorphic (pre-hatched) individuals with shell lengths of 1.5 to 2.0 mm, locality: Tierra del Fuego, Argentina, scale bars = 10 pm. C-D. Front and lateral views of rachidia of adult specimen with shell length of 30 mm, locality: Golfo Nuevo, Argentina, scale bar = 100 pm.

been no studies of radular morphology during early ontog- eny. Carriker (1969) figured the mature radula of a subspe- cies, Urosalpinx cinerea var. etterae Baker, 1955, including eight scanning electron micrographs of radulae from various angles and two light micrographs of a rasping radula in the process of drilling a shell. The radula of this subspecies ap- pears to differ from that of the nominate form in having one or two extra marginal denticles.

Subfamily MURICOPSINAE?

Vitularia salebrosa (King and Broderip, 1832)

(Eigs. 8A-I)

Material examined

One dried juvenile specimen of Vitularia salebrosa (9.9 mm shell length) and ten dried adult specimens (30-50 mm shell length) from various tropical eastern Pacific localities were obtained from the personal research collections of E. H.

Vokes and G. 1. Vermeij. No other specimens of this species were available at the time the study was conducted.

Ontogeny

At the small juvenile stage (one 9.9 mm specimen), the rachidian is just over 5 pm in width and resembles the 3-D type in having a short, beak-like central cusp (Eigs. 8A-C). However, the rachidian base is Hat ratber than rectangular, a condition typically associated with flattened rachidia. Adja- cent to the central cusp are three pairs of short, conical “cusps.” The intermediate denticle is unusual for the 3-D type in projecting further from the rachidian base than the adjacent “lateral” cusp. The attachment site for the denticle is separate from the lateral cusp as in some muricopsines. The outermost cusp sits far from the margin endpoint, which curves into a pseudo-cusp. The rachidian ot a larger specimen (31 mm shell length) differs in having a slightly longer outermost cusp.

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Figure 7. Radular ontogeny of the ocenebrine miiricid Urosalpinx cinerea. A-B. Front and lateral views of rachidia of early post- metamorphic individuals with shell lengths of 1.5 to 2.0 mm, locality: San Francisco Bay, California, scale bar = 5 pm. C-D. Front and lateral views of rachidia of adult specimen with shell length of 25 mm, locality: San Francisco Bay, California, scale bar = 50 pm.

The largest individuals of Vitularia salehrosa available to us (40 to 50 mm specimens) failed to produce a radula in eight of the nine specimens we examined (89%). A single radula from a shell 50 mm in length shows rachidian teeth to be approx. 70 pm in width with long, tusk-like outermost (marginal?) cusps situated far from the base endpoint (Figs. 8G-I). The overall morphology of the central cusp is roughly of the 3-D rachidian type, but the outermost cusps are mas- sive and elongated as in the typical dagger-type rachidian.

Remarks

D’Attilio’s ( 1991 ) investigation of the radula of Vitularia salehrosa showed that this species has at least two different radular morphotypes, including a “normal” rachidian with seven cusps and a second rachidian with only three cusps, which he described as “extremely aberrant resembling noth- ing else known to me.” The latter morphotype has one sub- obsolete central cusp and two massive, incurved, tusk-like lateral cusps. D’Attilio did not provide information on the

sizes of the “normal” and “aberrant” rachidia, but the pres- ent data suggest they could represent end-members of a single ontogenetic sequence.

A second “aberrant” feature of the radula of Vitularia salehrosa is its occasional absence. D’Attilio (1991) reported that his own efforts to recover a radula from this species were successful only twice out of ten total attempts, which is similar to the success rate of one out of nine attempts re- ported in this study. This species is parasitic on oysters and attacks by pushing the proboscis between the valve margins, aided initially by drilling (G. S. Herbert and G. P. Dietl, pers. obs.). Attacking oysters at the edge could result in amputa- tion of the proboscis and radular loss when the oyster closes its valves. Another possibility is that the radula is used only to initiate an edge-drilled hole, and afterwards, the animal reabsorbs used teeth and stops forming new ones as it begins a parasitic existence. A major group of muricid parasites, the coralliophilines, lacks a radula, but these are obligate para- sites, whereas V. salehrosa is not.

DEVELOPMENl' AND EVOLUTION OF THE MURICID RADULA

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Figure 8. Radular ontogeny of the muricopsine? muricid Vitularin saicbwsa. A-C. Front and lateral views of racliida of small juvenile with shell length of 9.9 mm, Fig. C shows worn portion of radula presumably used in feeding, locality: Venado Beach, Panama, scale bar = 5 pm. D-F. Front and lateral views of rachidia of medium-sized juvenile 31 mm in shell length, locality: Venado Beach, Panama, scale bar = 20 pm. G-I. Front and lateral views of rachidia of mature specimen with shell length of 50 mm, locality: Panama, scale bar = 50 pm.

We place this species tentatively in the Muricopsinae after Radwin and D’Attilio (1976), although this assignment was and remains controversial due to shell and opercular similarities of this genus to some members of the Ocenebri- nae (Vokes 1986). In a cladistic analysis of the Muricidae based on morphological characters of the shell, ovocapsules,

and radula (D. Merle and G. S. Herbert, unpubl. obs.), Vitularia is unec]ui vocally placed outside the Muricopsinae and may prove to be a sister group ol the Ocenehrinae. Further phylogenetic investigations are necessary to clarify the systematic position of this problematic genus within the Muricidae.

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Subfomily MURICINAE

Chicoreus (Phyllonotus) ponium (Gmelin, 1791)

(Figs. 9A-D)

Material examined

Ten adult specimens of the muricine muricid Chicoreus (Phyllonotus) poniurn were collected from shallow subtidal seagrass beds in December 2002 in St. loseph’s Bay, Florida, USA and transferred to aquaria, where they were monitored and fed regularly. Females deposited communal masses of egg capsules, and offspring hatched within several weeks. Several hundred individuals (1.0-1. 5 mm shell length) hatched with a brief pediveliger stage of approx. 24 hours before absorbing the velum and using only the foot for lo- comotion. After a week, many juveniles began cannibalizing one another by drilling. These fully metamorphosed juve- niles were collected then and preserved in 75% ethanol. Five adults (60-70 mm shell length), including males and females.

were also preserved in ethanol after being relaxed in a 7.5% isotonic solution of magnesium chloride.

Ontogeny

Early post-metamorphic juveniles (1.0-1. 5 mm shell length) have a rachidian that is 15 pm wide and has only three cusps of similar lengths and lying in the same plane, with intermediate denticles usually absent (Figs. 9A-B). Each rachidian of an adult snail is roughly 200 pm in width, and has a wider base, more massive cusps, a new intermediate denticle, and a more elongate central cusp (Figs. 9C-D).

Remarks

Radwin and Wells (1968) and Radwin and D’Attilio (1976) figured line drawings of the mature radula of Chicoreus (Phyllonotus) ponnim. There are no other pub- lished illustrations of the early post-metamorphic or juvenile stage radular morphologies of this species.

Figure 9. Radular ontogeny of the muricine Chicoreus (Phyllonotus) ponnim. A-B. Front views of rachidia showing irregular presence of intermediate denticle in an early post-metamorphic individual with shell length of 1.5 mm, locality: St. loseph’s Bay, Florida, scale bar = 10 pm. C-D. Front and lateral views of rachidia of mature individual with shell length 63 mm, locality: St. Joseph’s Bay, Florida, scale bar = 100 pm.

DEVELOPMENT AND EVOLUTION OF THE MURICID RADULA

29

The absence of intermediate denticles and overall ap- pearance of the rachidian in young individuals of Phyllono- tus pomum gives this radular element a generalized neogas- tropod or “buccinoid” appearance. Similar rachidia occur in buccinids but also in olivids, melongenids, and turbinellid neogastropods, for example. Most interesting is the fact that this rachidian type has not been documented previously within the Muricidae. It is the first-known link in rachidian form between muricids and non-muricids.

DISCUSSION

Fujioka (1982, 1984a, 1984b, 1985a, 1985b) was the first to report that the muricid radula has the capacity to undergo

radical transformation between subfamily-level bauplane during ontogeny. He found, specifically, that the ergalatax- ine monocusped rachidian likely evolved through peramor- phic heterochrony, i.c., extension of an ancestral rapanid ontogeny characterized by progressive reduction of all but the central cusp. With the exception of a brief treatment ot the radular ontogeny of the muricine Chicoreus (Triplex) torrefactiis by Houart (1992), however, we have, until now, known nothing of the ontogenies of the radulae of other muricids.

The present study demonstrates that major transforma- tions between radular bauplane are nearly pervasive within the Muricidae, with transformations occurring in four ol the five subfamilies studied (Fig. 10). The ontogeny ot Phyllo- notiis pomum also links the seemingly disparate rachidian

Rachidian type:

Three-Dimensional

Dagger

Concholepas concholepas

Stages

Trophon geversianus

Young

Adult

Urosalpinx cinerea

Vitularia salebrosa

Rachidian type:

Buccinoid

Flattened

Chicoreus (Phyllonotus) pomum

Figure 10. Generalized patterns of radular ontogeny in five species of muricid gastropod. A-B. Most of the species studied herein begin ontogeny with the three-dimensional rachidian tooth as found in the Ocenebrinae and Muricopsinae but end with a dagger rachidian typical of the Rapaninae and Ergalataxinae as well as some Trophoninae and the Thais-Wke Ocenebrinae (;.e., Nucella, Acanthina, etc.). C-D. One species, Phyllonotus pomum, begins ontogeny with a generalized neogastropod (“buccinoid”) rachidian and shifts to a flattened, pentacusped rachidian with intermediate denticles typical of the Muricidae.

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morphologies of muricid and non-muricid neogastropods, and suggests that the primitive, pentacusped rachidian of the Muricidae evolved through extension of development. Equally interesting is the widespread occurrence of the 3-D rachidian that, until now, has been considered a defining trait of only the Muricopsinae and the Ocenebrinae. Data presented or reviewed in this paper demonstrate that the 3-D rachidian also occurs in at least some species of the Muricinae [Chicopinnatus and Chicoreus (Triplex)], Rapani- nae (Concholepas), and Trophoninae (Trophon).

Feeding observations for one of the species studied, the rapanine Concholepas concholepas, suggest that the 3-D ra- chidian could be a functional specialization for scraping against hard substrata, whereas the dagger-type rachidian that occurs later in ontogeny could be specialized for non- drilling modes of prey subjugation. Individuals of C. con- cholepas drill shelled prey or rasp rock surfaces in search of algae exclusively during their early post-metamorphosis stage, when the animal has the 3-D rachidian, but shift to attacking prey by stabbing or lacerating the soft parts with the radula through natural orifices by the time they reach 10 to 15 mm in shell length, when it has the dagger-type ra- chidian (Castilla et al. 1979, Paine and Suchanek 1983, DiSalvo 1982, 1988, Dye 1991, DiSalvo and Carriker 1994).

Muricids that possess a 3-D rachidian as adults, i.e., most species of the Ocenebrinae and Muricopsinae, also use drilling as their primary or exclusive mode of attack (G. S. Herbert, pers. obs.), again tying this rachidian bauplan to a specific drilling function. Ocenebrine and muricopsine mu- ricids also tend to be exceedingly small relative to other taxa in the Muricidae and, thus, most similar ecologically to ju- venile rather than large adult individuals of Concholepas con- cholepas. It may well be that young or small muricids lack the power to incapacitate prey using faster techniques, such as toxins or the brute force of chipping and prying (Herbert 2004), thus requiring slower methods for feeding such as drilling (Diet! and Herbert 2005) and, hence, a specialized radular type. Studies using computer modeling of the vari- ous radular morphologies within the Muricidae will be nec- essary to understand the exact functional basis of these radu- lar types.

It is striking that essentially the same ontogenetic trend toward a more flattened rachidian base and elongation of just one or two cusps occurs in species that differ at all stages in important details of cusp number, position, and shape. Such differences suggest that basic structural similarities (i.e., 3-D vs. flattened vs. dagger forms) among adult ra- chidia may be the result of independent innovation. Once the first 3-D-to-dagger ontogenetic trajectoiy evolved, any descendent lineages possessing this generalized ontogeny would have had the opportunity to retain the 3-D rachidian

into adulthood independently through evolutionary trunca- tion or a slowing of development. For these reasons, it is important to revisit past systematic assignments based on the bauplane that are the focus of the present study (see background section).

The repeated evolution of new traits, or innovations, has been a central theme of the muricid radiation (Vermeij 1998, 2001, Marko and Vermeij 1999, Vermeij and Carlson 2000), perhaps more so than in any other neogastropod clade. Although this phenomenon has been examined in the past from the standpoint of extrinsic factors, such as envi- ronmental conditions (e.g., temperature, productivity) and community dynamics (e.g., presence of incumbents, compe- tition intensity), the ontogenetic data presented herein point to a complementary process, namely the evolution of devel- opmental timing. Major morphological transformations during ontogeny increase the amount of phenotypic varia- tion in the population upon which natural selection can act and, thus, the intrinsic capacity of a species to create new characters or transform existing ones. They can also reduce genetic constraints on repetitive innovation by allowing morphologies or structures already present at one stage of development to shift to later (or earlier) stages through small-scale changes in the rate or timing of development.

ACKNOWLEDGMENTS

The authors would like to thank Emily Vokes, Geerat Vermeij, and Gregory Dietl for providing some of the ma- terial used in this study; Louis DiSalvo for providing useful information about the biology of Concholepas concholepas; Maroniae Oleson for assisting us with lab and field work; and Jerry Harasewych for kindly inviting us to participate in the Relationships of the Neogastropoda symposium at the 2004 American Malacological Society meeting at Sanibel. We also thank Geerat Vermeij and two anonymous reviewers for providing helpful comments on earlier drafts of this paper.

