THE NAUTILUS

Volume 128, Number 1

April 16, 2014

ISSN 0028-1344

A quarterly devoted

to uuilacology.

9i

8oi

NSli

12-

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The Bailey- Mattliews Shell Museum

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Department of Invertebrate Zoology

National Museum of

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Smithsonian Institution

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Department of Invertebrates

Field Museum of

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North Carolina State Museum of

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THE^NAUTILUS

Volume 128, Number 1

April 16, 2014

ISSN 0028-1344

CONTENTS

Emilio Roliin Three new species of tlie genus Clieilea from the Atlantic Ocean

Raiil Feriiijindez-Garces (Chistropoda: Ilipponiciciae) 1

Kazutaka Amano Miocene vesicomyid species (Bivalvia) from Wakayama in

Robert G. Jenkins southern Honshu, Japan 9

Masaaki Ohara

Steffen Kiel

M.G. Harasewveh On the authorship and primar\’ type of Cerion incdnum

(Ciastropoda: Oriouidae) 18

Maria Dolores Bragado The freshwater mussel collection (Bivalvia; Unionida) of the

Rafael Araujo Museo Nacional de Ciencias Naturales (Madrid, Spain) 22

Arthur E. Bogan

Javier de Andres

Book Review 28

Notice 29

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THE NAUTILUS 128(l):l-8, 2014

Page 1

Three new species of the genus Cheilea from the Atlantic Ocean

(Gastropoda: Hipponicidae)

Emilio Kolan

M useo de Uistoria Natural,

Campus Universitario Norte, Vista Alegre

15782 Santiago de Compostela, SPAIN

er()[email protected]

Raul Fernandez-Garees

Centro de Estiidios Ainhientales de Cienfuego.s (CEAC)

13i\isi6n de Gestion Ainbieiital (DGA)

Calle 17, esq. Ave. 46

Cienfuegos, CUBA

ABSTRACT

Indisidiials from populations presiously thought to represent

CJieilea equestris (Linnaeus, 1758) are studied, showing that

they belong to diffferent species. Three new species are

described and named herein: one from the eastern Atlantic,

another from the western Atlantic and a third one with

amphiatlantic distribution. A lectotv'pe for the ty|re species of

the genus is designated.

INTRODUCTION

Since the latter half of the 20^'* Century, the genus Cheilea

Modeer, 1793 underwent severiil changes both in taxo-

nomic status and systematic position: Thiele (1929) and

Wenz (1940) placed it in Amaltheidae (= Hipponicidae);

Kuroda (1941) in Cdyptraeidae; Ludbrook (1941) in the

family Cheileidae Macpherson and Chaple, 1951, super-

family “Cheileacea.” Iredale and McMichael (1962) and

Kuroda et al. (1971) kept it in Cheileidae. Rehder (1980)

placed it in Crepiduhdae. Vaught (1989) and more

recently Boucliet and Rocroi (2005) placed it in the family

Hipponicidae Trosehel, 1861.

Linnaeus (1758) described the species Patella

equestris from the Indian Ocean. His types, deposited at

the Linnean Society (Figures i-3) seem to belong to two

distinct species, but they do not bear detailed locality

data. Moreover, they are difficult to study due the ero-

sion of their protoconch.

Linnaeus mentioned “O. Indico” as the type locality

for the species; despite that, several populations from

different areas and oceans have been associated to the

name Cheilea equestris. We list some of them herein:

Indo-Pacific: California to Chile (Abbott, 1974); Red

Sea (Sharabati, 1984); Japan (Fukuda, 1990); India

(Subba Rao, 2003); China, Japan, Philippines (Zliongyan,

2004); E. Pacific, Indo-Pacific (Rios, 2009); Easter Island

(Rehder, 1980); Hawaii (Severns, 2011).

Atlantic: Florida, West Indies and Brazil (Abbott, 1974);

Yucatan, Mexico (Yokes and Yokes, 1983); Caribbean Dutch

islands (de Jong and Coomans, 1988); Florida, West Indies,

Brazil: Amapa to Espirito Santo, Fernando de Noronha

Archipelago, Davis Seamount (Leal, 1991); Sao Tome and

Principe (Fernandes and Rolan, 1993); Baliamtis (Redfeni,

2001); Brazil, West Atlantic, Canary, Madeira, Ascension

(Rios, 2009); Cape Verde Archipelago (Rolan, 2005).

Some records for Canary and Madeira (Cosel, 1982)

have not been confirmed by later general works.

Pantoli and Riggieii (1988) mention the species for tlie

Italian Plicxieiie and consider it to be a cosmopolibrn species.

Espinosa and Ortea (2011) described a new genus

(Milicheilea) with shells similar to that of a Cheilea but

smaller and having a very large internal prominence. The

protoconch is apparently similar anti it should be studied

before acceptance of a distinct genus, as the internal

prominence may be induced by the environment (for

example, if the animal lives in cavities resulting in a wide

shell to keep the body closed and occluded, or in acid

environments that can dissolve the external shell while

tire internal one is protected by the soft parts).

Study and comparison of specimens from some of

tliese populations have shown that they represent iliffer-

ent species, which are similar in general externa! appear-

ance only. Some of these species are described as new in

the present paper.

MATERIALS AND METHODS

Descriptions of protoconch moiphology follow the meth-

odology and nomenclature described by Verduin (1986).

Sex of some individuals in this study was inferred based

on attachment of smaller males to the shells of feiniiles

(Redfern, 2001; 48, fig. 206A) Institutional and other

abbreviations used are: AMNH, American Museum of

Natural Histoiy, New York, USA; BMSM, The Bailey-

Mattliews Shell Museum, Sanibel, Florida, USA; lES,

Institute de Ecologia y Sistematica, Havana, Cuba; LS,

Linnean Society, London, UK; MHNS, Museo de

Historia Nattiral, Santiago de Compostela, Spain; MNCN,

Museo National de Ciencias Naturales, Madrid, Sptiin;

MNHN, Museum national d’Histoire iiaturelle, Paris,

France; MNRJ, Museu Nacional/Universidad Federal,

Page 2

THE NAUTILUS, Vol. 128, No. 1

Riode |aiieiro, Brazil; NHMUK, Natural Hi.storv' Museum

United Kingdom, London, UK; CCH, collection of Colin

Hedlern, Boca Baton, Llorida, USA (recently incoiporated

to BMSM); CLC, collection of Baul Lernandez-Garces,

Cienfuegos, (dilia; CPB, collection of Peter Byall, Maria

Biiin, Austria; (iSC, collection of Sandro Gori, Livorno,

Sicilv, Italv; Iv, live-collected; s, empty shell;], juvenile.

SYSTEMATICS

Lamily Hipponicidae Troschel, 1861

Genus Cheilea Modeer, 1793

Chi’ilea Modeer, 1793: llO.

Mitnilaiia Schumacher, 1817: 56.

Lithedaphii.s Owen, 1843; 147.

CdUjptra II. Adams and A. Adams, 1854: 364.

Type Speeie.s: Patella eqiiesiris Linnaeus, 1758, hv

subsecjiient designation (Modeer, 1793)

Cheilea eipiesths (Linnaeus, 1758)

(Ligures 1-8)

Patella ecpiestiis Linnaens, 1758: 780; “O. Indico.”

Type Material: In LS (A-L 0020204). Lectotvpe here

designated (Ligures 1-3).

Original Description: “P. testa iiite^ra orhicnlata,

lahio fomicali petyeadicalari”.

Description: Shell cap-shaped, usuallv vei'v irregular,

with rugose, iixial sculpture. Internally with an erect and

flat curved process that begins under apex. Protoconch

(Ligures 7-8) with 1.75 w4iorls, the first 0.75 wliorl with

slightly rough surface, followed by smooth surface.

Protoconch about 600 pm in miiximum diameter. Best

of surface smooth. Teleoconch begins with about 25-35,

ven’ fine spiral lines.

Dimensions: About 37 mm in maximum diameter.

Distribution: Cheilea ecpiestris has been considered

to be a cosmopolitan species. Lurther studies may well

show that it comprises a complex of several similar-

looking species.

Lectotype Designation: In the LS collection the t\pe

material from Linnaeus evidently includes txvo different

spt'cies. W'e have selected as the lectotype (Ligures 1-3)

Figure.s 1-12. Cheilea species. 1-6. Cheilea ecpiestris (l.innaeus, 1758). 1-3. Patella ecpiestris Linnaeus, 1758. Lectotype selected

from the material in the LS (A-1" ()()2()204), 37 mm in diameter (Indian Ocean). 4-6. Bali, 37.8 mm (MHNS). 7-8. Cheilea cf.

ecpiestris, Panglao, Philippines (MHNS). 9-12. Cheilea striata Nowell-U.sticke, 1959. 9-10. jnvenile, 1.5 mm, Ahaco, Bahamas

(CCR). 11-12. jnvenile watli protoconch, 12.5 mm, Cienfuegos, Cuba (MHNS).

E. Rolan and R. Fernandez-Garces, 2014

Page 3

the shell that most closely agrees with the description

and shells usually associated witli the nominal species.

Remarks: The type locality is Indian Ocean. The Liii-

nean material is badly eroded, so that the protocoiichs

could not be examined. We have presented (Figures 4-6)

a shell from Bali (Indonesia) with the same diameter of

the lectotype and witli part of the protoconch preser\'ed.

This is similar to the protoconch of a juvenile from

Philippines (Figures 7-8).

Cheilea striata Nowell-Usticke, 1959

(Figures 9-12)

Cheilea equestris striata Nowell-Usticke, 1959: 47, pL 2,

fig. 19. [Type locality: Judith’s Fancy, St. Croix, U.S.

Virgin Islands, Atlantic Ocean],

Type Material: Holotype: AMNH (198509). Paratype;

AMNH 294373 (1 specimen). Not examined.

Other Materiai Examined: Cuba: 2 s, 2 j, Rancho

Luna, Cienfuegos, 10-20 m (MHNS). Bcihamas: 3 j, Abaco,

lOm(CCR).

Description: Tlie protoconch (Figures 11-12) has

only one whorl, tlie nucleus is wide, more than 250 pm

an d tl le entire protoeonch is al)()iit 600-700 pm in diam-

eter, with very fine spiral line's. The teleoconch is similar,

with numerous radial striae crossed !)y isolated concen-

tric lines.

Di.stributior!: Known from the Virgin Islands (t\pe

locality), Bahamas, and Cuba.

Kemarks: Nowell-Usticke (1959) described tins species

as a “fomi” having “strong raised radiating ribs roughly

equal in size. On some shells these radiating ri!)s cross

rather weak concentric lines”. Nowell-Usticke (1959)

referred to material ex-Usticke Collection (AMNH

198509) and Schwartz Collection (AMNH 294373). The

repository of additional paraty|3es is unknown; its exis-

tence is indicated by Usticke (1959: 47) and the size

range for the taxon is given as 20-40 mm length. Faber

(1988: 90) considered this taxon to be a synonym of

C. equestris (Linnaeus, 1758); we cannot agree with

this view due to the consistent differences in proto-

conch morphology.

Figures 13-22. Cheilea arnericana new species. 13. Holoty|3e (female) (BMSM 17939) and paratype (male) on the dorsum, 19 and

7 mm, Abaco, Baliamas, ex-CCR. 14. Ventral wew. 15. Male on the dorsum of the feinHe shell. 16. Protoconch of female (holotype).

17. Protoconch of male (paratype). 18. Protoeonch of a juvenile, Abaco, Bahamas (CCR). 19. Paratyyre, juvenile 2.4 mm. Rancho

Luna, Cuba (MHNS). 20. Paratype, juvenile, 3.5 mm, Maria la Gorda, W Cuba (MHNS). 21-22. Protoconchs in two positions

(Figure 21 from the shell of the Fig. 19).

