Volume 133, Number 1

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TH E

CONTENTS

NAUTILUS

Volume 133, Number 1

April 30, 2019

ISSN 0028-1344

Mauricio Romulo Fernandes

Juan Francisco Araya

Kristine Greke

Kazutaka Amano

Tomoki Kase

New species and records of Triphoroidea (Gastropoda) from Chile l

New species and record of Diplommatina Benson, 1849

(Gastropoda: Diplommatinidae) from Java, Indonesia ..........00 0 14

A Paleocene species of Procardia (Bivalvia: Anomalodesmata:

Parilimyidae) from the Katsuhira Formation in Urahoro Town,

easter IEIOeattlo, [EGE ..-cooocesosoccsaeeeces030;03008065a0793esona520600094909009090309r0555G8C8T0500000000 DY)

A large new Wareniconcha (Bivalvia: Vesicomyidae) from a

Shinji Isaji Pliocene methane seep deposit in Leyte, Philippines wie agai acted aasae ino ance eer oer 26

Yolanda M. Aguilar

Steffen Kiel

INOS HYBD, noconnscshasnnosadabedeoacdesoodkenscégoouseeesautlesoucbasdpenescbsad sognost Ede ebne eeeas sooon de econSbaddecouncce cGnDECe caopSeaEB EE SSb cud co adeacr aapececaasenoccrao nao aaceet a: 30

Sponsored in part by the State of

Florida, Department of State,

Division of Cultural Affairs and the

Florida Council on Arts and Culture

FLORIDA DEPARTMENT of STATE

DIVISION of CULTURAL AFFAIRS

THE NAUTILUS 133(1):1-13, 2019

age

Page |

New species and records of Triphoroidea (Gastropoda)

from Chile

Mauricio Romulo Fernandes

Departamento de Zoologia

Universidade Federal do Estado do Rio de Janeiro

Rio de Janeiro, BRAZIL

Juan Francisco Araya

Centro de Investigaciones Costeras de la Universidad de

Atacama (CIC-UDA)

Universidad de Atacama

and Copiap6, Chile

Departamento de Invertebrados and

Museu Nacional

Rio de Janeiro, BRAZIL

[email protected]

Programa de Doctorado en Sistematica y Biodiversidad

Departamento de Zoologia, Facultad de Ciencias Naturales y

Oceanograficas

U averse de Concepcion

Concepcion, Chile

Oi eee.

ABSTRACT

The present study aims to describe specimens of Triphoroidea

recently sampled from Chilean waters, with two species from the

northem continental shelf and one from the Juan Fernandez

Archipelago. Cerithiopsis eiseni Strong and Hertlein, 1939, originally

dlexeativadl from western Panama, has its protoconch, operculum and

radula illustrated for the first time. Monophorus monocelha new

species is recognized by having a very thick shell; yellowish pro-

toconch and cower watcocondie median spiral cord emerging be-

tween the sixth/seventh whorl of teleoconch; a brown mark a tween

the eyes; and radular formula 4-1-1-I4. Inella basalis (Odhner,

1922) new combination is recorded almost a century after its

original description, including data on intraspecific variation

in adult size, shape of protoconch, number of axial ribs on

teleoconch, and number of basal and supranumerical cords.

Cerithiopsis eiseni and Monophorus monocelha new species

share very thick shells and narrow apertures, suggesting a

converge nt defensive mechanism against predation.

Additional Keywords: Triphoridae, Cerithiopsidae, taxonomy,

marine gastropods

INTRODUCTION

The superfamily Triphoroidea has species with high-spired

and comparatively small shells, with the adult she I usually

being less than 10 mm long (Wells, 1998). Their spe cialized

feeding mode on sponges produced an amazing diversity of

castnlbne morphologies, especially in the left-coiled triphorids

(Marshall, 1983). Three families of Triphoroidea are cur-

rently recognized; Newtoniellidae is often prese nt in the

deep sea, valitile Triphoridae and Cerithiopsidae are usually

found in shallow and tropical/temperate waters (Fe smmandes

and Pimenta, 2017a). Triphoridae and Cerithiopsidae belong

to the group of the “Big Five”, representing the five most

species-rich families of marine gastropods in the world

(Bouchet et al., 2002; Albano et al., 2011).

Most taxonomic works of Triphoroidea from the

eastern Pacific were published from mid-19th to mid-20th

centuries (e.g., Adams, 1852; Bartsch, 1907, 1911; Baker,

1926: Baker et al., 1938), with the most recent ones dating

back to almost four decades (e.g., DuShane and Draper,

1975). The majority of these species was de scribed from

the northeastern Pacific, with few studies covering the

Southern Hemisphere. Species were described almost

exclusively based on shells, with spare exceptions in-

cluding a drawing of the head-foot in one species

(DuShane and Draper, 1975), and the investigation of the

reproductive system of three species (Houston, 1985).

The knowlec Ige about other important structures (such as

radula and operculum) for the taxonomy of this super-

family in the eastern Pacific is lacking.