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Accepted: 26 September 2006

Amer. Make. Bull 23: 33-42

Phylogenetic relationships of the columbellid taxa Cotonopsis and Cosmioconcha (Neogastropoda: Buccinoidea: Columbellidae)'^

Helena Fortunate^

Smithsonian Tropical Research Institute, Center for Tropical Paleoecology and Anthropology, P.O. Box 0843-00153,

Balboa, Panama RP, fortunae@si.edu

Abstract: Phylogenetic reconstructions are still lacking for many molluscan groups, making evolutionary inferences much weaker. The genera Cotonopsis Olsson, 1942 and Cosmioconcha Dali, 1913 are part of the so called Strombina group, and as such have been used as models to study patterns of speciation and extinction brought about by the rise of the Central American gateway. Earlier work, based on a few species of each genus, pointed towards a very close relationship of these genera, which prompted a complete cladistic analysis, including all species of both genera to evaluate the level of relationship. Cladistic analyses based on shell morphology support the monophyly of the group composed by Cotonopsis -I- Cosmioconcha. Cotonopsis as currently defined is paraphyletic and includes Cosmio- concha. Cotonopsis (Tuirina) keeps its constituency and may retain its subgeneric status. Cotonopsis sensu stricto should he redefined to include part of Cosmioconcha. Cosmioconcha should be subdivided into two groups. One of these groups should be included in Cotonopsis sensu stricto. The second group should be given subgeneric status. Cotonopsis has a much earlier time of origination and most probably derives from Cosmioconcha. Obtained results give support to some of the evolutionaiy patterns documented earlier tor the Neogene molluscan faunas of tropical America and contribute to a better understanding of the Plio-Pleistocene divergence and turnover events related to the rise of the Panamanian land bridge.

Key words: gastropods, phylogeny, columbellids, morphology

The family Columbellidae is one of the most diverse and abundant shallow-water gastropod groups. The family has undergone rapid radiation, with over 400 species having evolved since the Danian Paleocene (Keen 1971, Abbott 1974, Radwin 1977, Tracey et al. 1993). The Strombina group sensu Jung, 1989, is one of the best known columbellid taxa, as it has been used as a model system to study evolutionary trends in species composition, diversity, and ecological pat- terns related to the Neogene rise of the Panama land bridge (VermeiJ 1978, Jackson et al. 1993, 1996, Fortunato 1998,

1999). Despite these studies, only recently have the phylo- genetic relationships of these taxa been investigated (de- Maintenon 1994, 1999, 2005, Fortunato and Jung 1995,

Fortunato 1998). Cotonopsis Olsson, 1942, and Cosmio- concha (Dali, 1913) are among the genus-level taxa that be- long to this group. They are abundant and include mostly recent species with a predominantly tropical American distribution.

In his latest revision, Jung (1989) included Cotonopsis but excluded Cosmioconcha from the Strombina group.

Work on the anatomy as well as preliminary cladistic analy- ses based on a subset of taxa (Fortunato and Jung 1995) confirmed Radwin’s ( 1977) hypothesis of a possible relation-

ship between Cosmioconcha and Strombina Morch, 1852 based on radular and shell morphology. These results pointed to a veiy close relationship between Cotonopsis and Cosmioconcha. The objective of this paper it is to investigate the phylogenetic relationships of these genera, including all fossil and living species, based on shell morphology in order to better understand their history and evolution.

MATERIALS AND METHODS

This analysis includes all known fossil and Recent spe- cies of the genera Cotonopsis and Cosmioconcha (Table 1). Cotonopsis is a very young genus (Jung 1989), with the first known species dating from the early Pliocene of Ecuador. Most of the diversity within Cotonopsis developed during the Plio-Pleistocene turnover, around the time of formation of the Panama land barrier. Cotonopsis includes 18 species grouped in two subgenera. Only one species is known ex- clusively as a fossil. Of the 17 living species, 13 inhabit the eastern Pacific basin (Jung 1989), two were reported from the Caribbean region (Houbrick 1983, Petuch 1988, Fortunato 2002b), one was described from the west coast of Afi'ica (Emer- son 1993), and a fourth species was found in the Andaman Sea (Kosuge et al. 1998, Kronenberg and DeJeker 1998, 1999).

From the symposium “Relationships of the Neogastropoda” presented at the meeting of the American Malacological Society, held 31 July-4 August 2004 at Sanibel Island, Florida.

' Current address: Institut ftir Geowissenchaften Universitat Kiel, Ludwig-Meyn-Strasse 10, D-24118 Kiel, Germany.

33

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AMERICAN MALACOLOGICAL BULLETIN 23 • 1/2 • 2007

Table 1. Taxa included in the phylogenetic analyses. Stratigraphic and geographic ranges are given only for ingroup taxa. LM, late Miocene; EP, early Pliocene; LP, late Pliocene. See Table 2 for more information on species of the genus Cosmioconcha. Extinct species indicated by

an n.

Species

Status

Stratigraphic

range

Geographic range

Nassarius hiteostoma (Broderip & Sowerby,

Outgroup

1829)

Nassarius antillannn Orbigny, 1842 Canthanis ringens (Reeve, 1846)

Outgroup

Outgroup

Latirus amceutncus (Reeve, 1847) Cotonopsis {Cotonopsis) panacostaricensis

Outgroup Type of species of

LP — Recent

Eastern Pacific (Costa

(Olsson, 1942)

Cotonopsis

Rica — Colombia)

Cotonopsis (Cotonopsis) edentida (Dali, 1908)

Olsson, 1942

Recent

Eastern Pacific (G. of

Cotonopsis (Cotonopsis) argentea (Houbrick,

Recent

California — Panama) Caribbean (Dominican Republic)

1983)

Cotonopsis (Cotonopsis) crassiparva (lung.

Recent

Eastern Pacific (Galapagos Is.)

1989)

Cotonopsis (Cotonopsis) deroyae (Emerson 8t

Recent

Eastern Pacific (Galapagos Is.)

D’Attilio, 1969)

Cotonopsis (Cotonopsis) aff. deroyae (Emerson

Recent

Eastern Pacific (Peru)

& D’Attilio, 1969)

Cotonopsis (Cotonopsis) esnieraldensis (Olsson,

EP

Eastern Pacific (Ecuador)

1964)*

Cotonopsis (Cotonopsis) jaliscana (lung, 1989)

Recent

Eastern Pacific (Mexico)

Cotonopsis (Cotonopsis) inendozana (Shasky,

Recent

Eastern Pacific (Mexico — El Salvador)

1970)

Cotonopsis (Cotonopsis) skoghindae (lung,

Recent

Eastern Pacific (Gulf of California)

1989)

Cotonopsis (Cotonopsis) siiteri (lung, 1989)

Recent

Eastern Pacific (Gulf of

Cotonopsis (Cotonopsis) aff. suteri (lung,

Recent

California — Mexico)

Eastern Pacific (Mexico — Costa Rica)

1989)

Cotonopsis (Cotonopsis) phiiketensis (Kosuge,

Recent

Andaman Sea (Phuket Is.)

Roussy & Muangman, 1998)

Cotonopsis (Cotonopsis) lindae (Petuch, 1988)

Recent

Caribbean (Barbados)

Cotonopsis (Cotonopsis) njonfilsi Emerson,

Recent

Western Africa (Senegal)

1993

Cotonopsis (Turrina) hirundo (Gaskoin, 1852)

Pleistocene —

Eastern Pacific (Gulf of

Cotonopsis (Turrina) radwini (lung, 1989)

Recent

Recent

California — Ecuador)

Eastern Pacific (Mexico — Panama)

Cotonopsis (Turrina) tiirrita (G. B. Sowerby I,

Recent

Eastern Pacific (El Salvador —

1832)

Cosmioconcha modesta (Powys, 1835)

Type species of

Recent

Colombia)

Eastern Pacific (El Salvador —

Cosmioconcha pahneri (Dali, 1913)

Cosmioconcha Dali, 1913

LM — Recent

Ecuador)

Eastern Pacific (Gulf of

Cosmioconcha parvula (Dali, 1913)

Recent

California — Panama) Eastern Pacific (Gulf of

Cosmioconcha rehderi (Hertlein & Strong,

Recent

Galifornia — Panama )

Eastern Pacific (Mexico — Ecuador)

1951)

Cosmioconcha pergracilis (Dali, 1913)

Recent

Eastern Pacific (Mexico)

Cosmioconcha nitens (C. B. Adams, 1850)

Recent

Caribbean (Cuba, Puerto Rico)

Cosmioconcha calliglypta (Dali & Simpson,

Recent

Caribbean (Florida, Texas, Puerto

1901)

Rico)

PHYLOGENY OF COTONOPSIS AND COSMIOCONCHA

35

The earliest known Cosmioconclia species dates from the middle Miocene. Cosinioconcha was first described as a subgemis of Amphissa H. & A. Adams, 1853 (Dali, 1913). Radwin (1978) elevated it to generic rank. Cosmiocouchn includes seven described species, two inhabiting the Caribbean Sea and five the eastern Pacific region (Table 2, Figs. 1-2). Recent patterns of diversity and abundance of this taxon indicate a radiation similar to other paciphile genera.

Outgroup taxa were selected from Nassariidae, Buccini- dae, and Fasciolariidae. Four common taxa from three buc- cinoidean families were selected as outgroups: Nassariidae — Nassarius luteostoma (Broderip & Sowerby, 1829) and N. autillarum d’Orbigny, 1842; Buccinidae — CatJtlmrus ringens (Reeve, 1846); Fasciolariidae — Latirus concentricus (Reeve, 1847). These taxa were selected based on availability and not on the presumption of close phylogenetic relationship.

Forty- two qualitative characters were identified (Ap- pendix 1). Shell sculpture is one of the most characteristic elements of this group, and provides numerous diagnostic characters. Fourteen characters code for type and sculptural details of the teleoconch and body whorl. Presence of shoul- der, constriction, inflation, and angulation of the whorls, as well as presence and strength of humps were also coded. Apertural elements (thickness, denticles, apertural and colu- mellar calluses and plicae, parietal ridge) used in traditional taxonomy of this group of gastropods are included here as well. Other characters are general shell shape, type of spire, type and depth of suture, and the relation between the total height and the height of the body whorl. All taxa were coded from direct observation.

MacClade 3.0 (Maddison and Maddison 1992) was used to create the data matrix of 25 taxa and 42 morphological characters (Appendix 2). The heuristic search in PAUP 4.0b 10 (Swofford 2001 ) was used for the analyses, using a random addi- tion sequence with ten repli- cate searches performed. All characters were unordered and weighted equally. Glade support was assessed through a bootstrap procedure (100 bootstrap repli- cates with 10 random addition sequences). Tree support was determined using Bremer decay analysis (Bremer 1994) in which progressively longer trees are saved and their consensus calcu- lated in order to see how many more steps are required to col- lapse branches.

RESULTS

Cladistic analyses of the data matrix in Appendix 2 yielded six most-parsimonious trees (F=218 steps, Cl=0.303, Rl=0.513, and RG=0.155). Only the strict consensus tree (Fig. 3) is presented here (the 50% majority rule consensus tree shows exactly the same topology).

The ingroup is monophyletic in all trees, consisting of a single clade grouping all Cotonopsis and Cosmioconclia spe- cies. This clade is defined by fusiform shells with high spire and mostly un-sculptured earlier teleoconch whorls, body whorl mostly un-sculptured, apertures with moderately thickened outer lips, and a well developed, recurved anterior canal.

The genus Cotonopsis, as traditionally constructed, is paraphyletic and includes the polyphyletic Cosmioconclia. The subgenus Cotonopsis [Turrina) emerges as a monophy- letic crown group. Cosmioconclia species are grouped in two separate clades within Cotonopsis. One clade, which contains the type species of Cosmioconclia, emerges in an unresolved trichotomy with a small clade containing the type species of Cotonopsis and a large clade that includes Cotonopsis, the remaining Cosmioconclia, and Cotonopsis (Tiirriiui).

All species assigned to Cotonopsis sensu stricto emerge as a grade that also includes a small clade of three species of Cosmioconclia, including its type species. These species have mostly stout shells with axially sculptured late spire whorls and well defined cords at the base of the body whorl. I'hey have broad apertures with denticles and thin outer lip edges.

This grade (all Cotonopsis sensu stricto -t- three Cosniio- conclia taxa) has several smaller groupings. Its base is weakly resolved with several Cotonopsis species branching succes-

Table 2. Synopsis of species belonging to the genus Cosmioconclia Dali, 1913. *, type species.

Genus

Species

Author & Year

Synonyms

Cosmioconclia

modesta *

Powys, 1835

Buccinum modestiim Powys,

1835; Stromhina lacvistriata Li, 1930

palmeri

Dali, 1913

parvula

Dali, 1913

relideri

Hertlein & Strong, 1951

pergmcilis

Dali, 1913

nitens

C. B. Adams, 1850

Fusiis nitens C. B. Adams, 1850;

Coliimbella (Astyris) perpicta Dali 8c Simpson, 1901; Mitrclla perpicta (Dali 8c Simpson) Woodring, 1928

calliglypta

Dali & Simpson, 1901

Anacliis calliglypta Dali 8c

Simpson, 1901

36

AMERICAN MALACOLOGICAL BULLETIN

23 • 1/2 • 2007

10mm

5mm

L

Figure 1. Species of Cosmwconcha Dali, 1913. A-C Cosnnocoucha modesta (NMB 17442). D-F Cosmiocoucha pahnen (NMB 17793) G-1 Cosmiocoiicha parvida (NMB H18181 ). 1-L Cosmwconcha rehden (NMB H18182). M-O Cosmwconcha mtens (NMB 18567). Views are front, rear, and from right side. All specimens belong to the Gibson-Smith Recent collection housed at the Naturhistorisches Museum Basel,

Switzerland.