Page 4

THE NAUTILUS, Vol. 128, No. 1

Redfern (200]) illustrated tlie protoconcli of Baha-

mian specimens externally and internally. These are very

distinctive from the protoconch of C. equestris. Shells

from Cu!)a are equal to those from Bahamas. Other

species of this genus can he distinguished by their dis-

tinctive protoconchs.

Cheilea americana new species Rolan, Redfem,

and Fernandez-Garces

(Figures 13-18, 42^3)

Ty]je Material: Holotcpe (female, sp) and a parahqre

(male, sp) in BMSM (17939) (Figures 13-14). Other

paratyi^es: MNCN (15.05/60102, 1 j); MHNS (100593,

1 j. Figure 16) both from Maria la Gorda, 10-15 m,

W' Cuba); MNCN (15.05/60103, 2 s); MNHN (IM-

2012-2112, 3 j), OCR (2 j), CFG (2 s, 2 j) and lES (1 s,

1 j), all from Rancho Luna, 10-20 m, Cuba).

Type Locality: Reef off Great Guana Cav, 26°42'30" N,

77°09'46" W, Abaco, Bahamas, 4.5 m.

Etymology: The specific name refers to the conti-

nents off w'liich the species was collected.

Other Material Examined: 1 sp, Chapeu Nordeste

Island, Fernando de Noronha Archipelago, Brazil (MNRf).

Description: Shell rather irregular, conical-depressed,

as an inverted cup, whitish, with numerous ratliating ribs.

Protoconch (Figures 16-17) with 1.75 whorls, 530 pm in

maximum diameter, nucleus about 100 pm. First 0.75

wliorl very rough, subsequent whorl with zig-zag lines,

often eroded (Figure 18). Initial part of the teleoconch

with about 16-23 spiral cordlets; teleoconch expands

(|uickly to partially envelop the protcK-onch. Rest of

teleoconch veiw irregiihir, with prominences, undulating

areas, and concentric scaly lines. Edge very iiregular and

inner surface witli a prominence. Internally with an erect

cui^’ed process, which l)egins under apex and is character-

istic of genus.

Radula (Figures 42-43) with very wide rachidian

teeth, each with very prominent central cusp and 8-9

smaller lateral cusps. Lateral teeth elongated, with about

5 cusps along cutting edge, most prominent one very

large and wide; inner marginal teeth elongate, with wide

and prominent cusp and fG8 verv small fine cusps on

each side; outer marginal teeth similar to inner marginal

teeth but narrower and witli less cusps.

Figures 23-32. Cheilea species. 23-27. Cheilea atlantica new species. 23. Holotype, 7.2 mm, Montague Seamount, Brazil (MNRJ

2.5542). 24. Protoconch. 25. Detail of the iiiicrosculpture. 26. Shell, 3.2 mm, Lagoa Aziil, SaoTome I. (MHNS). 27. Protoconcli. 28-

32. Cheilea africana new species. 28-29. Paratyjies, 2.7, 2.57 min, Regona, Cape Verde Archipelago (MHNS). 30-32. Protoconcli

and detail.

E. Roliin and R. Fernaiulez-Garces, 2014

Page 5

Dimensions: Shell of female indiGdual 19 mm in m:ix-

inium diameter; m;ile vvitli 7 mm maximum diameter.

Distribution: Brazil, Bahania.s, Cuba.

Eemarks: Cheilea equestris has a more finely striated

sliell (Figures 1-3); the protoconch is larger (about

650 pm. Figure 7) and is less sculptured (Figures 7-8).

For the differences wdth the other species see below.

Cheilea atlantica new species Rolan, Leal, and

F ernandez-Garces

(Figures 23-27)

Type Material: Holotype (Fig. 23) in MNR| (25542).

Paratypes: MNCN (15.05/60101, 1 s) from Santo

Antonio de Pale, Annobon, Equatorial Guinea; MHNS

(100594, 1 j, Figure 26) Lagoa Azul, Sao Tome.

Other Material Examined: Cuba: 1 j. Rancho Luna,

Cienfiiegos, 10-20 m (MHNS).

Type Locality: Montague Seamount, Brazil.

Etymology: The specific name refers to the Ocean

where the species was collected.

Description: Shell very irregular, appro.xiinatelv coni-

cal, solid, wliitish with radial ril),s. Frotoconcli (F’ig-

ures 23, 27) with 2.75 whorls, niaxiiimm diameter

800 j.UTi. Nucleus with about 130 pm. Nucleus and fol-

lowing 0.5 whorl lacking sculpture or c)nly veiw liglitly

sculptured. Reticulated sculpture appears after nucleus

and following 0.5 whorl. This lasts for 1.5 whorl. Last

protoconch 0.5 whorl witli very fine spiral striae and

growth lines. Beginning of teleoconch with numerous

(about 50-55) light spiral cordlets, which gradually sepa-

rate: a group of these cordlets l)orders tlie protoconch on

each side while others continue in front and centrally at

the end last protoconch whorl. Internal shell surface

with the erect and curved process, which begins under

the apex and is characteristic of the genus.

Dimensions: HoIoty|3e 7.2 mm in diameter; a

paratype (MNCN) reaches 11.7 mm in ma.\imum diam-

eter. The species may reach larger sizes, but one would

need to see a complete shell with well-preserved

protoconch ensure that it is actually the present species.

Figures 33—43. Cheilea species. 33-38. Cheilea africana new species. 33. Juvenile, 2.3 mm, Sao Tome 1. (MHNS). 34. juvenile,

2.4 mm, Principe (MHNS). 35-38. Protoconchs, Lagoa Azul and Esprainha, Sao Tome Island. 39-41. Cheilea africana new species.

39: holotype, 11 mm, Regona, Sal 1., Cape Verde Archipelago (MNCN); 40: juvenile, internal face, 2.6, Miamia, Ghana; 41: juvenile,

1.3 min, Lagoa Azul, Sao Tome Island (MHNS). 42-43. Radula of Cheilea aniericana new species, Chapeu Nordeste Island,

Fernando de Noronha, Brazil.

Page 6

THE NAUTILUS, Vol. 128, No. ]

Distribution: This is an Ainphiatlantic species known

from Brazil (liolotvpe), Cuba (a juvenile), and from the

islands of the Gtiinean Gulf (Sao Tome and Annobon Is.)

off the West African coast.

Remarks: CJieilea (ithintiai new species has a multi-

spiral protoconch with more whorls than otlier known

species and hpical microsculptiire not present on nucleus

and on first 14 wliorl, continuing w-itli a reticulated section

;uid ending on a distinctive section with veiy light sculp-

ture. The teleoconch !)egins witli veiy numerous light

spiral lines. Since it the teleoconch is so varial)le, only tlie

study of die protoconch allows for its conchologiciil sepa-

ration from other species. Apparently this species is li\ing

sympatrically with Cheilea africana in Sao Tome and

Annobon in West Africa and probably with C. americana

new species in Brazil. The protoconclis of these two latter

species are shorter, with only 2 and 1.75 whorls, respec-

tively. In addition, the beginning of tire prot(X.'onch of

Cheilea atlantica new species is more sculptured.

Cheilea africana new species Rolan and Fernandez-

Garces

(Figures 28-41, 44-48)

Type Material: Ilolotvpe (15.05/60100, s. Figure 39)

and 4 paratypes (15.05/60100, s) in MNCN, all from

t\pe localih'. Parahpes in following collections: MNHN

E. Rolan and R. Fernandez-Garces, 2014

Idige 7

(IM-2012-2013, 6 s), Praia, Santhiago, Cape Verde;

MHNS (100595, ex-12389, 5 s) (100595, ex-7031, 1 sp,

7 s) Mordeira, Sal Island, Cape Verde; CPR (1 s) Santa

Maria, Sal Island, Cape Verde; CFG (1 s), Sal-Rei,

Boavista Island, Cape Verde; MIINS (100595, 2 s.

Figures 28-29), Lagoa Azul, Sao Tome Island, Sao Tome

and Principe; CSG (6 j) Minerio, Sao Tome Island, Sao

Tome and Principe, 41 m deptii.

Type Locality; Regona, Sal Island, Cape Verde.

Other Material Examined: Senegal: 4 j, Dakar, 15 in

(MHNS). Cape Verde Archipelago: 9 j, Regona, Sal I.

(MHNS); 1 s, Palmeira, Sal 1. 30 m (MHNS). Ghana:

6 j, Miamia, 25-35 m (MHNS). Sao Tome and Principe,

Principe Island: 4 j, Santo Antonio, 4-10 m (MHNS). Sao

Tome and Principe, Sao Tome Island: 6 j, Minerio, 35-

41 m (MHNS); 6 j, Lagoa Azul, 15 m (MHNS); 1 s,

Esprainha, 8 m (MHNS). Aiinobon: 1 j, San Antonio de

Pale, 5 m (NHNS).

Etymology: The species is named after the continent

where the species was collected.

Description: Shell very irregular, approximately coni-

cal, solid, dirt-whitish. Protoconch about 650 maximum

diameter, with about 2 whorls; embryonic shell well-

defined including: nucleus measuring 140 pm and first

half-whorl with rough and very irregular sculpture with

rounded depressions, following 0.25 whorl presenting

about 5-6 strong spiral lines with small threads between

them. Separation between these tv/o parts is well-

defined. Last 1.25 whorls are smooth and have only

growth lines. As can be seen internally in juveniles (Fig-

ures 40-41), its surface originally present numerous

irregular spiral lines, which later disappear as a result of

erosion. Beginning of teleoconch with numerous spiral

lines (about 30-35) which gradually separate: a group at

each side bordering the protoconch and another con-

tinuing centrally in front of last section of last protoconch

whorl. Entire external surface irregularly striated, pre-

senting numerous elevations, shell edge very irregular,

conforming to substrate irregularities. Internally part

with the erect curved process, which begins under the

apex and is characteristic of the genus. Soft parts not

observed. Radula (Figures 44-48) v/ith wide central

teeth with very prominent central cusp and 8-9 smaller

lateral cusps; lateral elongate with about 6 cusps along

cutting edge, one larger, plus 2-3 smaller outward; mar-

ginal teeth elongate, with a vride and prominent cusp

and many, very small lateral cusps.

Dimensions: Largest shell measures 27.8 mm; the

holoty|3e is 11 mm.

Distribution; West Africa: Cape Verde, Senegal,

Ghana, Sao Tome and Principe, and Annobon.

Eemarks; Cheilea africana new species may be distin-

guished from C. striata Nowell-Usticke, 1959 as this

latter species has a protoconch with less than one whorl

and a wider nucleus. Chdlea aniericana new' species has

a very similar protoconch, l)ut witli only 1.5 whorls, w'ith

more marked, zigzag spiral lines (w'lieii not eroded); fur-

thermore, the protoconch is 100 pm smaller in diameter.

The shell is very irregular externally !)ut tlie radial striae

are less visible. Cheilea atlantica new' species has a

protoconch with 2.75 wliorls, the diameter is larger

(more than 100-150 pm difference between C. africana

and the larger C. atlantica), and tlie protocoiieh sculp-

ture is different with a nucleus and 0.5 whorl almost

smooth and subsequent 1.5 whorls strongly reticulated.

Coiiclusicms; Cheilea equestris (Linnaeus, 1758) has

been considered as a cosmopolitan species. The present

study shows that more than one taxon had been lumped

under that nominal species, most of them on the liasis of

shell characters only. This study may encourage similar

efforts for additional populations living in areas

geographically removed from the type locality of the

nominal species. Such efforts will inevitably reveal

additional unnamed species. It seems that, if nothing

else, SEM studies of protoconchs and comparison wdtli

previously known species could be a fine method

of separation.

ACKNOWLEDGMENTS

The authors thank all those who loaned material for

study (Colin Redfern, Jose H. Leal, and Sandro Gori).

Thanks are also due to Jesiis Mendez and Ines Pazos

from the Centro de Apoyo Cientffico y Teenologico a la

Investigacion (CACTI) of the University of Vigo who

made the SEM photographs; and to Antdnio A.