In Chilean waters, three species of Cerithiopsidae and

11 of Triphoridae were recorded from Easter Island

(Odhner, 1922: Rehder, 1980; Raines, 2002), in addition

to one species of each family from the Juan Fernandez

Archipelago (Odhner, 1922). With the exception of

Cerithiopsis magellanica Bartsch, 1911, described froma

locality near Punta Arenas, and the Miocene fossil

C erithiopsis pyrgiscus (Philippi, 1887), only larval forms

of both families were reported from coast al Chile (Romero

and Valdebenito, 2002), in addition to beach-worn or empty

shells without specific determination (Marincovich, 1973:

Guzman et al., 1998). Two Newtoniellidae species were

reported to occur in the continental shelf of Chile: Atax-

ocerithium pullum (Philippi, 1545), which has a mainly

magellanic distribution and was recorded as far north as

Chiloé Island in the Pacific (Cardenas et al., 2008), and

Eumetula michaelseni (Strebel, 1905), from the Straits of

Magellan (Strebel, 1905).

Page 2

The objective of the present study is to evaluate the

taxonomy of recently sampled species of Triphoroidea

from Chilean waters, with two species from the conti-

nental shelf of northern Chile and one from the Juan

Fernandez Archipelago.

MATERIALS AND METHODS

Part of the material was collected by trawls in the Juan

Fernandez Archipelago, a set of three volcanic islands sit-

uated 670 kn off continental (central) Chile, in 2004. Hand-

collected specimens were obtained from Caldera, northem

coastal Chile, in 2012 and 2016, by the second author. In the

last case, some live specimens were found among sponges.

Color photographs were obtained with a Zeiss AxioCam

ICc5 camera on a Zeiss Discovery.V20 stereomicroscope,

and scanning electron microscope (SEM) images taken

with a JEOL JSM-6390LV. Shell descriptions followed

Fernandes and Pimenta (2015), and were solely based on

the examined material. For specimens with soft parts, the

shell was photographed and cracked, then the external

morphology was photographed, and internal hard struc-

tures (operculum, jaw and radula) extracted following

Geiger et al. (2007). When necessary to avoid sonniaont

cusps of radular teeth were numbere d from the innermost

cusp (cusp 1), close to the rachidian/central tooth, toward

the outermost cusp, distant from the central tooth; cusps of

central teeth are numbered from left to right when many

cusps are present. The number of specimens or shells from

each lot is indicated between square brackets, and ‘d’ refers

to specimens partly or completely destroyed for the study

of internal hard structures. As a consequence of the 1 recent

fire at MNRJ on September 2018 (Zamudio et al., 2018),

only the type material of the new species herein described

was saved, whereas the ordinary material listed in this study

and stored at MNRJ was unfortunately lost.

Acronyms: CAS: California Academy of Sciences, San

Francisco, USA: MNR]: Museu Nacional do Rio de

Janeiro, Rio de Janeiro, Brazil; USNM: National Museum

of Natural History, Smithsonian Institution, W ashington,

USA: SBMNH.: Santa Barbara Museum of Natural History,

Santa Barbara, USA

SYSTEMATICS

Subclass Caenogastropoda Cox, 1960

Superfamily Triphoroidea Gray, 1847

Family Cerithiopsidae Adams and Adams, 1853

Genus Cerithiopsis Forbes and Hanley, 1850

Type Species: Cerithium tubercularis Montagu, 1803, by

original designation. Recent, Europe.

Cerithiopsis eiseni Strong and Hertlein, 1939

(Figures 1-18)

Cerithiopsis eiseni Strong and Hertlein, 1939: 216, pl. 20,

fig. 6; Hertlein and Strong (1955: 135, not figured).

Type Locality: Bahia Honda, 5-16 m, Veraguas, western

Panama.

THE NAUTILUS, Vol. 133, No. 1

Type Material: Holotype: CAS 65665. The additional

shells listed in Strong and Hertlein (1939) are also stored

at CAS. Although they do not yet have lot numbers

(Christina Protowel pers. comm. NT they can be regarded

as paratypes (ICZN, 1999: article 72.4. i),

Material Examined: MNRB]J 23163, 27°05'15" S, 70°51'29" W,

Calderilla beach, under sunken rocks at very low tide,

Caldera, Region of Atacama, J. F. Araya and M. Araya

coll., February 9, 2016, 5 specimens, 3 d.