PHYLOGENY OF COTONOPSIS AND COSMIOCONCHA

37

Figure 2. Species of Cosmiocondia Dali, 1913, protoconchs. A-R Cosniioconcha modesta. C-D Cosmioconcha palweri. E-F Cosmioconcha parvula. G-H Cosmioconcha rehderi. I-l Cosmioconcha nitens. Same specimens as in Fig. 1.

sively. Among these are C. inouftlsi and C. lindae, an African and a Caribbean species respectively. The next branch has two small subclades, one with three Cosmioconcha and a second one joining two Cotonopsis taxa. Next to diverge is the Cotonopsis living in the Andaman Sea, followed by an- other small group formed by two eastern Pacific Cotonopsis. The last grouping of this grade joins a Caribbean and two eastern Pacific Cotonopsis taxa.

The two other groups are sister clades and are located as crown groups. One of these clades groups, but does not resolve, the three Cotonopsis (Tiirrina) taxa. The second group is composed by four Cosniioconcha species, among which appear the two Caribbean species. Species of the crown clades have slender shells, almost no axial ornamen- tation on the spire whorls, and narrower apertures with a

thicker edge. The species of Cosmioconcha have more convex spires, numerous denticles on the ap>erture, and a collar like band below the spire suture. Cotonopsis {Tnrrina) taxa are characterized by taller, straight-sided shells and a well de- veloped thickening behind the outer lip. Results of the Bremer decay analysis are plotted onto the strict consensus tree (Fig. 3).

The first round resulted in 284 trees with 219 steps or less. These trees support the monophyly of the entire in- group as well as the monophyly of both crown clades, i.e. Cotonopsis (Tnrrina) and the four Cosmioconcha species. Two small groups that appear among the basal branches (one joining three eastern Pacific Cosmioconcha, and another with two Cotonopsis sensu stricto taxa) are equally supported here. The second round of the decay yielded 6,278 trees, 220

38

AMERICAN MALACOLOGICAL BULLETIN 23 • 1/2 • 2007

Latirus concentricus Cantharus ringens Nassarius luteostoma Nassarius antillarum Cotonopsis (Cotonopsis) jaliscana Cotonopsis (Cotonopsis) att.deroyae Cotonopsis (Cotonopsis) suteri Cotonopsis (Cotonopsis) monfiist Cotonopsis (Cotonopsis) lindae Cotonopsis (Cotonopsis) edentula Cotonopsis (Cotonopsis) crassiparva Cotonopsis (Cotonopsis) panacostaricensis Cosmiconcha palmeri Cosmiconcha modesta Cosmiconcha rehderi Cotonopsis (Cotonopsis) phuketensis Cotonopsis (Cotonopsis) esmeraidensis Cotonopsis (Cotonopsis) aft. suteri Cotonopsis (Cotonopsis) argentea Cotonopsis (Cotonopsis) mendozana Cotonopsis (Cotonopsis) skoglundae Cotonopsis (Cotonopsis) deroyae Cotonopsis (Turrina) turrita Cotonopsis (Turrina) hirundo Cotonopsis (Turrina) radwini Cosmiconcha pergraciiis Cosmiconcha parvula Cosmiconcha nitens Cosmiconcha calliglypta

Figure 3. Strict consensus tree of the six most parsimonious cladograms. Numbers below branch nodes are Bremer support values, i.e., the number of extra steps necessary to collapse that node. Nodes without values collapse with one extra step. Numbers above branch nodes are bootstrap support values for that node.

steps or less. The ingroup is still monophyletic but only the group with the two Caribbean Cosmioconcha species is sup- ported. The third round yielded 117,655 trees, 221 steps or less. It supports the monophyly of the ingroup but there is no resolution. The fourth round of decay analyses overflowed the memory with over 500,000 trees. The monophyly of the ingroup is still supported here. The bootstrap support for some of the clades is plotted in the strict consensus tree (Fig. 3).

DISCUSSION

The main objective of this work was to re-evaluate all species that have traditionally been assigned to the genera Cotonopsis and Cosmioconcha, in order to assess their rela- tionships and the true constituency of these genera. Earlier analyses based on a selected subset of species from each of these genera (Fortunato and lung 1995) suggested a close relationship of these taxa, confirming Radwin’s (1977) hy- pothesis of a relationship between Cosmioconcha and the

Strombina group of which Cotonopsis is part (Jung 1989). Our objective was to test this relationship, including in the analysis all known species currently included in both genera.

The results of this study indicate that Cotonopsis -l- Cos- mioconcha form a monophyletic group. Cotonopsis as it was initially defined by Jung ( 1989) is paraphyletic and contains Cosmioconcha. Of the two subgenera of Cotonopsis, only Cotonopsis (Turrina) is monophyletic and retains its entire constituency. Cotonopsis sensu stricto, as currently con- structed, is paraphyletic. Its status as a monophyletic taxon could be restored only by synonymizing both Cosmioconcha and Turrina. Alternatively, inclusion of more closely related outgroups might alter the rooting of the tree. Rooting at Position A (Fig. 3) would be required to retain monophyletic Cotonopsis and Cosmioconcha as sister taxa although the ma- jority of species currently assigned to Cosmioconcha would still emerge Cotonopsis.

Species assigned to Cosmioconcha are divided into two groups. The first group is composed of three eastern Pacific

PHYLOGENY OF COTONOPSIS AND COSMIOCONCHA

39

species, and is located near the base of the tree, among Cotonopsis sensu stricto taxa. This group includes the type species, Cosmioconcha modesta. The second group is one of the crown subclades and unites two Caribbean and two east- ern Pacific species. This group is sister of Cotonopsis {Tiirrina).

Within the grade Cotonopsis sensu stricto, C. monftlsi, a deep water species from West Africa, and C. lindne, a shallow water species from Barbados, form adjacent branches but are flanked by eastern Pacific species from California, Mexico, and Peru that have no known fossil record. It is tempting to speculate about the possible existence of geminate pairs [i.e., closely related taxa separated by a barrier (Jordan 1908)] among the extinct fossil ancestors of these taxa. These rela- tionships also suggest an earlier radiation of American spe- cies, probably from the eastern Pacific towards the Atlantic before the closure of the Panamanian Strait. Unfortunately, none of these species have a known fossil record which could help calibrate the time of such radiation. However, both the geographic distribution of these species, and the fact that several of the following taxa (within the context of this tree topology) have fossils dating back to the middle Miocene [i.e., Cosmioconcha palmeri (Dali, 1913)) suggest that such a radiation may have taken place during the middle Miocene. It is also reasonable to assume the possible existence of fossil lineages yet to be found. Molecular studies could provide an alternative tool to elucidate these relationships.

Three eastern Pacific species of Cosmioconcha^ including the type species, emerge as a clade. The stem species, C. palmeri, has the oldest fossil record of all the species in this study, being known from the middle Miocene deposits of Darien [Radwin, 1977; Panama Paleontological Project (PPP) data]. Based on these results, it is reasonable to as- sume that Cotonopsis sensu stricto is much older than pos- tulated by Jung (1989) in his revision of the Strombina group. Jung indicated an early Pliocene age for Cotonopsis, based on the occurrence of Cotonopsis esmeraldensis (Olsson, 1964) in the early Pliocene of Ecuador. Results of the present analysis indicate that Cosmioconcha is part of Cotonopsis sensu stricto, thus moving the time of origination of this genus most probably to middle Miocene.

Another closely related small clade unites the Recent Cotonopsis (Cotonopsis) crassiparva and a late Pliocene Coto- nopsis (Cotonopsis) panacostaricensis (Olsson, 1942), the type species of Cotonopsis.

Cotonopsis (Cotonopsis) phnketensis (Kosuge et ah, 1998), a shallow water species from the Andaman Sea, is the second species in this genus with a distribution outside of tropical America. There are infreciuent reports of plankto- trophic larvae crossing the central Pacific barrier (Scheltema, 1978). Most Cotonopsis have planktotrophic larvae (excep- tions are C. jaliscana, C. esmeraldensis, and C. argentea; au- thorities in Table 1 ) able to spend a considerable amount of time in the plankton (Fortunate 2002a). Again, the lack of

fossil data precludes the dating of this dispersal event. Nev- ertheless, the presence of an early Pliocene species within a sister clade indicates that it may date back to the early Pliocene, at the very least. Here again, molecular data would be useful to help resolve these events.

The next clade comprises Cotonopsis esmeraldensis (early Pliocene of Ecuador) and a recent eastern Pacific species, C. aff snteri. This is probably a case of speciation with a switch in developmental mode, as C. esmeraldensis is a non- planktotroph whereas its sister species has planktonic lar- vae. Cotonopsis esmeraldensis is the only extinct taxon in the analyzed data set. The basal species of this clade, C. pnkhe- tensis, is also a planktotroph. A trans-isthmian event in the history of the group is documented in the next branch of this phylogenetic tree. Cotonopsis argentea, a non planktotroph taxon found in deep water of the Dominican Republic coast is the sister taxon of two eastern Pacific species (C. mendo- zana and C. skoghmdae).

The crown of the tree is composed by two subclades with a relatively strong Bremer and bootstrap support. The stem taxon is Cotonopsis deroyae. One of the groups includes the three Cotonopsis (Tnrrina) species, confirming the com- position and monophyly of this subgenus. The second group, composed by four Cosmioconcha species, documents another trans-isthmian event: C. nitens and C. calliglypta are shallow water taxa inhabiting the Caribbean Sea that di- verged from an eastern Pacific taxon.

Based on the obtained results and the phylogenetic re- construction presented here, Cotonopsis sensu stricto, as presently understood, represents a grade that includes sev- eral Cosmioconcha taxa (i.e., C. palmeri, C. modesta, and C. rehderi), among them the type species of Cosntioconcha. All have stout shells with high spires, axially sculptured early teleoconch whorls, body whorls with strong cords on the base, and wide apertures.

The four “Cosmioconcha" species that constitute one of the crown groups of the tree are not closely related to the type species of Cosmioconcha. These species are characterized by smaller fusiform shells, absence of sculpture on the early teleoconch, absent or weak cords on the body whorl, and narrow apertures. The character “presence of a collar-like band below the suture”, traditionally used to unify Cosmio- concha taxa is not reliable and should not be given more value than any other morphological character.

Cotonopsis is a taxon that reflects the pulse of origina- tion that occurred in the eastern Pacific at the Pleio- Pliocene boundary. Most of the recognized taxa originated during the last two million years, probably along the shallow waters of the eastern Pacific coast. Unfortunately, the stratigraphic rec- ord of the eastern Pacific region is not very well preserved (Coates etal. 1992, Jackson et al. 1993, 1996) and there is no fossil record for most of the known species. Cosmioconcha also originated in this region and has a fossil record that

40

AMERICAN MALACOLOGICAL BULLETIN 23 • 1/2 • 2007

dates back to the middle Miocene. Based on the phylogenetic reconstruction presented here it is reasonable to assume that Cotonopsis derives from a Cosmioconcha-like ancestor. The group then radiates and speciates with the documented in- crease in species diversity towards the recent, a pattern well documented for the entire Strombina group (lung 1989, Jack- son et al. 1993, 1996).

The Strombina group has been used as a model system to document patterns of diversification during the Neogene rise of the Panamanian isthmus. Phylogenetic inferences have started to give historical support to earlier studies. The taxa studied here are part of this group and the results con- firm the validity of the evolutionaiy patterns documented earlier (Jackson et al. 1993, 1996, Portunato and Jung 1995, Portunato 1998, 1999). It is also reasonable to assume the existence of fossil lineages and even Recent taxa yet to be found that could contribute to a better understanding of the natural history of the molluscan fauna of the region and its relationships.

AKNOWLEDGMENTS

This work was presented at the molluscan phylogeny symposium organized by M. G. Harasewych during the AMS meetings, 2004. 1 thank the institutions that loaned materials for this study. A. Velarde, J. Jara, M. Alvarez, P. Rodriguez, and the Urraca’s crew helped with field collections and labo- ratory work. STRTs digital and SEM laboratory personnel helped with the illustrations. This work was supported by the Scholarly Studies and the Walcott programs of the Smithsonian Institution.

LITERATURE CITED

Abbott, R. T. 1974. American Seashells, 2”''* Edition. Van Nostrand Reinhold Company, New York.

Bremer, K. 1994. Branch support and tree stability. Cladistics 10: 295-304.

Coates, A. G., J. B. C. lackson, L. S. Collins, T. M. Cronin, H. J. Dowsett, L. M. Bybell, P. Jung, and J. A. Obando. 1992. Clo- sure of the Isthmus of Panama: The near-shore record ot Costa Rica and western Panama. Geological Society of America Bulletin 104: 814-828.

deMaintenon, M. J. 1994. Evolution of Columhella (Neogas- tropoda: Columbellidae) in the Neogene American tropics. Geological Society of America Abstract with Programs 26: A-53. deMaintenon, M. ). 1999. Phylogenetic analysis of the Columbel- lidae (Mollusca: Neogastropoda) and the evolution of her- bivory. Invertebrate Biology 118: 258-288. deMaintenon, M. J. 2005. Phylogenetic relationships of the tropical American columbellid taxa Conella, Eiirypyrene, and Para- mctaria (Gastropoda: Neogastropoda), fournal of Paleontology 79: 497-508.