Monteiro for the English revision. We are indebted

to Kathie Way (NHMUK) for the images of

the lectotype.

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THE NAUTILUS 128(1);9-17, 2014

Page 9

Miocene vesicomyid species (Bivalvia) from Wakayama

in southern Honshu, Japan

Kazutaka Amano

Department of Geoscience

Joetsu University of Education

Joetsii 943-8512, JAPAN

[email protected]

Masaaki Ohara

Wakayama Prefectiiriil Museum

of Natural History, Kainan City

W'akayama 642-0001, JAPAN

[email protected]

Robert G. Jenkins

School of Natural System

College of Science and Engineering

Kanazawa University Kanazawa City

Ishikawa 920-1 192, JAPAN

rol)ert [email protected]

Steffen Kiel

Georg- August University Gottingen

Geoscience Center, Geobiology Group

Goldschinidtstr. 3

37077 Gottingen, GERMANY

[email protected]

ABSTRACT

A fossil association of potentitilly chemosymbiotic bivalves is

reported from the lower Miocene Shikiya Formation in Kii

Oshima Island, Wakayama Prefecture, Japan. The association

is dominated by the elongate vesicomyid species Adulomija

uchimuraensis (Kuroda, 1931); a second vesicomyid of lower

abundance is here described as Archivesica sakoi new spe-

cies. It represents the oldest species of the genus Archivesica

and has a similar hinge structure as “Calijptogena” lauhieri

(Okutani and Metivier, 1986). Additional faunal elements

include chemosymbiotic bivalves, namely the solemyid

Acharax cf tjokosukensis Kanie and Kuramochi, 1995, and

the lucinids Lucinonui? sp. and Pomma? sp., as well as

the naticid gastropod Euspim meisensis (Makiyama, 1926).

This association of fossil bivalves till with well-known

chemosymbiotic li\4ng relatives suggests that these specimens

lived at an ancient cold seep, and their mode of occurrence

as mostly articulated shells parallel to the bedding plane

indicates that were transported a short distance and then

quickly buried.

Additional keywords: Miocene, Archivesica, Adulomtja

INTRODUCTION

Vesicomyidae is one of the six extant bivalve families

with a chemosymbiotic mode of life, with have a fossil

history ranging back to the middle Eocene some 47

Million years ago (Kiel, 2010a). The vesicomyid fossil

record of the Northwestern Pacific region is increas-

ingly well studied and revealed the successive appear-

ance, dominance, and disappearance of several genera.

Huhertschenckia Takeda, 1953 was the only genus present

during the late Eocene and Oligocene and it disap-

peared afterward; the elongate Adtilomya Kuroda,

1931 dominated during the early and middle Miocene;

and in the late Miocene both Archivesica Dall, 1908

and Calyptogena Dall, 1891 appeared and are the

dominant vesicomyid genera until the present day

(Amano and Jenkins, 2011; Amano and Kiel; 2007;

2010; 2011; 2012; Sasaki et al. 2005). Among those

two genera, Archivesica was, and still is, much more

species rich than Calyptogena .

When Amano and Jenkins (2011) investigated the fos-

sil record of extant vesicomyid species in Japan, they

questioned an identification by Katto and Masiida

(1978) of an early Miocene species from the lower

Miocene Shikiya Formation in southern Honshu as

Akebiconcha cf kawamurai Kuroda, which is an older

name for die extant Archivesica kawamurai . Amano and

Jenkins (2011) suspected that the Shikiya species may

belong to Archivesica, but was not A. kawamurai. More-

over, Katto and Masuda (1978) identified an elongate

bivalve from the same locality as Cultellus izunwensis

Yokoyama, although the lot documentation was poor.

Based on observations on the original material of Katto

and Masuda (1978) and new material collected at their

locality, both species are here identified as members of

the Vesicomyidae: the ovate specimens previously iden-

tified as Akebiconcha cf. kawamurai is named as a new

species of the genus Archivesica, and the specimens

assigned to Cultellus izumoensis are identified as

Adulomija uchimuraensis Kuroda, 1931. Here we

describe these species and discuss their evolutionary' sig-

nificance. We also describe the locality in more detail,

including its paleoecology and the mode of occurrence of

these fossils and additional taxa.

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THE NAUTILUS, Vol. 128, No. 1

MATERIALS AND METHODS

Tlu' .specimens were recovered from a large l)l()ck (ca.

7 X 4 X 4 in) and several small blocks in its viciniW along

the boulder beach near a cliff about 400 m north of

Oshima fishing port in Kii Osliima Island, Wakayama

Prefecture, Japan (Figure 1; Katto and Masnda, 1978,

Loc. no. 10). These blocks are derived from the cliff and

consist of sandv siltstone of the uppermost lower to low-

ermost middle Miocene Sliikiva Formation of tlie

Kmnano (Ironp (Hisatomi, 1981). All shell material is

dissolved anti rubber casts were made from the internal

and e.xternal molds to document the internal and exter-

nal morphologx' of the species, including the hinge struc-

ture. However, nianv small pvrite grains and the rough

surface of weathered shells prevent us from making rub-

ber casts without bubbles. All specimens reported here

are depositt'd at the W'akayama Prefectnral Museum of

Natural llistoiT (W'MNH).'

MODE OF OCCURRENCE

The bivalves occur in si lell beds consisting of mostly

articulatetl specimens lying parallel to the bedding plane,

although \alves that are opened to various degrees were

also found (Figures 2-5). The matrix of these shell beds

does not differ from the surrounding sandv siltstone.

This mode of occurrence indicates that shells are not

preserved in life position but may instead have been

transported for a short distance and then quickly buried.

In addition to the two vesicomyid species described

here, two specimens of the giant solemyid Aclwrax cf

t/okosukensi.s Kanie and Kuramochi, 1995 (reaching

about 30 cm in length), one specimen each of the lucinid

bivalves Ijicinonm? sp. and Pomnea? sp., and of the

naticid gastropod pAispira ineisensi.s (Makiyama, 1926)

were found (Figures 6—10).

SYSTEMATICS

Family \Tsicomyidae Dali and Simpson, 1901

Sulifamilv Pliocardiinae Waiodring, 1925

Genus Arc/iit'esica Dali, 1908

Txpe Species: Cdllocardia gig,as Dali, 1896, from the

Gulf of California.

Archivesica nakoi new species

(Figures 1 1-17)

Akehiconclia cf. kaivainurai Kuroda. Katto and Masnda,

1978: pi. 3, fig. 7.

Diagnosis: Medium-sized Archivesica with elongate

ovate shell, no lunular incision, shallow and wide pallial

sinus, and very shallow subumbonal pit; snbumbonal

cardinal tooth consisting of ver\’ small anterior (3a) and

rather thick posterior (3b) teeth and thick cardinal tooth

Locality

LocalitvtoTOmano):eu^.(20.'l 3),

Toshimasaki

Oshima fishing. port

^Kii Oshinna Island

Oshima port

IS Kushimoto Bridge

Figure 1. Localitx' of the vesicoinvid fossils (left-side base map is from the “Kushimoto”, scale 1:25,000 topographic map published

by tlie Geospatial Information Authoritv' of japan).

K. Ainano et al., 2014

Page 1 1

Figures 2—5. Field images of the outcrop. 2—4. Large block yielding vesicomyid fossils, white arrow indicates a gregarious

occurrence; enlargement of the southern side of the block (3), and close up of the gregarious occurrence (4). 5. Cluster of Acltdomi/a

uchimuraensis Kuroda in a small block in the immediate Acinity of the large block.

(1) in right valve; subumbonal cardinal tooth consisting

of thin anterior ramus (2a) and very thick triangular

posterior tooth (2b) and posterior cardinal tooth (4b)

very thin in left valve.

Description: Shell medium-sized for genus (up to

80.4 mm in length), moderately inflated, elongate-ovate

(height/length = 0.48-0.56), equivalve and inequilateral.

Antero-dorsal margin concave and continuing to rounded

anterior margin; postero-dorsal margin nearly straight,

posterior margin subtruncated; ventral margin broadly

arcuate. Beak prominent, prosogyrate and located at ante-

rior one-fifth to one-tliird of shell length (U%=22-34).

Luiiule absent. Surface ornamented with imegiiiar corn-

marginal lines. Right valve hinge vride for size, with three

cardinal teeth and very shallow subumbonal pit. Anterior

tooth (3a) in right valve very short and thin, connected

with posterior tooth (3b). Posterior tooth (3b) rather tliick,

oblique posteriorly; middle cardinal tootli ( 1 ) moderately

thick, oblique anteriorly. Left valve hinge wide, with three

cardinal teeth. In left valve, anterior tooth (2a) thin, con-

nected with middle tootli (2b); middle tootli (2b) triangu-

lar and very thick; posterior cardinal tooth (4b) very thin.

Anterior adductor scar semicircular and deeply excavated;

anterior pedal adductor scar narrow, quadrate and deeply

excavated, distinct from adductor muscle scar; posterior

adductor sear indistinct, ovate, situated directly above a

weak ridge riiniiiiig from umbo to postero-ventral corner;

pallial sinus shallow and wide.

Type Material: Holotype: Right valve, length, 63.2 mm,

height, 31.3 nun, WMNH-Ge-5. Paratypes; Left valve,

length, 48.7 mm+, height, 26.0 nini, WMNH-Ge-6;

left valve, length, 59.5 mm, height, 30.5 mm + , WMNH-

Ge-7; left valve, length, 80.4 mm, height, 39.2 mm,

WMNH-Ge-8.

Type Locality: The coastal near cliff about 400 ni

north to Oshima fishing port in Kii Osliinia Island,

Kushimoto-clio, Wakayama Prefecture, fapaii.

Page 12

THE NAUTILUS, Vol. 128, No. 1

Figures 6—10. Mollusks as.sodated with tlie vesicoimicl hi\alve.s described liere. 6, 7. The luciTiid bivalve Pouniea? sp. (W’MNH-

Ue-1 120210284): 6. rubber cast of the iiuier surface of a left valve showing elongate anterior adductor .scar (AAS); 7, intenuil mold. 8.

'fhe naticitl gastropod Euspira meisetisi.s (.Makivaina) (\\’M\H-Ge-1 120210285), rubber cast. 9. The lucinid bivalve Lucinoma? sp.

(\\’MNH-Ge-1 1202 10287), rul)ber cast of the external surface of a right \aK'e. 10. The solemyid bivalve Acharax cf t/okosukensis

Kanie and Kuramochi (WMNU-Ge-1 120210291), internal mold of a left valve.

Material Examined: Twelve .specimens from the

tvpe locality.

Remark.s: Archivesica sakoi new species wtis prexionsly

reported as Akehianu'lui cf. kaicaDuinii Kuroda, 1943 by

Katto and Masuda (1978), but their illustration did not show

tlie hinge stnicbire. The outline oi Arc! livesica hnoamurai

is indeed somewhat similar to that of A. sakoi except for

being liigher and larger. However, the subumboniil cardiniil

tooth of right N'iilve enables ns to easily separate botli spe-

cies. The riglit v;ilve of the present new species has a sm:ill

anterior tootli (3a) and posteriorly obli(|ue posterior tooth

(31)), while that of A. kaioaniurai lias a larger anterior tootli

and a posterior tooth that is slightly inclined anteriorly

(Kuroda, 1943; Sasaki et al., 2005).

Comparisons: The riglit salve hinge of Arcliivesica

sakoi is most similar to that of the Recent “Cahjptogena”

laabieii Okutimi and Metivier, 1986 in having a small 3a

tooth and a shallow subumbonal pit (Figure 18). A recent

molecular phylogenetic analysis (Audzijonyte et ak, 2012)

indicates that tliis species is phylogenetically close to the

t\pe species oi Arcliivesica, A. gigas (Dali, 1895). How-

ever, “C.” laiilneri cmi lie separated from the present new

species by basing a distinct blunt ridge running from beak

to posterior end, a tapering posterior eiul, a very narrosv

hinge plate, a less stout middle tooth of left vals-e, and

many distinct iiTegular grossih lines on the outer surface

(Figures 19, 22). The extant Arcliivesica ochotica Scarlato,

1981 also resembles this nesv species in a having broadly

rounded ventral mai'gin, a shallow palliiil sinus, a rather

small 3a tooth, a shallow subumbonal pit in the right valve,

and a stout 2b tootli in the left s’alve (Figures 20, 21, 23).

However, A. ochotica has a larger (]()5.() mm long) and

higher shell (H/L = 0.58), an overhanging posterior tooth

above anterior and middle ciu'dinal teeth of left valve, and

K. Ainaiio et al., 20] 4

Page 13

Figures 11 — 17. Archivesica sakoi new species. 11, 12, 15. Faratyj^e (WMNH-Ge- 1 1202 10286); 1 1, a rubber cast showing details

of the hinge of a right valve; SP=subuinboiui] pit; 12, ventral \aew showing the anterior pedal retractor scar (APRS); 15, internal

mold of left valve. 13, 16. Paratyjie (WMNH-Ge-1120210288); 13, rubber cast showing details of the hinge area of a left \alve;

16, view on the left valv'e of an internal mold, white arrow indicates the ver\- shallow pallial sinus; PAS=posterior adductor scar.

14. Ilolotvpe (WMNH-Ge-1 120210283); view on the right valve of an internal mold; AAS=anterior adductor scar. 17. Paratvpe

(WMNH-Ge-1 120210289), rubber cast showing the outer surface of a left valve.

a more rountUy curved po.ster()-dor.sal margin than

Archivesica sakoi new' specie.s.

Distribution: Lower Miocene Shikiya Formation at

the tvqoe localit)'.

Etymology: Named after Mr. Yukio Sake who col-

lected the type specimens and kindly offered them to

the authors for study.

Genus Adulomya Kuroda, 1931

Type Species: Aduloim/a uchiumraensis Kuroda,

1931, from tlie middle Miocene Bessho Formation, cen-

tral Honshu, Japan.

Adulomya uchimiiraensis Kuroda, 1931

(Figures 24-29)

Adulomya uchiinuraensis Kuroda, 1931; 27-28, pi. 13, figs. 1 1 1-

114; Tanaka, 1959; 117-118, pi. 1. fig. 1-10; Tanaka, 1960:

24—26. pi. 32, figs. 1-7; Amano and Kiel, 2011: 77-80, fig. 2.

Page 14

THE NAUTILUS, Vol, 128, No. 1

Figures 18, 19, 22. “Cahjptogena” lauhieri Okutani and Metivier, 1986, H()lot}'j3e (NSMT Mo 64158); 18, hinge dentition

of a right valve; SP=siihiiinbonal pit; 19. hinge dentition of a left valve; 22, right side \iew of the holoLpe. Figures 20, 21, 23.

Archivesica ochotica Scarlato, 1981, Holotvpe (ZIN AN SSSR no. 9912); 20, hinge dentition of a right valve; 21, hinge dentition

of a left valve; 23, left side view of the holoHpe.

Cahjptogemi {Adttlomija) udiimuraensis Kiiroda. — Kanno and

Tanaka in Kanno et ah, 1998: 20-22, figs. 7-8.

Cali/ptogena {Adulomtja) uchimuraensis kurodai Kanno and

Tanaka in Kanno et ah, 1998: 22-25, figs. 9-10.

Akehiconcha chitanii (Kanehara). — Kanno and Ogawa, 1964:

pl.l, figs. 17-18.

~ AduJomifa uchimuraensis Kiiroda. — Hayashi and Miura,

1973: pi. 1, fig. 15.

Cultelhis iziimoensis Yokoyania. — Katto and Masuda, 1978:

pi. 3, figs. 8-9.

non Akehiconcha uchimuraensis Kuroda. — Matsiiinoto and

Hirata, 1972: 755-757, pi. 1, figs. 1-8, pi. 2, figs. 1-2.

Material Examined; Fifty'-three specimens were

examined.

Hemarks; Katto and Masuda (1978) illustrated two

elongate specimens as Cidtellus iziiitmensis. Our exami-

nation of the liinge of their illustrated specimen (Katto

and Masuda, 1978: pi. 3, fig.9; Figures 15, 19 herein)

indicates that they belong to Adulomija uchimuraensis

for the following reasons: the left valve has an anterior

(2a) and a middle cardinal (2b) tooth and a low posterior

tootli (4b), and the right valve has onlv two teeth (cardi-

nals 1, 3b); the anterior retractor scar is very deep, and

some examined specimens show that the pallial line is

entire (Figure 16). Such characteristics are never seen

in any cultellid species. Cultellid species have only

two protruded cardinal teeth in each valve and pallial

sinus present.

Comparison: As discussed by Amano and Kiel (2011),

Adulomija uchimuraensis can be separated from the

early to middle Miocene Adulomtja chitanii Kanehara,

1937 by its larger size and the lack of a pallial sinus.

Although the maximum size of Ad ulointf a uchimuraensis

at Kii Oshima (94.1 mm long) is smaller than the maxi-

mum size of the specimens of A. uchimuraensis at its

type locality in the Bessho Formation (180 mm; Amano

and Kiel, 2011), it is still larger than A. chitanii,

which reaches only up to 70.4 mm in length (Amano

and Kiel, 2011).

Measurements: Left valve, length, 83.2 mm, height,

21.4 mm, WMNH-Ge-9; right valve, length, 89.7 mm,

height, 17.8 mm, WMNH-Ge-10; right valve, length,

94.1 inm, height, 23.9 mm, WMNH-Ge-11.

Distribution; Lower Miocene Shikiya Formation in

Wakayama Prefecture, lowest middle Miocene Takinoue

Formation in Hokkiiido and middle Miocene Bessho

Formation in Nagano Prefecture.

K. Amano et al., 2014

Page 15

Figures 24=29. Adulmmja uchimuraemis (Kiiroda), 24, 26,

28. Right valve (WMNH-Ge-1 120210293); 24, rubber cast

showing the hinge dentition and the anterior pedal retractor scar

(APRS) of a right valve; 26, rubber cast showing the anterior

(AAS) and posterior adductor scars (PAS) in a right vidve, white

arrow indicates the posteriormost portion of tlie entire pallial

line; 28, rubber cast showing the outer surface of this shell. 25,

29. The specimen illustrated by Katto and Masiida (1978, pi. 3,

fig. 9)(WMNH-Ge-1 120210292); 25, rubber cast showing

details of the hinge area; 29, left valve. 27. Rubber cast of the

outer surface of a right valve (WMNH-Ge-1 120210290).

DISCUSSION

Recent studies on presumed Paleogene members of

Archivesica from western North America (Amano and

Kiel, 2007; Kiel and Amano, 2010) indicate that these

species may not belong to Archivesica but instead to the

genus Pliocardia or a related new genus (Amano and

Kiel, 2012). Thus, the early Miocene Archivesica sakoi

new species described here from southern Japan repre-

sents the oldest species of the genus Archivesica . How-

ever, this does not change our earlier observation that

Adulomija was the dominant vesicomyid taxon during

the early and middle Miocene in Japan because

Adulomtfa uchinmraensis is far more conmion at the

Kii Osliiina site than Archivesica sakoi.

The association reported here from Kii Osliiiua

consists almost exclii.sively of potentially clienio.syni-

biotic bivalves (e.g., Taylor and Glover, 2010): domi-

nant are the vesicoinyids Adidonitja ucliiiuaraetisis

and Archivesica sakoi with a minor number of lucinids

(Lucinonia? sp. and Poamea? sp.) and a solernyid

(Acharax cf. yokosukensis). Such an association domi-

nated by vesicomyid bivalves is tvpical for extant cold

seep communities (Pauli et al. 1985; Le\’in 2005; Majima

et al. 2005; Campbell, 2006) and suggests that the

Kii Oshiina fauna miglit have lived at an ancient cold

seep. Predatory gastropods siicli as the naticid Euspira

nieisensis found along with the cliemosymbiotic bivalves

are common in soft sediments in many marine environ-

ments, but they are also frecpuently found at ancient

cold-seep communities, especially in the Nortli Pacific

realm (Amano et al. 2010; Kiel, 2010b). Due to the

transported nature of the association and the lack of

carbonate, the usual approach to identify an ancient

cold-seep deposit by stable carbon isotope analysis

(cf. Peckmann and Thiel, 2004) is not possilile. This is

so far the only record of a vesicoinyid-dominated

faunule from the Shikiya Formation and from southern

Honshu (cf., Majima et al. 2005).

Only about 4 km west of the Kii-Oshima locality, a

hydrocarbon-seep fauna has been recently been reported

by the present authors from the upper Eocene to lower

Oligocene(?) Tanamigawa Formation at Tanosald

(Amano et a!., 2013). That Paleogene community differs

from the early Miocene fauna from Kii Oshiina

reported here by consisting of a different vesicomyid

genus {Hubert.schenckia) , the batliymodiolin mussel

Bathijmodiolus , the thyasirid bivalve Conchocele and the

absence of lucinids. As already pointed out by Amano

et al. (2013), Paleogene seep communities in Japan

generally lack lucinid bivalves, wliile lucinids are generally

present at Miocene seep coinmimities as in the the

Kii Oshima fauna reported here.

ACKNOWLEDGMENTS

V/e thank Yukio Sako (Kushimoto-cho), Yiikito Kurihara

(Mie University) and Hiroshi Saito (National Science

Museum) for their help in examining some fossil or

recent specimens; Elena Krylova (P.P. Shirshov Institute

of Oceanology, Russian Academy of Sciences) for

discussions on vesicomyid systeniatics, for her helpful

review of the manuscript, and for allowing us to use !ier

images of A. ochotica. An anonymous review'd' is

thanked for suggestions that helped iinprov'e on the

manuscript. This study was partly supported by a Grant-

in-aid for Scientific Research from the Japan SocieO for

Promotion of Science (C, 23540456, 2011-2013) to KA,

Grant-in-Aid for JSPS Fellows to RGJ, and by the

Deutsche Forscliungsgemeinschaft through grant

Ki802/6-l to SK.

THE NAUTILUS, Vol. 128, No. 1

Page 16

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THE NAUTILUS 128(1):18-21, 2014

Page 18

On the authorship and primary type of Cerion incanum

(Gastropoda: Cerionidae)

M. G. Harasewyeh

Department of Invertebrate Zoolog}'

National Museum of Natural Histor\'

Smithsonian Institution

F.O. Box 37012

Washington, DC 20()13-7()12 USA

1 1 arasewych @si . edu

ABSTRACT

Although the author of the species Cerion incatinni (originally

proposed as Pupa incana) has been cited as Binney, 1851, a

re\iew of his multiple uses of the name do not satisfy the

criteria of availability set forth by the International Code of

Zoological Nomenclature (Article 12), until the publication of

illustrations of three shells labelled Pupa incana in 1857. In a

separately authored paper pul)lished as part of Binney’s multi-

volume work on the Terrestrial Mollusks of the United States,

Leidy published detailed descriptions and illustrations of

several organ systems oi'Pupa incana in 1851. Thus, the author-

ship o( Pupa incana must lie attiilnited to Leidy, 1851, rather

than to Binney. As Leidy did not retain the dissected tissues,

there is no sunixing type material for Pu])a incana Leidy. A

neotype is designated for Pttpa incana Leidy, 1851, from the

subse(|uently designated type locality in order to proxide an

ol)jectix’e standard of reference for this species-group taxon

that may be used to ex'aluate the relationship of the nomino-

typica! subspecies to three siibsecpiently described subspecies,

as well as to two introduced Bahamian species with xvhich it is

documented to hax'e hybridized.

Additional Ket/tvords: Pupa incana. authorship, neotype,

Florida Keys

INTRODUCTION

Tlie family Cerionidae is well known for its remarkable

moiphological diversity and large number of tiixa, partic-

ularly in the recent faunas of Cuba and the Bahamas.