Emerson, W. K. 1993. A new species of columbellid gastropod from the old world tropics. The Nautilus 106: 147-151.

Portunato, H. 1998. Reconciling observed patterns of temporal oc- currence with cladistic hypotheses of phylogenetic relation- ship. American Malacological Bulletin 14: 191-200.

Portunato, H. 1999. Biogeography and the tempo of speciation in strombinid gastropods. Abstracts, 7'*" Congress of the Euro- pean Society for Evolutionary Biology, II: 107.

Portunato, H. 2002a. Reproduction and larval development of the Strombina group (Buccinoidea: Columbellidae) and related gastropods: Testing the use of the larval shell for inference of development in fossil species. Bollettino Malacologico 4: 111-126.

Portunato, H. 2002b. The systematic position of Strombina {Coto- nopsis) Undae Petuch, 1988 (Gastropoda: Columbellidae). The Nautilus 116: 59-61.

Fortunado, H. and P. lung. 1995. The Strombina-group (Neogas- tropoda: Columbellidae): A case study of evolution in the neotropics. Geological Society of America Abstracts with Pro- grams 27: A-52.

Eloubrick, R. S. 1983. A new Strombina species (Gastropoda: Pro- sobranchia) from the tropical western Atlantic. Proceedings of the Biological Society of Washington 96: 349-354.

lackson, I. B. C., P. Jung, and H. Portunato. 1996. Paciphilia revis- ited: Transisthmian evolution of the Strombina-gvoup (Gas- tropoda: Columbellidae). In: I. B. C. Jackson, A. F. Budd, and A. G. Coates, eds.. Evolution and Environments in Tropical America. The University of Chicago Press, Chicago. Pp. 234-270.

lackson, I. B. C., P. lung, A. G. Coates, and L. S. Collins. 1993. Diversity and extinction of tropical American mollusks and emergence of the Isthmus of Panama. Science 260: 1624-1626.

Jordan, D. S. 1908. The law of geminate species. American Natu- ralist 42: 73-80.

lung, P. 1989. Revision of the Strombina-group (Gastropoda: Co- lumbellidae), fossil and living. Distribution, biostratigraphy and systematics. Memoires Suisses de Paleontology 111: 1-298.

Keen, A. M. 1971. Sea Shells of Tropical West America, 2"'^ Edition. Stanford University Press, Stanford, California.

Kosuge, S., P. H. Roussy, and P. P. Muangman. 1998. Report on the fauna of Thailand ( 1 ) with the description of a new species (Columbellidae and Buccinidae). Bidletin of the Institute of Malacology of Tokyo 3: 75-76.

Kronenberg, G. C. and H. Dekker. 1998. A new species of Coto- nopsis Olsson, 1942, from an unexpected locality (Gastropoda Prosobranchia: Columbellidae). Vita Marina 45: 11-16.

Kronenberg, G. G. and H. Dekker. 1999. Cotonopsis vanwalleghemi Kronenberg & Dekker, 1998, a junior synonym of Strombina phuketensis Kosuge, Roussy & Muangman, 1998, with some notes on the generic position and colour pattern (Gastropoda Prosobranchia: Columbellidae). Vita Marina 46: 69-72.

Maddison, W. P. and D. R. Maddison. 1992. MacClade: Analysis of phylogeny and character evolution. Version 3.0. Sinauer Asso- ciates, Sunderland, Massachusetts.

Petuch, E. J. 1988. Neogene History of Tropical American Mollusks. The Coastal Education and Research Foundation (CERF), Charlottesville, Virginia.

Radwin, G. E. 1977. The family Columbellidae in the western At- lantic. The Veliger 19: 403-417.

Radwin, G. E. 1978. The family Columbellidae in the western At- lantic. Part Ilb. The Pyreninae (continued). The Veliger 20: 328-344.

PHYLOGENY OF COTONOPSIS AND COSMIOCONCHA

41

Scheltema, R. S. 1978. On the relationships between dispersal of pelagic veliger lar\'ae and the evolution of marine prosobranchs gastropods. In: B. Battaglia 1 A. Beardmore, eds., Marine Organ- isms. Plenum Publishing Corporation. New York, New York.

Swofford, D. L. 2001. PAUP: Phylogenetic Analysis Using Parsimony, Version 4.0bl0. Illinois Natural History Survey, Champaign, Illinois.

Tracey, S., ). A. Todd, and D. H. Erwin. 1993. Gastropoda. In: M. J. Benton, ed.. The Fossil Record 2, Chapman and Hall, London. Vermeij, G. I. 1978. Biogeography and Adaptation. Harvard Univer- sity Press, Gambridge, Massachusetts.

Accepted: 27 March 2007

Appendix 1. Character and character state list.

1- Shell shape: (0) fusiform (elongate, spire high); (1) strombiform, spire low; (2) buccinoid (stout, spire tapering); (3) columbelloid (stout, spire high)

2- Shape of spire whorls: (0) straight sided; (1) straight going to convex; (2) straight going to concave

3- Depth of suture: (0) shallow; (1) impressed; (2) incised

4- Shoulder on spire whorls: (0) absent; (1) present, inconspicuous; (2) present, strong

5- Number of whorls in protoconch: (0) <2; (1) 2-3; (2) >3

6- Axial sculpture on early teleoconch whorls: (0) absent; (1) present, inconspicuous and subordinate; (2) present, well developed

7- Spiral sculpture on early teleoconch whorls: (0) absent; (1) present, inconspicuous and subordinate; (2) present, well developed

8- Axial sculpture on late spire whorls: (0) absent; (1) present, inconspicuous and subordinate; (2) present, well developed

9- Spiral sculpture on late spire whorls: (0) absent; (1) present, inconspicuous and subordinate; (2) present, well developed

10- Spiral sculpture on body whorl: (0) absent; (1) present, inconspicuous and subordinate; (2) present, well developed

11- Axial sculpture on body whorl: (0) absent; (1) present, inconspicuous and subordinate; (2) present, well developed

12- Shoulder on body whorl: (0) absent; (1) present

13- Cords on base of body whorl: (0) absent; (1) present, weak; (2) present, well developed

14- Concavity on central part of body whorl: (0) absent; (1) present

15- Constriction on lower part of body whorl: (0) inconspicuously constricted; (1) strongly constricted

16- Inflation of body whorl: (0) not inflated; (1) inflated

17- Type of sculpture on early vs. late spire whorls: (0) same; (1) different

18- Shape of aperture: (0) broad; (1) narrow; (2) slit-like

19- Thickness of outer lip: (0) not thickened; (1) slightly thickened; (2) conspicuous thickness

20- Teeth on inner surface of outer lip: (0) absent; (1) present, small and inconspicuous; (2) present, strongly developed

21- Number of teeth on inner surface of outer lip: (0) none; (1) few (1-5); (2) numerous (>5)

22- Posterior canal: (0) absent; (1) present, inconspicuous; (2) present, well developed

23- Apertural callus: (0) absent; (1) present, as a slight thickness; (2) present, continuous, well developed

24- Columellar denticles: (0) absent; (1) present;

25- Parietal callus: (0) absent; (1) present, slightly thickened; (2) present, well developed

26- Parietal denticles: (0) absent; (1) present

27- Parietal ridge: (0) absent; (1) present, small and inconspicuous; (2) present, well developed

28- Sinus on outer lip: (0) absent; (1) present

29- Flaring of outer lip: (0) absent; (1) present

30- Length of anterior canal: (0) short; (1) intermediate; (2) long

31- Width of anterior canal: 90) wide; (1) narrow

32- Extension of adapical part of outer lip (aperture edge at suture): (0) outer lip not extended; (1) outer lip somewhat extended after suture

33- Shape of anterior canal: (0) slightly curved; (1) strongly curved; (2) straight

34- Notch of anterior canal (at the end): (0) shallow; (1) deep depression

35- Thickening behind outer lip: (0) absent; (1) present, slight thickness; (2) present, well developed

36- Dorsal hump: (0) absent; (1) present, slight thickness; (2) present, well developed

37- Edge of outer lip: (0) sharp; (1) rounded

38- Hump on left side of outer lip: (0) absent; (1) present, slight thickness; (2) present, well developed

39- Repeated thickenings behind outer lip: (0) absent; (1) present

40- Plicae on columella: (0) absent; (1) present

41- Relation aperture height/total height: (0) aperture <‘/2 total shell height; (1) aperture much smaller than */: total shell height; (2) aperture bigger than Vi but smaller than Va total shell height

42- Gollar-like band below spire suture: (0) absent; (1) present

42

AMERICAN MALACOLOGICAL BULLETIN 23 • 1/2 • 2007

Appendix 2. Character matrix used for analyses. Species

Characters

Nassariiis luteostoma

2111221222

2220120012

2220202010

0022101001

00

Nassarius aiitilinnim

2111222222

2120120002

2220202010

0022101001

00

Canthams ringens

2102021222

2220120001

2211112010

0012200000

00

Lati nis co iicen trie us

0202021222

2120110001

2010100012

2020000002

10

Cotoiwpsis (Cotoiwpsis) argentea

0100020200

1010100111

1011101002

0011100000

00

Cotonopsis (Cotoiwpsis) crassiparva

0120120211

0020110022

1120202011

0011100000

00

Cotoiwpsis (Cotoiwpsis) deroye

0100120011

0010100121

1120102002

1011100000

00

Cotoiwpsis (Cotoiwpsis) edentula

0120120000

0020111010

0110101012

0111100000

00

Cotoiwpsis (Cotoiwpsis) jaliscaiia

0121020000

0110111121

2120102010

0000100000

00

Cotonopsis (Cotonopsis) mendozana

0100120000

0010111021

1020201010

0011100000

00

Cotoiwpsis (Cotonopsis) panacostaricensis

0120120211

1020110011

2120102012

0011100000

00

Cotonopsis (Cotoiwpsis) skoginiidae

0100?22021

0010001012

2020201002

1011100000

00

Cotoiwpsis (Cotonopsis) suteri

0111210000

0120111021

1120121002

0010100000

00

Cotonopsis (Cotoiwpsis) esmeraldensis

0100020111

0120111122

1020101002

0111101000

10

Cotoiwpsis (Cotonopsis) aff. deroye

olomoloo

1100111011

2110102000

0000101000

10

Cotoiwpsis (Cotonopsis) atf. suteri

0110120111

0120101011

1220001001

0010100000

00

Cotonopsis (Cotonopsis) phuketensis

0120110010

0020101111

2020100001

0001100000

00

Cotoiwpsis (Cotonopsis) lindae

0100110100

0020111012

2211102001

0101100000

00

Cotonopsis (Cotonopsis) inonfilsi

0121212011

0120100022

2111101000

1121101000

00

Cotonopsis (Turrina) turrita

0000100010

0010001120

0210102002

0001200100

00

Cotonopsis (Turrina) hiriindo

0000100010

0010101020

0210001012

1011200000

00

Cotonopsis (Turrina) radwini

0000100010

0010111020

1210102010

0011200000

00

Cosmioconcha palineri

0220120010

0120111032

2221200011

0011100000

01

Cosmioconcha nwdesta

3220101011

0010001022

2220201010

0001200000

01

Cosmioconcha rehderi

3200220210

2020011031

1220201011

0121200000

11

Cosmioconcha nitens

0210100010

0010001021

2110100000

0021100000

11

Cosmioconcha calliglypta

0200110110

0010011021

2110100000

0021101000

11

Cosmioconcha parvula

0200100011

0010101022

2110100002

0001101000

01

Cosmioconcha pergracilis

0200200010

0010101021

1110100002

0001101000

01

Ainer. Maine. Bull. 23: 43-78

Family Pseudolividae (Caenogastropoda, Muricoidea): A polyphyletic taxon’^

Luiz Ricardo L. Simone

Museu de Zoologia da Universidade de Sao Paulo, Cx. Postal 42494, 04299-970 Sao Paulo, SP, Brazil, lrsimone@usp.br

Abstract; A detailed morphological study was performed on the following taxa normally considered to belong to the family Pseudolividae: (1) Zemira australis (Sowerby, 1833) from Australia; (2) Fulmcntuin ancilla (Hanley, 1859) from South Africa; and (3) Melnpiinii lineatum (Lamarck, 1822) from South Africa. Two additional species of pseudolivids, Bentlwbia atafona Simone, 2003 and B. complexirhyiia Simone, 2003, from Brazil and New Zealand respectively, are considered. Two other muricoideans are included in this study: ( 1 ) Nassodonta dorri (Watteblet, 1886) [Nassariidae] from Vietnam (morphological study also included) and (2) Sirntiis senegaleusis (Gmelin, 1791) (Muricidae) from Brazil (published elsewhere). Both species are outgroups, but operationally included as part of the ingroup in order to test the monophyly of the Pseudolividae. In particular, N. dorri has a shell very similar to a pseudolivid. A complete taxonomical and morphological treatment of each species is included, as a scenario of a formal phylogenetic analysis. Additional outgroups considered include a pool of Tonnoidea (the root) and Conoidea. The cladogram is: (Tonnoidea (Conoidea {{Beutlwhia atafona-B. complexirhyna) {Nassodonta dorri (Zemira australis (Fulmentiim ancilla {Siratiis senegalensis-Melapium lineatum))))))). Analyses of each important character and of the cladogram were performed. Some of the conclusions include that the family Pseudolividae, as presently understood, is polyphyletic, as it

would include a nassariid {N. dorri) and a muricid (S. senegaleusis). Key words: Neogastropoda, polyphyly, morphology, phylogeny

The taxon Pseudolividae Fisher, 1884, had been previ- ously used by several researchers {e.g., Cossmann 1901, Goli- kov and Starobogatov 1975, Squires 1989), but it was better defined as a family by Kantor (1991), based on anatomical features of basal neogastropods. The family reunites genera previously considered as belonging to several other families, including, e.g., Cancellariidae {Bentlwbia Dali, 1889), Buc- cinidae {Buccinorbis Conrad, 1865), and Olividae (Melapiwn Adams and Adams, 1853; PseudoUva Swainson, 1840; Sylvn- nocochlis Melvill, 1903; Zemira Adams and Adams, 1853). This taxonomy was followed by some researchers {e.g., Ver- meij and DeVries 1997, Bouchet and Vermeij 1998, Pacaud and Schnetler 1999, Nielsen and Frassinetti 2003). More- over, Vermeij (1997, 1998) revised the family Pseudo- lividae, including fossil species, establishing its origin in the late Cretaceous. A more complete history of the concept of the family can also be found in that paper. However, some authors still considered the family as a subtaxon of Olividae {e.g., Hayes 1994, Smith 1998) (Pseudolivinae). Although our knowledge of pseudolivid species is relatively rich, particularly with regard to anatomy (e.g.. Ponder and Darragh 1975, Kantor 1991, Simone 2003), the defini- tion of the family remains unclear, and no phylogenetic analysis has yet been performed, other than that of Kantor (1991).