In contrast, this family is represented in the Florida Keys

by a single native species. The authorship of this species,

Cerion incamim (originally proposed as Pupa incana),

has universally been attributed to Binney, 1851 (e.g.,

Maynard, 1889; Pilsbiy', 1902; Bartsch, 1920; Abbott,

1989), despite recognition by several subsequent authors

of difficulties with the manner in which it was published

(e.g., W. G. Binney, 1859; 142; Clench, 1957: 148;

Pilsbry, 1946: 162).

A review of Amos Binneys posthumously published

work (Binney, 1851-57) in the context of the require-

ments of the International Code of Zoological Nomen-

clature (ICZN, 1999) reveals that his uses of the name

Pupa incana do not occur in a way that satisfies the

criteria of availability (Article 12) until the publication

of plate 68 (Binney, 1857), which associates images of

hx'o adult and one juvenile shell with the name Pupa

incana (Article 12.2.7, ICZN, 1999: 17). The first usage

of the name Pujta incana in conjunction with a descrip-

tion and illustrations was in a portion of the work written

bv Joseph Leidy (Leidy, 1851) reporting on the anatomy

of several terrestrial gastropods. Thus, the authorship of

this species must be attributed to Leidy, 1851. Leidy s

description pre-dates the publication of Pupa detrita

Pfeiffer, 1854, which is a junior synonym of Pupa incana

Leidv, 1851, but a senior synonym of Pupa incana

Binney, 1857.

HISTORICAL REVIEW

Amos Binney, M.D. (1803-1847), a physician, business-

man, and naturalist had an axfd interest in the terrestrial

Mollusca of North America, accumulated a sizeable col-

lection, and had undertaken the production of a major

publication on the subject. Despite his untimely death

(February 18, 1847), he had made provisions for the

completion and publication of this work in his will.

Augustus A. Gould was engaged by the executors of

Binney’s estate to edit and publish this work. Gould

(in Binney, 1851a: ix-x) reported that “The work was

found to be less complete than anticipated . . and

acknowledged that there are some discrepancies

between the first x'olume and the second, portions of

wliich were written by Gould (clearly identified by

‘annexation of the letter “G.”’)

The work, titled The Terrestial Air-Breathing Mollusks

of the United States, and the Adjacent Territories of

North America, was published in five volumes, the first

three attributed to A. Binney (1851-1857) although

M.G. Harasewych, 2014

Page 19

edited by A. A. Gould. Tlie first volume (1851) contains a

series of essays by Amos Binney on topics ranging from

the history of zoological investigations in America to the

zoogeography and ecolog)' of terrestrial Molliisca, and

also includes a section, authored l)y Josepli Leidy, on

the anatomy of the terrestrial gastropods of the United

States. Tlie systematic treatment of taxa begins in the

second volume (1851) and continues into the third vol-

ume (1957), which also conhiins the plates illustrating

the treated species. The fourth and fifth volumes were

autliored by W. G. Binney (1859-1878), the son of Amos

Binney, and are a continuation and expansion of the first

three volumes. W. G. Binney further continued his work

on this fauna, pii!)lishing three supplements to the fifth

volume (Binney, 1883; 1886; 1890).

Chronological appearance of the name Pupa

incana in The Terrestial Air-Breathing Mollusks

of the United States, and the Adjacent Territories

of North America

Volume 1

Pupa incana Binney, 1851a: 109 {nomen nudum).

The name appears in a “Table of Distribution of Spe-

cies in Several Geographic Regions,” listing Pupa

incana as one of the “Species noticed in the Atlantic

Region Only”.

Pupa incana Binney, 1851a: 119 {nomen nudum).

Pupa incana is listed as among the land mollusks

that are indigenous to Cuba but also occur in south-

ern Florida.

Pupa incana Binney, 1851a: 128 {nomen nudum). Pupa

incana is listed in a table as one of the species in

“Section 1.” Identified on page 108 as occurring on

“The Penimula of Florida”. The species name is not

italicized in the table, indicating that it is not “intro-

duced from foreign countries” (see p. 126).

Pupa incana Binney, 1851a: 153 {nomen nudum). Pupa

incana is listed as one of the species “undoubtedly

from Cuba”, transported on floating debris, that has

flourished in southern Florida.

Pupa incana Binney, 1851a: 162 {nomen nudum). Pupa

incana is listed in a “Table of Distribution of Species

Inhabiting the United States, in other parts of

America” indicating that it also occurs in Cuba.

Pupa incana Leidy, 1851: 199 {nomen nudum). Pupa

incana appears in a list of animals dissected.

Pupa incana Leidy, 1851: 216 {nomen nudum). The

name appears (“In Vaginulm, Pupa incana, etc.”)

in a description of the “tissues of the digestive appa-

ratus”. This is a generalized description that does

not distinguish Pupa incana from Vaginulm.

Pupa incana Leidy, 1851: 230-231. A detailed descrip-

tion of the genital apparatus of Pupa incana is pro-

vided. Tliis is the first association of tlie name

Puj)a incana witli a decscription (of the reproductive

system) that meets tlie reijuirements of Article 12.1

(ICZN, 1999:16), making tlie name Pupa incana

available as Pupa incana Leidy, 1851.

Pupa incana Leidy, 1851:259. Piipa incana appears in

captions to figures ii-iv of plate 15, as well as on the

plate itself. This plate includes three illustrations

of the dissected soft tissues (Article 12.2.7; ICZN,

1999:17), making the name Pupa incana availalile.

VOLU.ME 2

Gould in Binney, 1851b: 314. Uses the liinonien Pupa

incana in a discussion of supraspecific relatioii.ships

within Pupa. The annotation “[G.]” identifies this as

the work of Gould.

Gould in Binney, 1851b:316-319. This description was

no doubt intended by Binney to be the formal

description of Pupa incana. However, Gould replaced

the name with Pupa nuirifinui Pfeiffer. The following

parentlietical note, identified as being by Gould [-G.]

follows the description.

[This shell was descri})ed by Dr. Binney as a new species,

and is referred to in the first volume under the name,

P. incana; and the plate representing it is so lettered;

but it is so evidently the P. maritima of Dr. Pfeiffer

that it would be deviating from scientific rules to intro-

duce it under any other name. It is most likely that

P. cijclo.stoma, Kiister, is merely a small specimen of this

species, its variation in size being very considerable. -G.]

VOLU.VIE 3

Gould in Binney, 1857: 39. “Index to Plates” says “Pupa

maritima Pfeiffer” with a note that the plate itself is

lettered Pupa incana.

Binney, 1857; pi. 68. Plate illustrating two adult and

one juvenile specimen is labeled Pupa incana.

This makes the taxon Pupa incana Binney available

(Article 12.2.7; ICZN, 1999:17) as of 1857.

Volume 4

Binney, W.G., 1859: 142. W. G. Binney commented

“In 1851, this species was only designated by name

in vol. i; the full description was given in vol. ii, the

name being changed by Gould to P. maritima.

Shuttlewortli’s description of detrita has not prioritv

over the name I have retained [P. incana].”

Binney, W.G., 1859: 206, pi. 79, fig. 17. The sub-

species Pupa incana fasciata is introduced in the

“Explanation of the Plates” for a variet}' from Key

Biscayne, Florida.

Volume 5

Binney, W.G. 1878: 213. Included the note “Pupa incana,

= Strophia”

Page 20

THE NAUTILUS, Vol. 128, No. 1

Binney, W.C. 1878: 220-222, figs. 125,126. A synonymy

and description of the shell, jaw and anatomy of

Strophia inaina is provided.

TYPE MATERIAL

It is not clear if Leidy’s descriptions of several organ

systems of Pupa incana were based on one or multiple

dissected specimens. It is probable that the shell(s) were

cnished in order to access the soft tissues, and the ana-

tomical material was not saved following dissection. Tlius,

there is no siiniUng tyjie material for Leidy’s taxon.

Gratacap (1901) reported tliat the Binney and Bland

collection of North American land shells, which included

specimens used by Amos Binney and A. A. Gould in the

preparation of Terrestrial Air-Breathing Mollusks of the

United States, as well as specimens that W. G. Binney

incoqoorated from his collection and from tlie cabinet

of Thomas Bland, came to the American Museum of

Natural History in 1882. In the published catalog of this

collection, Gratacap (1901: 369) listed ten lots of

Strophia incana Cerion incamim = Pupa incana],

from Florida, Key Biscayne, and Key West. Among

these, he refers to two lots [Cat. Nos. 795, 796]

as “Type”, citing the pagination for the description

“(as nuiritinm Pfeiff.)” and referencing the plate pub-

lished in 1857, illustrating two adult specimens and a

juvenile over the name Pupa incana. A specimen in the

collections of the Museum of Comparative Zoology

(MCZ 149431) is also catalogued as a svntvpe of Pupa

incana Binney, 1851.

However, these specimens were not seen by Leidy. As

neither the description (as rnaritinia) nor the illustrations

of these shells were published until after Leidy s anatom-

ical descriptions, tliey cannot be considered as part of a

synU'i^e series of Pupa incana Leidy, 1851 (Article 73.2.1,

ICZN, 1999: 81). Thus, there are no name-bearing type

specimens of Pupa incana Leidy, 1851.

Early authors regarded the geographic range of Cerion

incanum to span Cuba, the Bahamas, and soiitliem

Florida, both the mtiinland and the Keys. Pilsbry (1902:

Figure.s 1-10. Pupa mcana Leidy, 1851. Neotyjre, USNM 1231355. Little Hamaca City Park, Government Road, Key West,

Florida, 24°33.410' N, 81°46.37()' W. In roadside leaf litter, under rock, at base of grass clumps. Febriiaiy 8, 2013. M.G. Harasewych

coll. 1-5, Shell. 1. Apertural, 2. Lateral, 3. Dorsal, 4. Apical, and 5. Basal Hews. 6-7. Scanning electron micrographs of protoconch.

6. Apical, and 7. Lateral Hews. Arrows indicate protocondi/teleoconch transition. 8-9. Dorsal views of jaw. 8. Photomicrograph.

9. Scanning electron micrograph. 10. Scanning electron micrograph of portion of radular ri!)bon. Entire radula consists of 135 rows,

each row with formula 24-1-24. r, racliidian tooth.

M.G. Ilarasewydi, 2014

Page 21

214) restricted the t\pe locality of Cerion incanmn to the

area around the salt ponds on Key West. Subse(|iient

workers furtlier subdivided tliis species into subspecies.

W.G. Binney (1859) proposed the viiriety Pupa incana

fasciata for a form from Key Biscayne, Florida. Pilsbry

and Vanatta (1899) added Cerion incamim saccharimeta

from Siigarloaf Key, and Pilsbr)' (1902) described Cerion

incanmn vaccinum from Key Vaca.

A neotype for Pupa incana Leidy, 1851, is designated

from a population [Little Hamaca City Park, Government

Road, Key West, Florida, 24°33.410' N, 8r46.370' W]

inhabiting the edges of salt ponds in Key West, Florida

that corresponds to the type kx-cility reported by Pilsbry.

This neotype (USNM 1231355) consists of 2.0 protoconch

whorls and 8.5 teleoeonch whorls (Figures 1-7), has a

shell length of 24.18 mm, a shell width of 9.47 mm, and a

maximum aperture length of 6.64 mm. It corresponds to

the Cerion incanum incanmn of authors, and generally

conforms to Leidys anatomical descriptions (although

Pilsbry 1946: 162, noted some inaccuracies in Leidys

description of the pallial organs) as well as to the numer-

ous detailed subsequent descriptions of tire anatomy and

shell iTior|ihology (e.g, W. G. Binney, 1878: 220; Bartsch,

1920: 7; Pilsbiy, 1902: 213; 1946: 160-165).