The main difficulty in studying pseudolivids is finding preserved animals. Pseudolivids are normally rare and found in deep waters, which precludes obtaining a large set of samples for an extensive anatomical study. Although the pseudolivids are more abundant as fossils, with about a hun- dred species (Vermeij 1998), they are relatively poor in di- versity in the Recent fauna, with about 10-15 living species. As about a third of the species are available for study, be- longing to the different branches of the family, a study of them appears to be worthwhile, at least in terms of testing the monophyly of the group and identifying anatomical characters that would better define it.

This paper is part of a larger project on the phylogenetic definition of the Caenogastropoda based on detailed morphology, this time focusing the Pseudolividae. One of the genera, Bentlwbia Dali, 1889, was published else- where (Simone 2003), and the species of remaining genera are included herein.

Nassodonta dorri (Watteblet, 1886), from Vietnam, one of the few freshwater neogastropods known, belonging to the family Nassariidae, has a shell similar in morphology to those of pseudolivids (Kantor and Kilburn 2001). This taxon is also included in this study to test the monophyly of the Pseudolividae, as the shell characters certainly can converge.

* From the symposium “Relationships of the Neogastropoda” presented at the meeting of the American Malacological Society, held 31 luly-4 August 2004 at Sanibel Island, Florida.

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AMERICAN MALACOLOGICAL BULLETIN 23 • 1-2 • 2007

MATERIALS AND METHODS

Most specimens used in this study belong to institu- tional collections. The specimens were dissected by standard techniques, under a microscope, with the specimens im- mersed in water. Some organs such as the oviduct and foregut were processed by standard histological tech- niques to obtain serial sections 5 pm in thickness and stained using Mallory trichrome. Hard structures, such as shells, radulae, and jaws were also examined using the SEM in the “Laboratorio de Microscopia Eletronica do Museu de Zoo- logia da Universidade de Sao Paulo”. The descriptive part of this paper provides a complete description of the first species; the remaining species are described under a com- parative aspect, with most of similar features omitted. This measure is adopted for decreasing the length of this paper and for optimizing the data. The same approach is adopted in the figures. A detailed list of examined species follows each species description.

The section on comparative morphology is organized as a phylogenetic analysis. The account on each character be- gins with abbreviated descriptive sentence followed by plesiomorphic and derived conditions(s); also included are Cl and RI (consistency and retention indices, respectively) and values for the character under the most parsimonious hypothesis. Following the apomorphic state(s), a list of ter- minal taxa with the apomorphic condition is presented. Hundreds of characters were selected, based on the exam- ined samples. Those that emerged as autapomorphic, highly variable, or overlapping, were selected but not included in the cladistic analysis. The remaining characters were orga- nized into states, coded, polarized comparing with out- groups, and a cladistic analysis was performed.

Other previously studied Caenogastropoda were selected as outgroups. They are mainly the following; Ceri- thioidea (Simone 2001), Stromboidea (Simone 2005), Cypraeoidea (Simone 2004a),

Calyptraeoidea (Simone 2002), and architaenioglossans (Si- mone 2004b). In the discus- sion, some specific outgroup taxa are mentioned, based upon observed or published data. However, in the matrix of characters (Table 1 ) only 4 taxa are shown, the ground plan of the Conoidea and Tonnoidea.

The ground plan of these su- perfamilies are chosen in the sense of being representative; however, the final result is the same if the ground plan were

substituted by any one of the 64 (terminal) species present in those papers. Two additional species are included, Siratiis senegalensis (Gmelin, 1790) [Muricidae], based on unpub- lished observations from another, ongoing study, and the nassariid Nassodonta dorri, mentioned above. Each charac- ter, state, and polarization is justified in the discussion sec- tion which includes a concise explanation when warranted.

The discussion of each character is also based on the analysis of the resulting tree (Figs. 1-2) although the matrix of characters (Table 1) and the subsequent tree (Figs. 1-2) are shown only in the following section.

The synapomorphies of the ingroup (superfamily auta- pomorphies) are preserved in the present paper, because they are the main concern as referred to in the introduction. The ingroup autapomorphies are the basis for a better es- tablishment of a still imprecise taxon. They confirm the internal position of some possible “outgroups” such as tonnoideans and conoideans. They will be useful in the on- going phylogenetic study of the entire order Caenogas- tropoda as the ground plan of the superfamily. Additionally, they are in agreement with some phylogenetic approaches used in studies of other groups [e.g., Yeates 1992, Pinna 1996).

Some multistate characters are here analyzed under an additive (ordered) approach. In each case, the additive concept is justified in the discussion and is always based on the ontogeny, or because each state represents a clear modification of the preceding one. Additionally, each addi- tive multistate character was also analyzed as non-additive, and any fortuitous change in the result and/or indices are also reported. The cladistic analysis was performed with the aid of the computer program “Tree Gardner 2.2” (Ramos 1997), which basically works as an interface of the Hennig86 (Farris 1988). The used algorithm was “ie”. The computer program PAUP was also used, mainly to determine the ro- bustness of the nodes. Both programs presented the same result.

Table 1. Matrix of characters of ingroup and two outgroups (bottom).

Taxon/character

1

2

3

12345

67890

12345

67890

12345

67890

1234

Bentiwbia atafona

10101

00121

10001

00011

21010

20101

1111

Benthobia complexirhyna

10101

00121

10001

00011

21010

20101

1111

Zemira australis

11010

01120

01111

10011

01121

11101

0111

Melapium Uneatum

00100

13010

01111

mil

11121

21010

0011

Fiihnentiim ancilla

11110

00010

01101

11011

21111

11110

1111

Nassodonta dorri

10110

01011

00111

10111

?1110

2??01

0?11

Siratus senegalensis

00010

12000

01111

11111

mil

OHIO

0211

Conoidea

00100

00000

00001

00001

00010

00001

0011

Tonnoidea

00000

00000

00000

00000

00000

00000

0000

PSEUDOLIVIDAE: A POLYPHYLETIC TAXON

45

Figure 1. Single most parsimonious tree based on the data matrix in Table 1, with three outgroups operationally analyzed as part of the ingroup (Conoidea, Siratiis, and Nassodonta). Length: 62; Cl = 66; R1 = 69. Each symbol indicates a synapomorphy supporting each node (only the homoplastic autapomorphies are shown) as follows: full sc]uare = non-homoplastic synapomorphy; circle = convergence; empty square = reversion.

Figure 2. Single most parsimonious tree (same of Fig. 1, excluding both more basal out- groups) (Length; 62; Cl = 66; R1 = 69), with the nodes numbered. The gray branches represent the non-pseudolivid taxa, left branch a member of the Nassariidae, right branch a member of the Muricidae. The black branches represent the taxa mostly considered to be Pseudolividae, showing the polyphyletic nature of the taxon.

SYSTEMATICS

Genus Zemim Adams and Adams, 1853

(Type species Eburna niistmlis, by monotypy)

Zemim australis (Sowerby, 1833) (Figs. 3A-F, 4A-B, 5A-7H)

Synonymy; see Ponder and Dar- ragh 1975: 101.

Complement:

Zemira australis: Ponder and Dar- ragh 1975: 89-97, 101-104 (text figs. 1, 2, pi. 7 fig. 1-2. pi. 8 figs. 12-24); Smith 1998: 835-836 (fig. 15.165-C).

Description

Shell (Figs. 3A-C, 3F). Fusiform, pale brown, opaque. Protoconch, spherical, smooth, opaque, of about one whorl; boundary between proto- conch and teleoconch unclear. Spire pointed, about half of length of body whorl. Suture well-marked by a subsu- tural, concave, wide groove, from pro- toconch up to outer lip; surface of groove smooth, external edge elevated, forming a low carina. Remaining re- gions sculptured by uniform, spiral, narrow furrows, about nine in pen- ultimate whorl, about 10 in body whorl; one of these furrows, located between middle and anterior thirds of body whorl, deeper and wider (Figs. 3B-C). No umbilicus except a narrow furrow in inferior third around inner lip (Fig. 3A). Peristome oval, white, glossy (Figs. 3A, 3F). Canal short and narrow, left edge truncate, right edge wanting, as continuation of outer lip. Outer lip simple, cutting edge, rounded; very short tooth between middle and inferior thirds correspon- dent to deeper spiral furrow of body whorl (Figs. 3C, 3F); wide notch in su- perior region, at some distance from suture correspondent to sub-sutural Carina. Inner lip simple, callus narrow, slightly more transparent than inner region of peristome.

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AMERICAN MALACOLOGICAL BULLETIN 23 • 1-2 • 2007

Figure 3. Shells and opercula. A-F, Zemira australis, AMS 333288. A-C, shell (specimen #1 ), female, apertural, dorsal and lateral views, length = 18.6 mm; D-E, operculum, outer and inner views, arrow indicates separation of two scar regions, scale bar = 2 mm; F, detail of aperture closed by operculum (specimen #4), showing labral tooth and opercular sculpture. G-I, Nassodonta dorri, MZSP 53533; G-H, shell, apertural and dorsal views, length = 13.6 mm; I-|, operculum, outer and inner views, length = 5 mm.

Head-foot (Figs. 5A, 5C, 5F). Head weakly protruded, bilobed, with single region pigmented by dark brown, the rest pale cream. Tentacles located close to each other and close to median line; tentacles’ base very wide, flat, flap-like, outer edge rounded; dis- tal half of tentacles marked by abrupt narrowing of the base, narrow, taper- ing gradually; tentacles’ tip rounded. Foot broad, of about half whorl when retracted. Sole oval, edges thick and rounded. Anterior furrow of pedal glands deep, straight, thick superior and inferior edges, not reaching lat- eral-anterior end. Lateral region of the sole of the foot dearly extending be- yond remaining dorsal regions of foot, division marked by a shallow longitu- dinal furrow lying somewhat in middle region between sole edge and dorsal region of foot (Fig. 5A). Opercular pad elliptical, almost as wide as the dorsal surface of the foot; possessing clear, median, oblique difference in levels (Fig. 5C); posterior half of this division forming small area with different, iri- descent color. Columellar muscle thick, of about one half whorl. Male with large penis in posterior-right region behind the right tentacle described below.

Operculum (Figs. 3D-F). Elliptical, horny, pale to reddish brown. Nucleus sub-terminal, located closer to interior- inner edge. Outer surface with normal concentric growth lines, and series of radial lines produced by minute, aligned scales located on the growth lines, from nucleus to edges (Fig. 3F). Low carina running at some distance from inferior and inner edges, from nucleus up to middle level of inner edge. Inner sur- face glossy. Scar elliptical, occupying about 2/3 of inner area, somewhat dis- located closer to inner edge. Scar having two different levels of about the same area, one superior and another inferior; both separated by a wide chevron, marking a low step (Fig. 3E, arrow); a small notch in region where the chev- ron touches outer scar edge.

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47

Mantle organs (Figs. 5B, 5D). Mantle edge simple, thick. Siphon small, not extending beyond mantle edge. Osphradinm about 1/3 the width of the pallial cavity and V4 of its length. Osphradinm filaments tall, central re- gion scalloped by 5 folds in the left and 6 folds in the right filaments (Fig. 5B: os). Osphradinm filaments widely at- tached along mantle roof. Osphradinm anterior end curved to the left, with left filaments clearly smaller than right filaments; remaining osphradial re- gions with somewhat symmetrical fila- ments (left filaments slightly smaller). Very narrow area between osphradinm and gill. Ctenidial vein narrow, dislo- cated weakly beyond left gill edge, to- wards right edge of osphradinm. Gill slightly longer than the osphradinm and of about the same width; its ante- rior end broadly pointed, located closer to the mantle edge, far from an- terior end of osphradinm; posterior gill end located slightly posterior to that of osphradinm. Afferent gill vessel very narrow, lying at a short distance from the right edge of the gill. Between the gill and the right edge of pallial cavity there is an area ec]iiivalent in width to that of the gill. The hypo- branchial gland is thin, greenish beige, covering most of the area between the gill and the rectum, including the left and ventral surfaces of the rectum; the anterior region of the hypobranchial gland tapers gradually. Rectum nar- row, running along the right edge of the pallial cavity (Figs. 5B, 5D-E). Anus simple, sessile, locateci between middle and anterior thirds of pallial cavity. Pallial gonodncts located be- tween rectum and pallial fioor, de- scribed below.