A neotype is necessary in order to provide an objective

standard of reference for this species group taxon that

may then be used to evaluate the relationships of the

nominotypical subspecies to three subsequently described

subspecies that are native to Florida. A precisely defined

concept ol Cerion incanmn incanum is of particular

importance in unravelling the relationships of tliis species

to several species that have been purposely introduced

into the Florida Keys from the Baliamas and elsewhere

during the early 20th century (see Bartsch, 1920; Pilsbry,

1946; Harasewych and Strauss, 2006), as some of these

have been documented as having hybritlized with Cerion

incamim (Woodruff and Gould, 1987).

In addition to the shell, radula, and jaw, genomic DNA

samples have been archived for this neotype. Results

of Next Generation sequencing of a subsample of this

DNA, including the reconstruction of a complete mito-

chondrial genome and the complete ribosomal operon,

will be published separately.

ACKNOWLEDGMENTS

I am grateful to Yolanda Villacampa for assistance with

tlie SEM imaging of the protoconch, radula, and jaw.

Richard E. Petit provided helpful comments on a draft

of this manuscript.

LITERATURE CITED

Abbott, R. T. 1989. Conipendiuin of Landsliells. American

Malacologists, Melbourne, 240 pp.

Bartsch, P. 1920. Experiments in the breeding of Cerions.

Papers of the Department of Marine Biology, Carnegie

Institution of Washington 14 (282): 1-54, pis. 1-59.

Binney, A. 1851-1857. [published po.stliimiously]. I’he 'lerre.stial

Air-Breathing Mollusks of the linited State.s, and the Adja-

cent Territories ol North America: Described and Illustrated

l)y Amos Binney. Edited by Agustiis A. Could. Charles Little

and James Brown, Boston, Volume 1 [ ]851a]:i-,xxix, 1-266

[last page inisnumbered as .366 (hpo)] pis. 1-16; Wlume 2

[1851b]: 1-362; Volumes [1857]:'l-4(). pis. 1-74.

Binney, W.G. 1859-1878. The Teirestial Air-Breathing .Mollusks

ol the United States, and the Adjacent Territories of North

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Terrestrial Air-Breathing Mollusks of the United States,

and tlie Adjacent Territories of North America. Bulletin

of the Museum of Comparative Zoology 11 (8): 135-166,

pis. 1-4.

Binney, W.G. , 1886. A Second Supplement to the Fifth Volume

of Tlie Terrestial Air-Breathing Mollusks of the United

States, and the Adjacent Territories of North America.

Bulletin of the Museiiin of Comparative Zoology' 13 (2):

23-48, pis. 1-3.

Binney, W.G., 1890. A Third Supplement to the Fifth Volume

of The Terrestial Air-Breathing Mollusks of the United

States, and the Adjacent Territories of North America.

Bulletin of the Museum of Comparative Zoology 19 (4):

183-226, pis. 1-1 1.

Clench, W.J. 1957. A catalog of the Cerionidae (Mollusca:

Pulmonata). Bulletin of the Museum of Comparative

Zoolo^ 116: 121-169.

Gratacap, L.P. 1901. Catalogue of the Binney and Bland Col-

lection of Terrestrial Air-Breathing Mollusks of the United

States and Territories in the American Museum of Natural

History, with enumeration of ty|)es and figured specimens,

and supplementary notes. Bulletin of the American

Museum of Natural History 14 (Article 23): 335-403.

Harasewych, M.G. and J. Strauss. 2006. A new record of intro-

duced Cerion (Gastropoda: Pulmonata: Cerionidae) in

southeastern Florida. The Nautilus 120: 94-100.

ICZN (International Commission on Zoological Nomenclature)

1999. Internationa] Code of Zoological Nomenclature,

Fourth Etlition. The International Trust for ZoologicN

Nomenclature, London, 306 pp.

Leidy, J. 1851. Special Anatomy of the Terrestrial Gastropoda

of the United States, pp. 198-260, pis. 1-16. [In] Binney,

A. 1851a. The Terrestrial Air-Breathing Mollusks of the

United States, and the Adjacent Territories of North

America; Described and Illustrated by Amos Binney.

Edited by Augustus A. Gould. Volume 1: 1-266, pis. 1-16.

Maynard, C.J. 1889. Monograph of the Genus Strophia.

[continued]. Contributions to Science 1(2): 68-79, pi. 7.

Pilsbry, H.A. 1902. Family Cerionidae. Manual of Conchology.

Series 2. Pulmonata. Academy of Natural Sciences of

Philadelphia 14(56): 193-286, pis. 27-47.

Pilsbry, H.A. 1946. Family IX.Cerionidae. Land Mollusca

of North America. Academy of Natural Sciences of

Philadelphia, Monographs 3, 2(1): 158-169.

Pilsbry, H.A. and E.G. Vanatta. 1899. Some Cuban species of

Cerion. Proceedings of the Academy of Natural Sciences

of Philadelphia [1898]: 475-478, figs. 1-9.

Woodruff, D.S. and S.J. Gould. 1987. Filtv years of inter-

specific hybridization: genetics and moqihometrics of a

controlled e.xperiment on the land snail Cerion in the

Florida Keys. Evolution 41: 1022-1045.

THE NAUTILUS 12S(l):22-27, 2014

Page 22

The freshwater mussel collection (Bivalvia: Unionida) of the

Miiseo Nacional de Ciencias Naturales (Madrid, Spain)

Maria Dolores Bragado

Rafael Araujo’

Mu.seo Nacional de Ciencias Naturales, CSIC

Jose Cntierrez Abascal 2

28006 Madrid, .SPAIN

[email protected],s

[email protected]

Javier de Andres

Mnseo Nacional de Ciencias Naturales, CiSlCi

Jose (intierrez Abascal 2

28006 Madrid, SPAIN

[email protected],sic.es

Arthur E. Bogan

North Carolina State Musenin of Natural Sciences

1 1 West Jones Street

Raleigh. NC 27601 USA

[email protected]

ABSTRACT

A review of the freshwater mussel collection housed at the

Mnseo Nacional de Ciencias Naturales of Madrid (Spain)

(MNCN) including complete specimen data for the 3,270 lots

is provided. The 10,009 specimens (single valves not included)

represent species from five continents and Oceania, with

342 species in 110 genera belonging to 6 currently recog-

nized families of the Order Unionida. This collection contains

42.9 % and 60.7 % of the total currentlv recognized unionoid

species and genera respectively. There are 152 lots with 30 spe-

cies from Africa, 1,359 lots containing 245 species from the

Americas, 125 lots including 55 species from Asia, 1,472 lots

representing 18 European species and 28 lots of 7 species from

Oceania. The Iberian Peninsula has the greatest representation

with 1,233 lots (114 Margaiitiferidae and 1,119 Unionidae).

'fhe alcohol-preserved collection includes 456 lots of European

ami 39 lots of North African mussels, and represents a sig-

nificant contemporary collection of freshwater mussels from

these areas. We also present an historic description of the

collecdon, with special emphasis on the South and Central

American mussels collected by the Comision Cientlfica del

Pacifico (1862-1866), aTid updated data on the presence

of threatened and endangered species following the Endan-

gered Species .Act (USA) and the Habitats Directive (Europe).

According to the IntemadoTuil Union for the Conservation of

Nature [lUCN] Red Ust, there are 30 % of the Vulnerable

species, 36 % of the EndaTigered species, 44,5 % of the Crid-

cally Etidangered and tlie .'34.5 % of todays E.xtinct freshwater

mussels represented.

' Corresponding author

INTRODUCTION

Biological specimen collections, conserved in natural

history museums and other institutions, provide a rich

source of data essential for numerous areas of research

including biogeographv to phenological or evolutionary

studies. These collections comprise samples collected

during the period of accelerated anthropogenic habitat

destruction, climate warming and freshw'ater habitat

destruction. These specimens and their associated data,

document baseline conditions before the major impact

of these factors (Nicholson, 1991; Allmon, 1994; Shaffer,

Fislier and Davidson, 199S; Lister et ah, 2011; Suarez

and Tsutsui, 2004). Museum collections are essential for

the future documentation of the abundance and dis-

tribution of species (Lips, 201 1 ). In order to achieve this

goal, it is necessaiy to curate existing specimens and tlieir

corresponding data in museums and make sure this data

is well organized and available for scientists (Solem at ah,

19S1) and others on the World Wide Web.

The objective of this paper is to update the data

associated watli each lot (taxonomic, faunistic, storage,

number of specimens, etc.) in the fre.shwater mussel

collection (order Unionida) (Bouchet and Rocroi, 2010;

Carter et ah, 2011) of the Mnseo Nacional de Ciencias

Naturales (Madrid, Spain) (MNCN), providing a com-

prehensive overview of this material, as well as docu-

menting the association of classic studies on taxonomy

and tlistribution with the specimens. Drv specimens mav

be used for studies on shell ultrastructure, trace ele-

ment analyses, sclerochronologv or climate change,

while alcohol-preserved specimens can be the source

of anatomical and molecular markers used in systematic,

phylogenetic or ev'olutionarv research. We present an

M.D. Bragadoet al., 2014

Page 23

historical description of this collection, with special

emphasis on the freshwater nmssels collected by the

“Comision Cientifica del Pacifico”, a natural histor\'

expedition made between 1862 and 1866, and provide

updated data on the presence of Threatened, Endan-

gered and Extinct species following the lUCN Red

List, the United States Endangered Species Act and

the Habitats Directive.

The mollusk collection of the MNCiN houses

aroimd 10(),()()() lots of the eight known mollnsk

classes: Solenogastra, Caudofoveata, Polyplacophora,

Monoplacophora, Gastropoda, Cephalopoda, Bivalvia,

and Scaphopoda. Although specimens have been col-

lected from around the world, the best represented

areas are the Iberian Peninsula, Philippines, Cuba,

E(juatorial Guinea, and South America. The terres-

trial mollusk collections from the Philippines, Cuba

and Pajuatorial Guinea are probably among the best

in the world. There is an important historical repre-

sentation of the South and Central American fauna

coming from the Comision Cientifica del Pacifico.

The core historical material of the collection is

from three Spanish malacologists: Paz (P. M. Paz y

Membiela, 1808-1874) wath 40,000 specimens represent-

ing 12,000 species anrl subspecies, Hidalgo (J. Gonzalez

Hidalgo, 1839-1923) with 8,000 species, and Azpeitia

(E. Azpeitia, 1859-1934) with 80,000 specimens of

8,171 species (6,594 gastropods and 1,577 bivalves)

(Barreiro, 1992). Other historical material comes

from the founder of the Museum, P. Eranco Davila

(1711-1786), and from M. Graells (1809-1898), who

w'as director of the museum betw'een 1845 and 1867.

More recently, the collections have growm with the

additions of A. Ortiz de Zarate (1887-1964) with

4,500 lots, A. Cohos (1922-1998) with 4,390 lots, and

J. Conde with approximately 20,000 lots. The type

collection incliuling primary and secondary types, con-

tains 1,192 lots corresponding to 946 taxa, all are com-

puterized, arranged in tiixonomic order, and housed

in all steel cabinets with metal drawers that meet con-

servation standards and are locked (Templado et ah,

1993; Villena et ah, 1997). Currently, only 50% of the

general collection is computerized.

The Comision Cientifica del Pacifico (the Comision)

collected 38,755 specimens belonging to 816 different

species (Almagro, 1866). Most of them were collected by

the President of the Comision, (P. M. Paz y Membiela),

and by E P. Martinez y Saez, but other members of tlie

expedition like Jimenez de la Espada, Isem and Almagro

also collected mollusks. Zameron, Barreiros, Philippi

and Richardson donated 767 specimens of 43 species,

and 37 specimens of 19 species were purchased (Almagro,

1866). All of the Comision specimens were studied by

Martinez y Saez, who was responsible for mollusks

during the expedition, and by j. Gonzalez Hidalgo

(Hidalgo, 1900; Galvo, 1994). Martinez y Saez and

Hidalgo wrote the three volumes set Mohiscos del viaje

al Pacifico, which included marine bivalves (Martinez,

1869), terrestrial gastropods (Hidalgo, 1872) and marine

gastropods (Hidalgo, 1879). The freslnvater bivalves

were studied bv Isaac L('a (1792-1886) (Lea, 1866a, b,

1867, 1869a, li) and Eritz Haas (Haas, 1886-1969), w'ho

pulilished "Ndtjades del viaje al Pacifico" (Haas, 1916).

Of the 60 new' mollusk spi'cies described from th('

material collected by the Comision, 18 w'(>re freslnvater

liivalves: 2 were described bv Hidalgo (CaK'o, 1994),

14 by Lea (1866a, b, 1867, i869a, b) and 2 bv Haas

(1916). Tlie 14 species describc'd by Lea (1866a, b,

1867, 1869a, b) lacked exact locality data, reporting

only South or Central America, but w'ere proliablv

collected bv Paz in the Uruguav River (Salto Oriental,

Uruguay) and were sent to Lea at the end of the (‘xpe-

dition (Haas, 1916). It was uncertain if Lea’s tvpes

were included in the MNCN or in the Lea collection

at tlie USNM (Smithsonian Institution, Washington, DC)

(Haas, 1916).

RESULTS

Detailed locality, taxonomic and historical on all the

freshw'ater mussel (Unionida) lots in the MKCN collec-

tion can be found at the Global Biodive rsitv Information

Eacility (GBIE). There are 1(),()()9 specimens (single

valves not included) in 3,270 lots representing 342 spe-

cies of 110 genera belonging to the currently recognized

6 families: Margaritiferidae, Uniouidae, Mycetopodidae

(= Mulleriidae), Etheriidae, Iridinidae and Ilyriidae

(Bouchet and Rocroi, 2010; Carter et ah, 2011). The

collection contains representative lots and specimens

of 60.7 % and 42.9 % of the total nnmber of described

unionoid genera anti species respectively (Graf and

Cummings, 2007; Bogan, 2008) (Table 1). There are

587 alcohol-preserved lots and 2,683 diy lots. Among

these specimens are glochidia, optical microscopical

slides of soft parts and shell thin sections.

Summarizing the geographical information on speci-

mens in the unionoid collection (Table 2), there are

134 lots without locality information, including the

continent of origin. The area represented by the most

specimens is Europe, with 18 species (including the

exotic Siruinodonia ivoodiana (Lea, 1834)) in 1,472 lots,

and 29 lots without country identification. The largest

number of lots comes from the Iberian Peninsula, w'ith

1,233 lots (114 Margaritiferidae and 1,119 Uniouidae):

1,131 from Spain and 102 from Portugal. During the

past 12 years (Soriano et ah, 2001), the Iberian material

in the family Uniouidae in the collection has grow'ii b\’

754 lots. Other European lots by countiv are: Albania

(3), Austria (7), Belgium (11), England (14), Croatia (6),

Czech Republic (l), Denmark (1), Estonia (2), Erance

(69), Einland (1), Germany (37), Greece (32), Hungaiv-

(7), Ireland (5), Italy (58), Latrta (2), Netherlands (1),

Poland (11), Romania (2), Russia (12), Slovakia (10),

Slovenia (4) and Sw'itzerland (5). Of the 152 African

lots, 42 come from Morocco, 19 from Egypt, 18 from

Senegal and 18 from Tunisia; tlie rest come from Algeria

(1), Burkina Easo (3), Buiundi (7), Dem(x.'ratic Republic

Page 24

THE NAUTILUS, Vol. 128, No. 1

I'ablc 1 . 4 otal numher of genera anti .species in families of fresliwater mussels in the MNCN collection in comparison with the

whole described taxa. * W'e consider all the li\ing Margaritiferidae belonging to the genus Mnroaritifera. (1) Graf & Cummings

(2007). (2) Bogan (2008).

of tlie Congo (6), Etliiopia (1), Gabon (1), Guinea (1),

Ivoix,' Coast (2), Lil)ya (2), Mauritania (1), Niger (1),

Nigeria (1) and Soutli Africa (6). Tlie alcoliol-preserv'ed

collection, witli 456 European and 39 lots of North

African naiads is one of the most significant conteinpo-

ran’ collections of freshwater inns.sels for these areas.

There are 1,358 American lots, which include 973 from

the United States of America, and 53 lots from Ctiha,

44 collected hv Poey already included in the Piiz v

Memhiela collection (Barreiro, 1992), and tlie aforemen-

tioned specimens collected hv the Comision Cientffica al

Pacffico (.see below). The nnmher of lots by countiw

is: Argentina (14), Bolivia (4), Brazil (60), Canada (2),

Chile (17), Colombia (2), Ecuador (18), Guatemala

(1), Me.xico (10), Nicaragua (S), Paraguay (1), Peni (3),

Uniguay (103) and \Tnezuela (1). There are 78 North

American lots wdthout a countn’ listed. The best repre-

sented Asiatic countries, with a total of 125 lots, are

Japan with 33 and Ghina with 21. Other A.siatic lots

by counti'v are: Cambodia (1), Philippines (8), India (15),

Indonesia (5), Irac] (6), Israel (2), Laos (3), Malaysia

(1), Myanmar (2), Singapore (2), Sri Lanka (2), Taiwan

(1), Tliailand (7), Turkey (9) and \ietnam (7). There

are 28 lots from Oceania, 8 from Australia and 20 from

New Zealand.

IIISTOBICAL OVER\4EW

Among the freslnvater mussel historical material, it is

important to document the presence of mid-nineteenth

centniT specimens collected in South America hv

the Comision Cientffica al Pacffico (1862-1866), which

included tlie Parana and Amazon basins, Chile and

the Pacific coast of Ecuador. The information about

tliese specimens comes from tliree different sources:

(1) Almagro (1866, page 162) published the list of the

freshwater mussels collected in the Comision, detailing

tlie localities (15), number of species (44), number of

specimens (300) and the name of the collectors. (This

information w'as based on specimens exhibited after the

Comision Expedition); (2) Lea (1866a, b; 1867, 1869a,

b) described 14 new' species from South America, pro-

cured by Paz during the Comision but failed to include

any reference as to the origin of the specimens. Fol-

low'ing Lea (1866a), some of the material sent by Paz

was preserv'ed in alcohol; (3) Haas (1916), who, during

his obliged stay in Spain due to the unfavorable politi-

cal climate in Germany (Haas, 1915), w'as invited to

the MNCN where he studied the mussels collected

during the Comision. Haas (1916) listed 32 different

ta.xa from the Comision collection, two he described as

new' species (Diplodon hidalgoi and Mijcetopoda holivari).

He added a summarx’ organizing the species by river basin

and included some biogeographic remarks and observa-

tions. Most of this material is in the MNCN collection

and has the original Haas label (Fig. 1). In his paper,

Haas mentioned that some specimens were sent by Paz

to Lea, who published several new species based on this

material (Lea, lS66a, b, 1867, 1869a, b) and Lea did not

return part of it. This included four types of species

Lea described, and Haas supposed they w'ere in the

Lea collection at the United States National Museum,

Washington, DC (Haas, 1916).

Table 2. Number of lots in families of freshwater mussels at the MNCN mollusc collection. In brackets the corresponding

numher of species.

Ml). Bragado et al., 2014

Page 25

Figure 1. Handwriting of P’ritz Haa.s on the labels of the

Coinision specimens.

We located 51 lot.s of mussels collected by the

(loniision (Table 3. Freshwater mussels from the

Comision Cienti'fica al Pacifico in the MNCN mollusk

collection. Posted online at http://nautilus.shellinuseum

.org); they include 223 specimens ol 22 species belong-

ing to 2 different families. We verified that the specimens

sent to Lea by Piiz were indeed absent; at least those

corresponding to the following seven species described by

Lea (1866a, 1869) are not in tlie MNCN. We have found

them at the mollusk collection of the Smitlisonian Institu-

tion, National Museum of Natural History (Washington,

USA): Unio ampidlaceus Lea, 1866. Tvpe: USNM 85614

(=Diplodou wijmanii (Lea, I860)), Unio Jocelhis Lea,

1866. Type: USNM 85189 {=Rlupidodonta variahilis

(Maton, 1811)), Unio parens Lea, 1866. Ty|3e: USNM

85198 {=R]iij)idodorita cJiarniana (Orbigny, 1835)), Unio

aaitirostris Lea, 1866. Type: USNM 85930 {—Diplodon

parallelopipedon (Lea, 1834)), Unio appriinus Lea, 1866.

Ty|:)e: USNM 85167 {=Diplodon wtpmmii (Lea, I860)),

Unio mgososidcaius Lea, 1866. Type: USNM 84392

{=P(ichijnaias mgososidcata (Lea, 1866)), and Unio

nifofuscns Lea, 1865. Type: USNM 84101 (—RJiipidodonta

grata (Lea, I860)). Another part of the Paz material

is separated between the Smithsonian and the MNCN

(Table 3. Freshwater mussels from the Comision

Cientifica al Pacifico in the MNCN mollusk collec-

tion, posted online at http://nautilus.shellmuseum.org):

Unio paraguaijensis Lea, 1866. Ty|ie: USNM 85068

i=Diplodon delodontiis (Lamarck, 1819)), Unio peadlaris

Lea, 1866. Type: USNM 85190 (= Diplodon wijuuinii

(Lea, I860)), "Unio finnus I^ea, 1866. Type: USNM 85113

(= Diplodon delodontiis (Lamarck, 1819)), Monocondijlaea

lentifomiis Leti, 1866. Type: USNM 86335 {—Monocondijlaea

comentesensis (Orbigny, 1835)), Monocondijlaea pazii

Lea, 1866. Ty|3e: USNM 86342 {—Monocondijlaea

niinuana (Orbigny, 1835)) and Anodorita pazii Lea,

1866. Type: USNM 86703 {—Anodontites trapezea (Spix

and Wagner, 1827)). HcUis (1916), cited Simpson (1914),

noting the tyjie of Anodorita napoensis Lea, 1868 was

also sent by Paz to Lea and it was present in the Lea

collections in the Smithsonian Institution, Wa.shington,

DC. This was an error by Haas (1916) because Simpson

(1914) did not cite P;i/. as donor but the name of Prof

Orton; therefore, the type' spe'cimen of this species at

tlie Smithsonian (USNM 25429) did not come from P;iz

l)ut from Dr. C. Strebc'l (Craf and Cummings, 2013).

Otlier material from the Comision can be' feMinel at the

Senckenberg Museum e)f Frankfurt (Haas, 1916; Zilch,

1967; Graf anel Cummings, 2013).

The hvo species described by Hielalge), Cast alia

crosseana Hielalge), 1865 anel Cast alia jrazi Hielalge),

1868, luwe been sy'ne)nvmizee! uneler Cast alia ainhigua

Lamarck, 1819 anel Rhipidodonta hi/laea (Orbignv,

1835) by Be)nnette) (1965) anel Siine)ne (2006), rexspec-

tively. One specimen of each was sent by Hielalge) te) the

Senckenberg Museum (Frankfurt am Main, Germany)

(Haas, 1916). SyTie)nymized by Haa.s (1969) are the tvve)

species described by himself (Hais, 1916), Diplodon

hidalgoi and Mijcetopoda bolivari, as Rhipidodonta

charniana (Orbigny, 1835) anel Mijcetopodella falcata

(Higgins, 1868).

It is worth ne)ting the presence of senne specimens

from the Comisiehi preserved in fluiels (currently in

70% ethane)!) at the MNCN ce)llectie)n that were

probably e)verloe)keel by Haas (1916): 7 specimens of

Anodontites trapesialis (Lamarck, 1819), 6 specimens

of Anodontites trapezea (Spix and Wagner, 1827) anel

2 specimens e)f Diplodon delodontiis (Lamarck, 1819).

The remaining 208 specimens are all elry.