Visceral mass (Fig. 5D). Anterior whorl mostly occupied by stomach, kidney, and pericardium (Figs. 5E, 6E). Digestive gland greenish brown, located along inferior region of each visceral whorl, covering middle diges- tive tubes and also two whorls poste-

Figure 4. Scanning electron micrographs of radulae. A-B, Zcmira iwstralis, scale bars = 30 pm. C-D, Fnimeiitum ancilla, scale bars = 50 pm. E-F, Melapiiiiu liiicalnm, scale bars = 50 pm. G-I, Nassodonta dorri, scale bars = 30 pm.

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*1

Figure 5. Zeniira australis anatomy. A, head-foot, male, frontal view. B, pallial cavity roof, transverse section at middle level of osphradium. C, foot, detail of opercular pad, dorsal view, operculum removed. D, pallial cavity, ventral- inner view, and visceral mass, male. E, region of kidney, ventral view, ventral wall of kidney and pericardium removed, anterior membrane partially deflected to right. F, head and haemocoel, ventral view, foot removed. Scale bars = 1 mm. Abbreviations listed in section with figure captions.

PSEUDOLIVIDAE: A POLYPHYLETIC TAXON

49

rior to stomach. Gonad pale beige, lying along superior and columellar surfaces of visceral whorls posterior to stomach.

Circulatory and excretory systems (Fig. 5E). Pericar- dium located just posterior to gill, along the left anterior region of the visceral mass (Fig. 5D). Auricle small, trian- gular, attached to anterior surface of pericardium, with the ctenidial vein entering from the left and the connection to kidney at its right end. Auricle connected to anterior surface of ventricle. Ventricle very large, filling most of pericardium volume. Aortas located along posterior region of the ven- tricle; anterior aorta about 4 times larger than posterior aorta, and located ventral to it. Kidney occupies about 1/3 of pallial cavity volume, located along middle and right regions of the anterior end of the visceral mass. Nephridial gland triangular in section, broader anteriorly, gradually narrow- ing posteriorly; lying along the dorsal region of the reno- pericardial wall. Renal lobe occupying most of the kidney’s interior volume, presenting two flaps of similar thickness, fused along right region; ventral flap shorter (about half of dorsal flap), intestine running through it; dorsal flap occu- pying most of renal dorsal surface. Afferent renal vessel large, running from the haemocoel, covering right side of nephropore, with some branches inserted in inner surface of dorsal flap of renal lobe.

Digestive system (Figs. 5F-8A). Proboscis relatively short (about 1/3 of haemocoel length) (Figs. 5F, 7A: pb). Mouth transverse along proboscis tip. Buccal cavity with pair of broad and tall lateral folds, each one dividing within a short distance, one branch running to the odontophore tube, the other to the esophagus (Fig. 5F). Ventral surface between buccal folds with a clear, low, flat, chitinous plat- form (Fig. 7F: ol). Odontophore oval, about half the length of the proboscis (Figs. 7A, 7E). Odontophore tube connect- ing it with buccal cavity. Odontophore muscles (Figs. 6A-D, 7E-F): ml, several small muscle fibers connecting buccal mass to adjacent inner surface of proboscis; mj, pair of peri- buccal muscles and protractor of odontophore, origin thin within dorsal wall of oral cavity, running along odontophore tube becoming thicker, inserting into outer surface of carti- lages, externally to m6 and medially to m4, in two branches, one anterior and another posterior, posterior branch about twice the size of and longer than the anterior branch; m2, pair of retractor muscles of odontophore, originating in ven- tral surface of haemocoel, in region just posterior to pro- boscis (when retracted), running dorsally, with median fi- bers running through nerve ring, inserting into posterior surface of odontophore, part into m5 and part into m4 regions close to median line; m2a, auxiliary of m2, being single and running between both m2, attached to ventral surface of anterior aorta; its fibers apparently originated ven- tral to nerve ring, not passing through it (Figs. 6A-D); m3.

pair of thin dorsal protractor muscles of odontophore, origi- nating in anterior-dorsal end of odontophore tube, at its juncture with the esophagus, running posteriorly, covering dorsal surface of odontophore tube, inserting into odonto- phore middle-dorsal surface (Figs. 7E-F); m4, strong pair of dorsal tensor radular muscles, originating in odontophore cartilages along a line surrounding their ventral surface, run- ning towards dorsal surrounding lateral surface of cartilages, inserting laterally along radular sac into the region near the buccal cavity; m5, pair of secondaiy dorsal tensor muscles of radula, originating in posterior and medial regions of the cartilages, running dorsal and medial as continuation from ni4, inserting into radular sac near the buccal cavity along- side and medial to m4 insertion; m6, thin horizontal muscle, uniting both odontophore cartilages, with about 3/4 of car- tilage length, inserting along a line into the ventral and ex- ternal surfaces of the cartilages, at a short distance from their inner-ventral edge, starting at the anterior end of the carti- lages and ending just before their posterior c]uarter (Figs. 6B-C); ml la, pair of ventral tensor muscles of radula, thin, somewhat broad, originating partly in the posterior-ventral end of cartilages and partly in the m2 insertion, running anteriorly covering m6, inserting into ventral edge of radula and subradular cartilage, and some inner portion preceding this (Fig. 6D); ml4, pair of ventral protractor muscles of odontophore, originating along ventral surface of oral fube and tube of odontophore, running posteriorly at short dis- tance from median line, covering central surface of odonto- phore, inserting into posterior-ventral surface of odonto- phore, close to m2 insertion (Fig. 7E). Other non-muscular odontophore structures: sc, subradular cartilage, expanding in exposed region of radula into buccal cavity, covering neighboring surface of radula (Figs. 6A, 6D, 7F); oc, odon- tophore cartilages, somewhat elliptical, flat, with medial- ventral edge slightly straighter than outer edge, posterior region clearly narrow (Figs. 6B, 6C); br, subradular mem- brane, covering inner surface of subradular cartilage and radula, m4, ni5, and ml la inserfions. Radula (Figs. 4A-B): rachidian toofh with short transverse base, spanning about 1/3 of radular ribbon, 3 long, tall (about 2/3 of base length), sharp pointed cusps somewhat ecyiidistant from each other, central cusp symmetrical, outer cusps weakly turned out- wardly; between rachidian and lateral teeth a distance equivalent to 1/3 of rachidian width; lateral tooth hook-like, base broad (equivalent to 2/3 of rachidian base width), gradually narrowing up to sharply pointed tip, height about 1.5 that of rachidian; straight to weakly curved inwardly. Salivary glands clustering along anterior region of valve of Leiblein and ventral ganglia of nerve ring, attaching to lateral surface of the anterior esophagus just anterior to the valve of Leiblein (Figs. 7A-B); their ducts very narrow, totally at-

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AMERICAN MALACOLOGICAL BULLETIN 23 • 1-2 • 2007

Figure 6. Zernim australis anatomy. A, odontophore, dorsal view, anterior odontophore tube removed. B, same, outer layer of muscles and radular apparatus removed and only partially shown (rs). C, odontophore, ventral view, only its right side shown, posterior muscles deflected. D, same, outer view, outer layer of muscles and membrane partially removed. E, midgut, ventral view as in situ, some adjacent structures also shown. Scale bars = 1 mm. Abbreviations listed in section with figure captions.

tached to anterior esophagus wall and to lateral wall of oral tube; opening very small (Fig. 7F: sa), into anterior region of lateral folds of buccal cavity, somewhat ventrally, just within the anterior end of a narrow furrow surrounding the ventral edge of the odontophore tube folds. Anterior esophagus with somewhat thick walls, length ec]uivalent to that of odontophore, inner surface with lateral, longitudinal, low, and flat folds that become narrower posteriorly. Between these folds are secondary, low, narrow folds (Fig. 7F). Valve

of Leiblein with about 1/4 of odontophore volume, anterior region with a transverse, white band into which long cilia insert, middle and posterior regions pale beige, correspond- ing to a tall inner gland occupying most of inner surface; oblique furrow (by pass) present, separating all valve regions (Figs. 7A-B); inner surface smooth, not glandular, bordered by pair of low and veiy narrow folds that diverge in its anterior region, and continuous with middle esophagus folds posteriorly. Middle esophagus about half as long as

PSEUDOLIVIDAE: A POLYPHYLETIC TAXON

51

Figure 7. Zemira australis anatomy. A, foregut, extended, ventral view. B, region of the valve of Leiblein, its oblique furrow (vf) seen by transparency. C, gland of Leiblein partially uncoiled, some adjacent structures also shown. D, transition between middle and posterior esophagus and duct of gland of Leiblein, opened longitudinally to expose inner tokis. E, buccal mass, left view. F, buccal mass, anterior region opened longitudinally and deflected to expose inner surfaces. Scale bars = 1 mm. Abbreviations listed in section with figure captions.

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AMERICAN MALACOLOGICAL BULLETIN 23 • 1-2 • 2007

anterior esophagus (Fig. 7A: em), walls thin; inner surface with longitudinal, low, narrow folds, a pair of close folds larger (Eig. 7D), running towards duct of gland of Leiblein. Gland of Leiblein triangular in situ (Figs. 5F, 7A), long and somewhat flat if uncoiled (Fig. 7C), becoming about as long as posterior esophagus; anterior aorta crossing between middle and posterior thirds of this gland. Duct of gland of Leiblein long and narrow (about as long as middle esopha- gus, and about half of its diameter) (Figs. 7A, 7D); having two origins, one sub-terminal in anterior end of the gland, the other in a portion more posterior (Fig. 7C); these two ducts unite within a short distance, remaining duct having pair of tall, longitudinal, narrow folds (continuation from larger folds of middle esophagus); these folds separate a narrow, white, multi-lobed secondary gland from a smooth, narrow area; this secondary gland occupies about 2/3 of duct volume, ending abruptly before the duct’s insertion into the esophagus (Fig. 7D). Posterior esophagus (Figs. 7A, 7D; ep) about twice as long as the anterior esophagus, inner surface with narrow longitudinal folds, some low, others taller (cov- ering lower folds) diverging and coalescing randomly; these folds disappearing abruptly before stomach. Stomach spherical, blind sac, about half the width of adjacent visceral whorl. Esophagus enters stomach along its left anterior re- gion (Figs. 6E, 8A: st), intestine originates to the right of the esophageal insertion. Gastric inner surface (Fig. 8A) mostly smooth, except for a pair ot low, narrow folds that run along its left surface, from the esophageal insertion, disappearing gradually into the posterior gastric surface. Duct to digestive gland single, wide, located between esophageal insertion and intestinal origin (Fig. 6E). Intestine with tall, dorsal, smooth, long and triangular platform adjacent to the stom- ach (Fig. 8A); left edge of this platform alongside a band of longitudinal, narrow folds; right edge of this platform serv- ing as insertion of several transversal folds, each about twice as wide as the longitudinal folds, becoming gradually ob- lic]ue, surrounding ventral surface of intestine, ending at left edge of band with longitudinal folds. Inner surface of intes- tine, beyond this platform, with only longitudinal folds, very close to each other, filling inner surface totally. Intestine runs almost straight anteriorly, crossing through anterior region of digestive gland, kidney lobe, and right edge of pallial cavity (Figs. 5E, 6E). Rectum and anus described above (pallial cavity).

Genital system. Male (Figs. 5A, 8B). Visceral vas def- erens begins a half whorl before anterior end of testis. Within a short distance it becomes a very broad, intensely coiled seminal vesicle, occupying about half of adjacent vis- ceral whorl (Fig. 5D). Seminal vesicle located in the ventral surface of last whorl of the visceral mass, posterior to kidney; becomes narrow at some distance posterior from pallial cav- ity, running about 1/6 whorl. Prostate gland relatively nar-

row, running along right edge of pallial cavity ventral to rectum, visceral vas deferens inserting posteriorly (Fig. 5D: pt); walls thick-glandular; no apertures to pallial cavity; in- ner lumen surrounded by muscle fibers. Prostate spans about Vs pallial cavity length, gradually becoming narrow, crossing to pallial floor. This region in the pallial cavity floor with thick, muscular walls, slightly convolute up to penis base (Fig. 8B). Penis slightly larger than half of pallial cavity volume, stubby, dorso-ventrally flat (Figs. 5A, 8B); base broad, with a large, broad right fold covering base of right tentacle; then twisting, remaining tall, flat and thick, nar- rowing gradually up to bluntly pointed tip (Fig. 8B). Pallial vas deferens within the integument, becoming penis duct. Penis duct running approximately along penis center, very narrow, weakly coiled. Penis aperture apical, very small.