\¥e ne)te some differences in the number e)f speci-

mens recordeel by Almagro (1866), Haas (1916) anel this

paper. Fe)r instance, regarding Anodontites patagonica

(Lamarck, 1819), Haas (1916) cite^d two le)ts at the

MNCN, one lot e)f 3 specimens from Rie) ele la Plata

and anotlier le)t with 2 specimens fre)m Miguelete River,

and we have fe)unel only one lot, labelleel as Ri'o de la

Plata, but with 6 specimens, four of them with “Rio de

la Plata” written insiele the shell. This may!)e e.xplained

because se)me material may have been lost or mixed

during the last 150 years, or because the data from

Almagro (1866) might not be accurate. Almagro ’s book

yvas wTitten in the last year of the Comision, so the

historical number of species and corresponding number

of specimens may be ine,xact. Another reason could be

the loaning or gifting of specimens between different

collections and/or the loss of old labels, as occurred

among the Paz, Hidalgo, and Azpeitia collections. We

have not found the specimens from Montevideo, Negro

Riy^er, Aguano and (3tavalo (Almagro, 1866).

Some years after the end of the Comision, Paz sent

Ix'a his collection of North American freshyvater mollusks

for expert identification (ACN()252/()()2; ACN()259/()13).

Lea returned the identified specimens (451 gastropods

and 287 bivalves) in 6 parcels to La Habana (Cuba),

yvhere thev yvere shipped again to Cadiz (Spain) arriving

on 12 July 1872. On 22 August 1872 they yvere moved

by train to Madrid yvhere they arrived oti 4 September.

One year later, in 1873, the Paz mollusk collection.

Page 26 THE NAUTILUS, Vol. 128, No. 1

O ’

Table 4. Niiinlier ol freshwater iiiussels species included in the lUCN in the MNCN collection. Number total of species included

in each categors' in brackets.

including this material, was sold to tlie Museo Nacional

de Ciendas Natiirales (Barreiro, 1992).

ENDANGERED SPECIES

Freshwater bix'alves are among the most imperiled

animal groups on tlie planet, declining due to pollution,

modified or disturbed habitat and reduced numbers

of host fish (Lydeard et ah, 2004; Strayer et ah, 2004).

In cases of imperiled fauna, the role of natural histoiw

collections is essential, being the last repositories where

scientists can study extinct or endangered species

(Nicliolson, 1991; Shaffer, Fisher and Davidson, 1998).

The MNCN collection has 82 % of all the Near Threat-

ened freshwater bivalve species, 30 % of the Vulnerable

species, 36 % of the Endangered species, 44.5 % of the

Critically Endangered and 34.5 % of the today’s Extinct

naiads recognized by the International Union for the

Consemition of Nature (lUCN) (Table 4). It also lias

nearly 60 % of the North American Endangered species

(Endangered Species Act) and numerous lots of all

the European species protected under the Habitats

Directive: M arrant if era margantifera, M. auricularia,

U. elongatiihis, and U. crassus (Tables 5, 6; Table 6.

Freshwater l)ivalve species in the MNCN collection

included in tlie lUCN, the Endangered Species Act and

the Habitats Directive. Posted online at http://nautilus

., shellmuseiim.org).

Margaritiferidae represents one of the most imperiled

naiad families. The MNCN mollusk collection contains

Table 5. Number of fre.shwater inus,sels species included

in the Endangered Species Act and in the Habitats Directive

in the MNCN collection. Number total of species included

in each category in lirackets. * We consider Unio mancus

and U. ravoisieri under the same Annex (III and IV) as

U. elongatuluH, and U. tuniidiformis under tlie same Annex (II

and IV) as U. crassus (Araujo et ah, 2009).

169 specimens of M. auricularia in 68 lots, 67 from Spain

and 1 from France, probably the largest collection of

this species in the world (Araujo and Ramos, 2000),

69 lots of M. niargaritifeni, 3 lots of the North African

M. marocana, 3 of M. inonodonfa, 1 of M. dahurica,

2 of M. laevis ant! 1 lot of the Asiatic M. laosensis.

Margaritifera laosensis is (]iiite rare in collections and

has only recently been collected alive in the northern

region of the Lao People’s Democratic Republic

(S. Schneider, Personal Communication).

AC K N OWLE DC M E NTS

C. Bogan, J. Smith, H. Leslie, A. Garcia- Valdecasas,

and K.S. Cummings reviewed earlier drafts of the

manuscript.

LITERATURE CITED

ACN()252/()()2. 1872. Archivo del Museo Nacional de Cieiicias

Natiirales, 16 pp.

ACN()259/013. 1872-187.3. Archivo del Museo Nacional de

Ciencias Natiirales. E.xpediente relative a la confusion

surgida en el envio de Rafael Arango al Museo de Ciencias

de una coleccion de conchas y caracoles, redamada por

Patricio Mana Paz como suya, y posterior coinpra de la

inisma para dicho establecimiento, 66 pp.

/Vllmon, W.D. 1994. The value of natural history collections.

Curator 37: 83-89.

.Almagro, M. 1866. Breve descripcion de los viajes hechos en

America por la Comision Cientifica enviada por el

Gobieriio de S. M. C. Durante los anos 1862 a 1866.

Mairena de Aljarafe, Sevilla, 174 pp.

Araujo, R., J. Reis, A. Machordom, C. Toledo, M.J. Madeira, 1.

Gomez, J.C. Velasco, J. Morales, J.M. Barea, P. Ondina,

and i. Ayala. 2009. Las nayades de la peninsula Iberica.

Iberus 27: 7-72.

Araujo, R. and M.A. Ramos. 2000. A critical revision of the

historical distribution of the endangered Margaritifera

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lie Jiu^ and Zhiiang Zimin. 2013. The Freshwater

Bivalve.s of China. Conch Books. Bahnhofstr. 117 D-

55296 Ilar.xJieiin, Germany.198 pages. 68 €.

This colorful volume is the most complete, well-illustrated

book to cover the freshwater bivalve fauna of China to

(late. The authors point out the early contributions/

foundation to our understanding of the Unionidae and

Corbicula species of China by Pierre Marie Heude

(1836-1902) illustrated by outstanding lithographs. His

ten-part series wus entitled CA)nchijliolo^ie flnviatile de la

pnmnce de Nankin^ et de la Chine central (1875-1885).

There liave been numerous journal articles covering new

species <uid notes on die liivalve fauna of particular Chinese

rivers but no compilation of the fauna. Liu Yue Ying (1979)

published the Economic Fauna Sinica - Freshwater Mol-

lu.sks, tlie first modem volume on freshwater mollusks

but it was illustrated with only line drawings and was not

nearly as detailed or in depth as the current vohime. He

Jing and Zhuang Zimin’s book treats all of the 126 recog-

nized freshwater bivalve taxa in seven families and lists

over 1000 smonyins. Five new species of Unionidae are

described by the authors. Liu (1979) listed a single species

of Spliaeriidae from China, while this work expands the

known fauna to include nine species. The Freshwater

Bivalves of China is the first modern treatment to s\ii-

thesize available information on this fauna with color pic-

tures and original lithographs where a\’ailable.

The front endpapers provide a physiographic map of

China with major cites marked. The volume begins with

an explanatory/ preface, acknowledgments, museum acro-

nvins and a regional map of Cliina with abbreviations used

for major faunal provinces. Species accounts begin with

Unionidae. Generic accounts are arranged alphabeti-

cally with all species in that genus listed alphabetically.

This is followed by the Margaritiferidae, Pharidae,

Mytilidae, Dreissenidae, Cyrenidae, and Spliaeriidae.

Generic accounts are ty|3ically short descriptions of diag-

nostic shell characters. Species accounts are usually

restricted to a single page, but may extend to three pages

and contain svnonvmy, distribution within China, types, a

shell description and figures. If original lithographs were

provaded with tlie description of the species, these are

reproduced, especially those species described by Isaac

Lea and Pierre Marie Heude.

It is great to see numerous color photographs of the

species where specimens were available and the repro-

duction of original lithographs of Heude, because his

work is not readily available. Pictures illustrating shell

shape variation and sculpture are veiw helpful.

Synonymies for each species appear to be complete and

wall be very useful in sorting out historic collections and aid

in developing future research problems. A synonymy is

very necessary w'hen dealing with the eastern Russian

named taxa and where diose taxa might belong. The distri-

bution table at the end of tlie species accounts lists species

by province and is very useful for distribution of a species

and a reference for the local fauna. I especially like the

complete and full citation style used in Bibliography. There

are no obscure abbreviations to provide puzzles on where

to find an article of interest. A list of the valid mussel

species is provided and has tire three introduced species

highlighted and a column on the right witli tire Cliinese

names. The most important part of any book is the index.

This index includes family and generic names, all species

names followed by the generic name under which it

appears in the text [valid names are in bold]. Synonymous

names are listed as well as tliose taxa recognized as valid.

As a reviewer, it is always easy to make a list of items

one would like to have seen included in the book, but you

are not the author of the book nor its editor. However, I

w'ould suggest the following changes for the authors to

consider for tire next edition of this fine volume. It would

help to have the photographic view of each valve stan-

dardized, outside of the right valve and the inside of the

left valve at least. A few of the color pictures are not as

shaqr as they might be, but all are very helpful. Type

section lists recognized specimens that are flagged as var-

ious tvpes. The tvpe status of each lot was not discussed

and is listed as it appears on lalrel without research. The

museum and literature research recpiired to determine

the correct status of a tvpe and who is responsible for the

correct tvpe determination is beyond the scope of this

book. I would caution anyone using this book not take

the tvpe status as resolved. Determining the ty|3e status

for each species will require a lot of work and time.

Remarks on each species are limited. In the next edition,

a more detailed comparison of similar species and how to

tell them apart would be useful. Detailed distributional

information for each species would be very helpful, but

this information may not be readily available at this time.

This volume should be on the bookshelf of all fresh-

w/ater biviilve biologists. I would recommend this volume

to anyone interested in freshwater bivalves, first of China

and secondly to anyone interested in the zoogeography of

freshwater bivalves in Asia. The pictures clearly illustrate

the issue of con\/ergence in shell shape and sculpture

between Asia and the rest of the world. But, it is also

just fun to page through the book looking at the pictures.

Hopefully, tliis volume will spark increased interest,

research, and conservation efforts on the freshwater

bivalves of China.

Arthur E. Bogan

Aquatic Invertebrates

North Carolina State Museum of Natural Sciences

11 West Jones St. Raleigh, NC 27601 USA

[email protected]

THE 2014 R. T ABBOTT VISITING CURATORSHIP

The Bailey-Matthews Shell Museum is pleased to invite applications for the 2014 R.T. Abbott Visiting Curatorship.

The Curatorship, established originally in accordance with tlie wishes of the late Dr. R. Tucker Abbott, Founding Director of the

Shell Museum, is awarded annually to enable malacologists to visit the museum for a period of one week. Abbott Fellows are

expected, by performing collection-based research, to assist with the curation of portions of the Museum’s collection and to provide

one talk for the general pul)lic. Tlie Museum collection consists of marine, freshwater, and terrestrial specimens. A large percentage

of our holdings have been catalogued through a computerized dataliase management .system; part of the catalogue is already

available for searches online at: www.shellmuseum.org/collection.cfm. The R.T. Abbott Visiting Curatorship is accompanied l)y a

stipend of $1,500.

Interested malacologists are invited to send a copy of their curriciiium vitae, a letter detailing their areas of taxonomic expertise and

research objectives, and to provide a tentative subject for their talk. Send materials to:

Dr. Jose H. Leal, Curator & Director of Education

The Bailey-Matthews Shell Museum

P.O. Box 1580

Sanibel, FL 33957

[email protected]

Applications for the 2014 Visiting Curatorship should be sent electronically to the alrove e-mail address no later than May 31, 2014,

or postmarked by that date if sent by regular mail. The award will be announced by late June 2014. Questions about the Visiting

Curatorship should be sent to the e-mail address above, or by phone at:

(239) 395-2233; fax (239) 395-6706

Sponsored in part by the State of

Florida, Department of State,

Division of Cultural Affairs and the

Florida Council on Arts and Culture

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