Female (Figs. 8C-F). Visceral oviduct very narrow, run- ning along middle region of columellar surface of the last whorl of the visceral mass, about W whorl preceding pallial cavity, gradually becoming thicker, inserting into pallial ovi- duct without clear separation. Posterior region of pallial ovi- duct protruding into kidney, having a narrow zigzag. Albu- men (whitish) and capsule (beige) glands adjacent, albumen gland spanning posterior 1/5 of pallial oviduct. Seminal re- ceptacle very small, located between albumen and capsule glands (Fig. 8C); flat to rounded; duct very narrow, attached along the dorsal surface of the pallial oviduct, opening into the vaginal furrow between the albumen and capsule glands. Capsule gland with flat lumen, vaginal furrow running along its left edge, with surface smooth (Fig. 8D). Female pore wide, protruded, with thick edges (Figs. 8E-F: fp). Bursa copulatrix small, short, located along left side of the distal, detached portion of the pallial oviduct (Fig. 8F: be); with thick muscular walls; its aperture turned anteriorly, occupy- ing about 2/3 of total female pore; inner surface with low, wide, longitudinal folds. Capsule gland aperture narrow, situated to the right of the bursa aperture; its walls thick muscular, protruding inside the chamber of the terminal atrium of the capsule gland. Terminal atrium, with thin walls, located between capsule gland anterior and female pore. Female pore with several wide, longitudinal folds. No cement gland in foot sole.

Central nervous system (Figs. 8G-H). Relatively well- concentrated. No distinction between pleural and cerebral ganglia. Cerebral ganglia broadly connected to each other. Pedal ganglia slightly smaller than cerebro-pleural ganglia; pedal commissure broad, but narrower than cerebral com- missure. Cerebro-pedal and pleuro-pedal connectives short, but distinguishable. Pair of buccal ganglia small, located close to posterior edge of the cerebral ganglia.

Measurements of shells (in mm). AMS C333288; ?1 = 18.6 by 12.0; 62 = 19.3 by 12.2; 63 = 16.6 by 9.8; 54 = 14.1 by 8.7.

PSEUDOLIVIDAE: A POLYPHYLETIC TAXON

53

Figure 8. Zemira australis anatomy. A, stomach, ventral view, its ventral wall and adjacent region of intestine (in) and esophagus (es) opened longitudinally, inner surface exposed. B, penis and adjacent region of its base, ventral view, penis partially deflected, its duct (pd) seen by transparency. C, pallial oviduct, dorsal view, detail of its posterior region. D, pallial oviduct, transversal section though its middle region. E, pallial oviduct, entire ventral view, including its portion in kidney chamber. F, same, detail of its anterior end, ventral wall removed, region of pore opened longitudinally and deflected to show its parts. G, central nervous system, ventral, slightly right oblique view, esophageal passage (es) shown by arrow. H, same, dorsal, slightly left oblique view. Scale bars = 1 mm. Abbreviations listed in section with figure captions.

Distribution. East coast of Australia.

Habitat. Sandy, from 3 to 146 m depth (Beechey 2005). Material examined. AUSTRALIA. New South Wales; Sydney (Shelf Benthic Survey col.), 1.6 km east of Malabar

outlet, 33°58.250'S 1 5 1° 1 7.000'E, 66 m depth, AMS C092091, 3? (Sta. 027653, SBS3, 26/jii/1973), 2.3 km east of Malabar outlet, 33°59.450'S 151°16.800'E, 66 m depth, AMS C333288, 3d, 6? (Sta. 002609E, SBS5, 25/x/1973), 2.6 km

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east of Cape Banks, 33°59.500'S I5I°I6.740'E, 66 m depth, AMS C406792, Id (Sta. 038772, SBS25, 26/1/1973).

Genus Fidmentiim Eischer, 1884

(Type species Biiccinwn sepimentum Rang, 1832, by monotypy)

Fulmentum ancilla (Hanley, 1859)

(Figs. 9A-E, 4C-D, 10A-13F)

PseiidoUva ancdla: Kantor 1991: 31-34 (figs. 12, 13); Hayes 1994: 77-78.

Sylvanocoddis ancdla: Lorenz 1989: 16-18.

Fiilmentwn ancdla: Vermei) 1998: 60.

Description

Shell (Figs. 9A-B, 9E). Large (about 60 mm), heavy, biconical. Color brown to beige. Periostracum velvet-like, rust colored, partially eroded. Spire conical, aperture span- ning 70% of shell length. Protoconch of IVi whorls, white, smooth, to wealdy reticulated in some areas, border with teleoconch unclear. Teleoconch of 5-6 whorls, each whorl with a weakly concave shoulder. Axial sculpture limited to growth lines. A wide spiral furrow runs along the anterior third of the shell, forming a low, broad labral tooth. Umbi- licus absent. Aperture elliptical, posterior end pointed. Siphonal canal wide, open, very short. Inner lip smooth, callus narrow and low except for a small, low node at some distance from posterior end. Outer lip with simple edge, thickened at the shoulder and in region of the labral spine. Thickening of the outer lip and the callus of the inner lip located just posterior to the spiral furrow on the previous whorl.

Head-foot (Figs. 9E, lOB-C, llA). Head not protruded. Color mostly pale beige, with dark brown, coalescent (form- ing wide transversal bands) spots in head and dorsal surface of foot (Fig. 9E). Tentacles relatively short, located close to each other; basal half clearly thicker than distal half. Eyes located along outer edge of tentacles at middle level, proxi- mal to their constriction; terminal portion of tentacle very short (Figs. lOB-C). Rhynchostome located ventral to and between the tentacles. Foot spans Vi whorl when retracted; sole simple; anterior edge with deep furrow that contains the pedal glands. Dorsal and ventral edges of this furrow rela- tively thick and rounded; both ends of anterior edge rounded. Columellar muscle spans Vi whorl, thick; posterior end with a wide, rounded component, and a small (about 10% of origin area) projection located along its left end. Haemocoel about 1/3 head-foot width along its anterior half, posterior half becomes very narrow and turned to the left (Fig. IOC).

Operculum (Figs. 9C-D). Elliptical, horny, brown, oc- cupying entire shell aperture. Nucleus terminal, inferior. Outer surface with concentric growth lines and narrow un-

dulations. Inner surface glossy. Scar occupying about 80% of inner surface, approx, central, slightly displaced closer to inner and superior edges. Scar divided into approx, two similar sized areas by oblique line. Inferior region broadly pointed, with wide free projection; triangular, wide furrow running along middle region of this projection, starting in the apex, finishing in the scar.

Mantle organs (Figs. lOA, lOE). Mantle edge simple, thick, transversally banded. Siphon relatively long (about half of pallial cavity length), edges simple (Figs. 9E, lOA); mantle edge surrounding its base. Pallial cavity with about Wi whorls. Osphradium about % pallial cavity length and about '/4 of its width, with curved, symmetrical filaments; anterior and posterior ends rounded; each filament relatively tall, projecting laterally; dorsal edge connected to mantle along Vi its length the rest supported by a rod; ventral edge thin, with small notch located approximately in central re- gion; right filaments covering ctenidial vein and part of gill filament bases (Fig. lOE: os). Gill about 3/4 of pallial cavity length and 1/4 of its width; anterior end pointed, posterior to that of osphradium; ctenidial vein extending a short dis- tance anterior to gill end; posterior gill end rounded, touch- ing pericardium. Gill filaments triangular and tall, curved to right; rod relatively broad; filament tip rounded, preceded by narrow region. Ctenidial vein narrow, lying along left edge of gill. Afferent gill vessel very narrow, lying at some distance from apparent right gill edge. Hypobranchial gland some- what tall, whitish, covering entire area between gill and rec- tum (about V2 pallial cavity width), anterior end at level of anus. Rectum relatively narrow, running along right edge of pallial cavity. Anus detached, situated between middle and anterior thirds of pallial cavity, with small papilla on its right side. Genital ducts lying between rectum and right edge of cavity, described below.

Visceral mass (Fig. lOA). Spans about three whorls. Reno-pericardial structures occupy anterior half whorl. Stomach located just posterior to reno-pericardial area. Di- gestive gland greenish cream, occupying all whorls, sur- rounding stomach. Gonad same color as digestive gland, occupying superior and columellar surface of each visceral whorl, terminating a short distance posterior to kidney.

Circulatory and excretory systems (Fig. lOD). Pericar- dium about '/4 of kidney volume. Heart similar to that of Zeniira. Kidney somewhat trapezoid, spanning half whorl. Nephridial gland flat, thicker anteriorly, lying along entire dorsal half of reno-pericardial membrane. Kidney lobe similar to that of Zemira. Afferent renal vessel large, with branches running between folds of both kidney lobe flaps, covering right side of nephropore, connected to membrane between kidney and pallial cavities, producing small urinary cavity.

Digestive system (Figs. 1 1A-12E). Rhynchostome forms a small opening located ventral to and between both cephalic

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55

Figure 9. Shells, opercula, and living specimens. A-E, Fiihuentuiu aiidihr, A-B, shell, NMNH E5279, female, apertural and dorsal views, length = 64.5 mm. C-D, Operculum, outer and inner views. E, Live, crawling specimen from Jeffreys Bay, South Africa, photo courtesy of Brian Hayes. F-K, Melapium Uneatum. F-G, shell, NMNH V9979, male, apertural and dorsal views, 2 egg capsules attached in anterior region of inner lip (total length = 26,2 mm). H, detail of egg capsules attached to shell, NMNH 59733, portions ot both capsules removed, showing young specimens inside. E), dorsal and apertural views ot a young specimen extracted from an egg capsule shown in the preceding figure, scale bar = 2 mm. K, live, crawling specimen from off Algoa Bay, South Africa, photo courtesy of Brian Hayes.

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Figure 10. Fulmentum nncilla anatomy. A, pallial cavity and visceral mass, male, ventral-inner view. B, head-foot, male, frontal view. C, head and haemocoel, ventral view, foot and columellar muscle removed. D, kidney and pericardium, ventral view, ventral wall removed, renal membrane with pallial cavity (km) deflected anteriorly (right in fig.). E, Transverse section through pallial cavity roof, at mid-length of the osphradium. Scale bars = 2 mm. Abbreviations listed in section with figure captions.

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ad sa

me

Figure 11. Fuhnentum ancilla anatomy. A, foregut partially uncoiled, ventral view, adjacent region of head also shown. B, buccal mass, left view, esophagus opened longitudinally. C, odontophore, dorsal view, some muscles deflected. D, same, dorsal view, superficial layers of muscles and membrane reflected, aorta shown as in situ. Scale bars = 2 mm. Abbreviations listed in section with figure captions.

tentacles (Fig. IOC). Proboscis short (less than 1/3 of hae- mocoel length) and broad (about haemocoel width) (Figs. IOC, llA); walls thick, muscular. Proboscis retractor muscles forming a main paair, one in each side, left retractor muscle slightly more dorsal than right, originating at middle level of haemocoel latero-ventral region, running towards

middle level of p^roboscis, inserting along its distal half (Fig. IOC: rm); several accessory proboscis retractor muscles along dorsal surface, thinner than main retractor muscles, originating in a virtual line connecting both main retractor muscles, surrounding dorsal haemocoelic wall. Mouth a transversal slit on proboscis tip. Oral tube with thick walls.

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about half the length of the odontophore. Oral cavity wide, with a pair of low, broad, dorsal folds, each occupying about 1/3 of the dorsal surface and 1/3 of area between them; each dorsal fold with rounded anterior end, at short distance from mouth (Fig. 1 IB); remaining fold characters similar to those of Zemira. Ventral chitinous platform present (Fig. IIB, ad). Odontophore about same length as proboscis (when extended), protruding beyond proboscis into haemo- coel when retracted (Figs. IOC, llA). Odontophore organi- zation and musculature similar to that of Zemira, with fol- lowing distinctions. Odontophore muscles (Figs. 11B-12A): m2 a single pair inserted into posterior region of m4, by side of radular sac; m2a absent; m5 pair wider and thicker; m6 with anterior end at some distance from that of odonto- phore cartilages (Fig. 10); ml la pair broader (Fig. IID). Odontophore cartilages elliptical, with posterior region simi- lar to anterior region (Fig. 12A). Radula (Figs. 4C-D): ra- chidian tooth with broad base (about 60% of radular rib- bon), close to adjacent teeth, with 3 tall (about 1/2 base width), triangular, sharply pointed cusps, somewhat equi- distant and separated from each other; distance between rachidian and lateral teeth area equivalent to half of rachid- ian base width; lateral tooth with base 60% of rachidian width, 2 tall (height equivalent to base width), triangular, sharply pointed cusps, one on each side of the tooth, inner cusp slightly smaller than outer cusp. Anterior esophagus shorter than odontophore, wall thick muscular, with some muscles connecting its latero-ventral region to the adjacent region of the haemocoel floor, sometimes passing through the salivary glands; inner surface with a pair of lateral folds, as continuations from buccal cavity folds, gradually narrow- ing (Figs. IIA-B: ae). Salivary glands paired, each an ellip- tical, separated mass located along each side of valve of Leiblein and close to the nerve ring (Figs. 1 lA-B, 12B: sg). Salivary ducts very narrow, originating in the anterior end of the gland, penetrating the lateral walls of the anterior esophagus within a very short distance; running within this wall up to salivary aperture. Salivary aperture very small, located in lateral edge of buccal cavity dorsal folds, at mid- level, just within the posterior end of a narrow and shallow furrow running anteriorly, surrounding antero-lateral edge of dorsal folds (Fig. IIB; sa). Accessory salivary gland single, elliptical, internally hollow, situated within the hae- mocoel near the middle region of odontophore’s ventral surface. Anterior region of gland gradually narrowing with- out clear division from its duct. Duct long and very narrow, equal to the length of odontophore, lying along ventral sur- face of the odontophore, odontophore tube and oral tube (Figs. IIA-B: ae); opening of duct very small, in median region of the ventral surface of the oral tube, just posterior to the mouth (Fig. IIB: ad). Valve of Leiblein with about half size of odontophore, inner organization similar to that

of Zemira, with narrow folds of the oblique furrow disap- pearing gradually at anterior and posterior ends (Figs. IIA- B, 12B: vl). Middle esophagus, narrow, roughly equal in length to the anterior esophagus, walls thin, inner surface smooth (Figs. IIA-B: em). Gland of Leiblein brown, long, triangular, anterior region wide, narrowing gradually to- wards posterior, posterior tip narrow and rounded; anterior aorta crossing through this gland between middle and ante- rior thirds (Figs. IOC, 1 lA: gl). The portion of the duct of the Gland of Leiblein that is free from the gland is relatively long (about half the length of the middle esophagus) (Figs. IIA, 12C: Id); inner surface with a longitudinal, white gland in its dorsal side, and a smooth, thin region on the ventral side; lacking transverse septa within (Fig. 12C). Posterior esopha- gus narrow, about three times as long as anterior esophagus, wall thin, inner surface smooth or with narrow longitudinal folds, close to each other (Figs. 11 A; ep; 12D-E: es). Stomach trapezoidal, weakly dorso-ventrally flattened, occupies about half of visceral whorl volume, is situated about 1/4 whorl posterior to kidney (Figs. 12D-E); esophagus joins the stom- ach at left-anterior end, the intestinal joins the stomach to the right of the esophagus, and is slightly wider. Duct to digestive gland single, located a short distance posterior to esophageal insertion and intestinal origin; duct with very wide and flat base, with branches running from opposite sides after short distance. Gastric walls thick muscular. Gas- tric inner surface with transversal, low, broad, somewhat irregular folds (Fig. 12E). Intestine almost straight, weakly sigmoid, running anteriorly (Figs. lOD, 12D-E); inner sur- face full of low, narrow, closely spaced longitudinal folds; A larger pair of adjacent folds run along the left side of the stomach, gradually disappearing into rectum. Intestine passes initially through digestive gland, then through the ventral flap of the kidney lobe. Rectum and anus as de- scribed above (pallial organs).

Genital system. Male (Figs. lOA-B, 13A-B). Testis as described above (visceral mass). Seminal vesicle very large, occupying the columellar surface of almost the entire last whorl of the visceral mass, forming a relatively narrow, ex- tremely convoluted tube (Fig. lOA: sv). Seminal vesicle abruptly terminates near the pallial cavity, giving rise to a very narrow vas deferens that crosses the afferent renal vessel dorsally, and becomes exposed in the pallial cavity, lying along the posterior and right pallial cavity edges, gradually becoming thicker (Fig. lOA). No prostate gland differen- tiable. Vas deferens descends to pallial floor at mid length of the pallial cavity. The portion of the vas deferens lying on pallial floor is open (furrow); with tall, thick edges (Fig. lOB), becoming convoluted near the base of the penis. Penis of about 1/4 of pallial cavity volume (Fig. lOB); twisted inwards in basal region, middle region slightly broader, con- stricting gradually up to narrow, rounded tip. Penis duct

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Figure 12. Fuhnentiim ancilla anatomy. A, odontophore, dorsal view, most of the superficial membrane and muscles removed, radular sac deflected to right, both cartilages (oc) deflected from each other, left ni5 turned downwards. B, region of the valve of Leiblein (vl), ventral view. C, region of duct of the gland of Leiblein (Id), ventral view, most tubes opened longitudinally. 1), midgut as in situ, ventral view, position of kidney indicated. E, same, most tubes opened longitudinally to expose inner surface. Scale bars = 1 mm. Abbreviations listed in section with figure captions.

open (a furrow), lying on inner penis edge, relatively deep, runs up to penis tip (Fig. 13A). Terminal papilla in penis tip, about 1/6 of penis length, located inside a chamber formed by terminal portion of penis (Fig. 13B).

Female (Figs. 13C-D). Visceral structures similar to those of males. Visceral oviduct very narrow, running along the middle of the columellar surface of the visceral mass. Visceral oviduct inserting in left side of pallial oviduct, an-

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Figure 13. FiilnienUuii ancilla anatomy. A, penis and adjacent region of its base, dorsal view. B, detail of penis tip, showing, partially by transparency, terminal papilla. C, pallial oviduct, ventral view, detail of its terminal region mostly opened longitudinally, walls reflected. D, entire pallial oviduct, ventral view as in situ, some adjacent structures also shown, a transverse section at indicated level also shown. E, central nervous system, ventral view, esophageal passage indicated (es). F, same, dorsal view. Scale bars A, D = 2 mm; others = 1 mm. Abbreviations listed in section with figure captions.

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terior to albumen gland, opening to vaginal duct. Albumen gland whitish, about 'A length of pallial oviduct, forming a blind-sac with thick walls and flat lumen that is continuous with the lumen of the capsule gland. Seminal receptacle triangular, located between albumen and capsule glands, close to right-dorsal side, no apparent duct, opening directly to vaginal duct between both glands. Capsule gland beige, inner lumen broad and flat (as wide as gland). Vaginal duct lying all along capsule gland left edge, separated from it by a low fold of dorsal lamina ot capsule gland. Capsule gland laminae terminating close to genital pore, without forming a vestibule. Female pore protruded, rounded, located to right of anus. Female pore walls thick muscular, inner lumen flat, curved from left to right, then to left again, expanding gradually, leading to pore (Fig. 13C). Female pore edges thick, preceded by longitudinal, broad folds. No cement gland in sole of foot.

Central nervous system (Figs. 13E-F). Located at base of proboscis, a short distance ventral to the rhynchostome. Very concentrated, practically no individual ganglia distin- guishable. Cerebro-pleural ganglia widely connected to each other along median line. Both also widely connected to pedal ganglia, no connective distinguishable. Pedal ganglia of about the same size as the cerebro-pleural ganglia. Commis- sure between both pedal ganglia relatively narrow and very short (both pedal ganglia maintained in contact). Pair of buccal ganglia small, close to each other, located oblic^uely (left ganglion slightly more anterior) in dorsal region of cerebro-pleural ganglia; connective with cerebral ganglia narrow and short, left connective a little longer and joining another secondary nerve, running anteriorly. Supra- and subesophageal ganglia about half the size of the main gan- glia, located near and ventral to right cerebral ganglion; sub- esophageal ganglion connected to cerebral ganglion by a narrow and short connective; supra-esophageal ganglion connected to subesophageal ganglion by a broad and very short connective, and also to left cerebral ganglion by a narrow and short connective.

Measurements of shells (in mm). NMSA E5279: 64.5 by 35.0.

Distribution. South Africa.

Habitat. Rocks and coarse sand, from 32 to 81 m depth.

Material examined. SOUTH AFRICA. Western Cape; 93 km SE of Mossel Bay, 68.4 m depth, NMSA E2770, 1 $ (no shell) (Exch. C. Marais col., 24/vi/1988), SW of Mossel Bay, Agulhas Bank, 81 m depth, NMSA E5279, 1 9 (Exch. C. Marais col., xi/1988); Struis Bay, 34°47.2'S 20°08.6'E, 32 m depth, NMSA S3578, Id (no shell) (dredged Sardinopsis, 08/vi/1991).

Discussion. Fulmentu?n ancilln has been mostly referred in the genus Pseiidoliva or Sylvanocochlis Melvill, 1903, for which it is type species. Recognition of Fidmentiim is based on the arguments of Vermeij (1998: 60), who considered

both genera {Sylvanococldis and Fidmentum) as synonyms. Kantor’s (1991) anatomical description was used as the ground plan for the anatomical study of this species. Results of the present study generally agree with Kantor’s data; the few cfifferent points include the presence of transversal dis- tinct folds in the hyp)obranchial gland found in his speci- mens (his fig. 12D), but lacking in those examined here.

Genus Melapiiim Adams and Adams, 1853

(Type species Pyrida lineata, by subsequent designation of Cossmann, 1901)

Melapium lineatum (Lamarck, 1822)

(Figs. 9F-K, 4E-F, 14A-17F)

Melapium lineatum: Liltved 1985: 9; Kantor 1991: 39-41 (figs. lA-B, 2B, 17, 18); Hayes 1994: 77-78; Vermeij 1998: 75.

Description

Shell (Figs. 9F-K). Of medium size (about 30 mm), rounded; wider than long. Walls thick. Color cream, with narrow axial bands, dark beige, with irregularly intercalated longer and shorter bands, mainly concentrated in a band along the middle of the body whorl; canal white, with purple pigmentation within the anterior edge. Protoconch flat, dome-shaped, of 1 ‘A whorls; transition to teleoconch indis- tinct (Figs. 9G-J). Spire flat, low, weakly elevated, with about 3 whorls. Suture relatively deep. Body whorl very large, sur- rounding almost completely the penultimate whorl. Surface glossy, lacking sculpture except for weakly visible growth lines. Anterior region with carina surrounding left edge of canal, projected forwards. Aperture rounded (Fig. 9F), lo- cated close to suture, peristome white, gradually becoming orange in interior regions. Outer lip simple, semi-circular; edge thick, rounded. Inner lip bearing thick callus, cover- ing roughly half of the ventral surface. Canal short, broad, relatively deep, projected forwards. Young specimens (about 2 whorls) antero-posteriorly longer, outline elliptical (Figs. 91-1); outer surface opaque, sculptured by a net ot thin and very narrow reticulation of spiral and axial lines (Fig. 91).

Head-foot (Figs. 9K, 14A-B, 14 D, 15B). Head weakly protruded, socket-like; basal region of head as short flap. Tentacles located in both ends of this flap; each tentacle long and narrow, with a broader region just above base; tip pointed. Color mostly beige-cream, with bluish band flanked by a narrow bands of red and yellow surround the dorsal surface of foot at its margins (Fig. 9K); tentacles with pale base and orange middle and distal regions. Eyes located on both ends of head-flap, below the base of each tentacle (Figs. 14A, 14D). Rhynchostome in the form of a transversal slit is located between the tentacles (Fig. 14D). Foot very wide and broad; thick in center, gradually becoming thinner toward the periphery, uniformly in all directions. Furrow of

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Figure 14. Melapiurn lineatiim anatomy. A, head-foot, male, frontal view. B, same, posterior view. C, pallial cavity and visceral mass, male, [;

ventral-inner view. D, head and haemocoel, ventral view, foot and columellar muscle removed. E, Transverse section through pallial cavity |i

roof, at mid-length of the osphradium. Scale bars = 2 mm. Abbreviations listed in section with figure captions. [j

pedal glands very thin, restricted to anterior half of foot edge (Lig. 14A: pg). Columellar muscle of about 1/3 whorl, having broad main flap in middle and right regions; secondary flap taller and longer, at its left end, projected deeper, weakly coiled (Fig. 12A: cm). Male penis relatively small, originating far removed posteriorly from right tentacle (described below). Haemocoel oval, broad, weakly curved to left (Fig. 14D).

Operculum. Absent.

Mantle organs (Figs. 14C, 14E). Mantle border simple.

wide, thick, unpigmented. Pallial cavity broad and short (just over Vi whorl). Siphon long (equal in length to pallial cavity) and slender; edges simple; inner surface with low transversal folds. Siphon base with pair of reinforcements extending as low folds beyond siphon base, parallel to mantle border, longer on right, about 1/3 of mantle border length, gradually diminishing. Osphradium slightly longer than '/2 pallial cavity length and about 1/5 of its width; anterior end rounded, posterior end pointed. Osphradium

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filaments tall, symmetrical, mostly free from attachment to mantle roof (connected mainly to osphradial ganglion), forming a longitudinal concavity along this ganglion. Each filament extends ventrally about twice the diameter of the osphradium ganglion, its edge reinforced by a rod that is weaker along inner regions closer to the ganglion. Gill ellip- tical and broad, slightly shorter than pallial cavity and about half as wide, slightly curved to left. Anterior and posterior ends broadly pointed, anterior end slightly forward of os- phradium margin, and separated from it by a low, broad fold of the siphonal base. Posterior end of gill extends beyond posterior end of osphradium, reaching the pericardium. Gill filaments triangular, relatively low, apex at or slightly to the right of center, rounded; rod broad, lying along left edge, extending slightly beyond the membranous portion of the filament. Ctenidial vein and afferent gill vessel narrow (af- ferent slightly narrower), running along respective gill edges. Hypobranchial gland moderately thick, cream colored, cov- ering most of the area between the gill and rectum (-1/3 of pallial roof), becoming narrow in the region anterior to the anus, surrounding right edge of gill. Rectum relatively broad, running along the right edge of the pallial cavity for about 2/3 of its length. Anus detached, located between middle and anterior thirds of pallial cavity; with small papilla along its left edge. Genital ducts lying between rectum and pallial floor, described below.

Visceral mass (Fig. 14C). Of about 2Vi whorls, rapidly enlarging and almost involute. Right portion of visceral structures encroaching into the right posterior portion of the pallial cavity. Kidney triangular, spanning ~Vi whorl along its right border, partially located inside the middle and right portion of the posterior pallial cavity. Pericardium located slightly to the left of the middle of the posterior portion of the pallial cavity. Stomach located about 1/3 whorl posterior to kidney, occupying about half of adjacent visceral whorl volume. Digestive gland orange, extending from apex to kid- ney, surrounding middle digestive tubes. Gonad within right and columellar surfaces of visceral mass, of the same color as the digestive gland all along its length. Ovary surrounded entirely by digestive gland, becoming internal to it.

Circulatory and excretory systems (Figs. 14C, 15A). Heart volume about 1/3 kidney volume; characters similar to those described for Zemira, auricle small, entirely attached along antero-dorsal region of pericardium; ventricle very large,