Description: Shell: dextral, ovoid to slightly elongate,

very thick, concave prone up to 3.4 mm long, 1.4 mm

wide, ratio length/width 2.3 to 2.5. Protoconch multispiral,

conical/cylindrical but slightly mammillated, 0.35 mm

long, 0. 30 mm wide: 3.5 much convex whorls, clear

transition between protoconch and teleoconch: embry-

onic shell dome-shaped; larval shell mainly smooth, except

by very fine, near orthocline to slightly prosocline, axial

threads covering entire length of last whorl. Teleoconch

with up to six whorls; two main spiral cords (median and

abapical) at beginning, with third one (adapical) very close

to median one, gradually developing and reaching nearly

same size of other cords on fifth whorl: on body whorl,

mean distance between cords is 1.1-1.4X higher than

mean width of cords; 15 to 19 orthocline to slightly

prosocline axial ribs on fifth whorl; rounded (particularly

on adapical and median cords) to slightly elliptical (par-

ticularly on abapical cord) nodules of large size; distinct

suture, with small, partially hidden sutural cord; thick,

wavy to weakly nodulose subperipheral cord, two smooth

to slightly wavy basal cords, adapical one considerably

antler small, elliptical aperture, 0.72-0.74 mm long,

0.50—0.52 mm wide, ratio length/vidth 1.4; anterior eave

very short, wide, completely open, 0.16—0.18 mm long,

0.19-0.23 mm wide; posterior canal as small notch,

0.11—-0.12 mm long. Teleoconch with reddish-brown to

dark-brown background, spiral cords dark brown with

yellowish nodules, protoconch yellowish to light golden.

External Morphology of Soft Parts: Color cream-

white, five to six light brown stripes mantle cavity roof;

adapical portion oF half to one whorl posterior to head

covered by several small black dots.

Opercalum: Elliptical, thin, semi-transparent, paucispiral,

2.5 whorls, nucleus eccentric, displaced 72% from center

toward margin; maximum length of operculum exceeds

diameter of opercular disc by 23% to 60% on anterior edge.

Radula: Only marginal teeth were discemible, 32.1—38.6 wm

long, usually bearing seven elongated and curved cusps on

alisis il portion of teeth (occupying 11% to 25% of their total

length, positioned almost perpendicular to main axis of teeth),

most external cusps much reduced.

Remarks: Cerithiopsis eiseni, originally described from

western Panama, is identical in teleoconch morphology

to shells herein studied from Chile. The thick sculpture

and ovate outline of the small shell, its dark brown color,

a moderately slow development of the adapical spiral

cord of teleoconch, the shape of the aperture and of the

M.R. Fernandes and J.F. Araya, 2019

Figures 1-9. Cerithiopsis eiseni Strong and Hertlein, 1939. 1-9. MNRJ 23163. 1, 4. 3.0 mm long. 2. 3.1 mm long. 3. 3.4 mm long.

5-9. Same shell as 4. 5. Development of the adapical spiral cord of teleoconch. 6-7. Body whorl, frontal and dorsal view. 8-9.

Protoconch, frontal and apical view. Scale bars: 1-4 = 1 mm; 5-7 = 500 pm; 8-9 = 100 pm.

Page 4

THE NAUTILUS, Vol. 133, No. 1

Figures 10-18. Cerithiopsis eiseni Strong and Hertlein, 1939. 10-18. MNRJ 23163. 10-14. External morphology of soft parts:

10-12, same individual, 13-14, another. 15. Operculum. 16-18. Marginal radular teeth. Scale bars: 10-14 = 500 pm; 15 = 100 ym; 16,

18 = 5 pm; 17 = 10 pm.

basal cords are the main features shared by them, in

addition to an apparent bathymetric re striction to the

shore zone. However, the type material of C. eiseni does

not have a protoconch, pre ‘cluding further comparisons

in relation to this structure. The original record of C.

eiseni from Galapagos by Hertlein aad Strong (1955)

requires confirmation, because no shell was fioured and

cerithiopsids may have a restricted larval dispersal in

M.R. Fernandes and J.F. Araya, 2019

Page 5

the open sea when compared to triphorids (Fernandes

and Pimenta, 2017b).

Cen ‘ithiopsis eiseni shares some features with the

Panamic species Cerithiopsis neglecta (C.B. Adams,

1852), including the brownish shell end similar teleoconch

sculpture, but C. eiseni can be distinguished by the larger

nodules on teleoconch, more ovoid shell shape, and light-

colored protoconch with less whorls and weaker orna-

mentation. The Californian specie Ss Cerithiopsis OXYS

Bartsch, 1911 also has a similar protoconch/teleoconch

sculpture, but has a more elongated and lighter-colored

shell, with the adapical cord of fale oconch strengthe ning

much earlier than in C. eiseni.

The operculum of C. eiseni follows the ‘pattern 2’

described by Marshall (1978), possessing a submarginal or

eccentric nucleus, which is found in several genera Dut not

Cerithiopsis sensu Marshall (1978)- ahitelh causes a

problem with that generic definition (Cecalupo and

Robba, 2010: Modica et al., 2013). The ovate shell outline

and presence of thick basal cords in C. eiseni (Figure 6)

are more similar to Joculator Hedley, 1909 her to

Cerithiopsis. These two genera certainly share consid-

erable conchological inter Teetons especially after recent

works (e.g., Jay el Drivas, 2002; Cecalupo and Perugia,

2012, 201 13, 2014) that substantially increased the amount

of known Joculator species, and both may be proved to

contain a number of non-natural groups after a robust

phylogenetic approach. Similar opercula to C. eiseni in-

clude those of Joculator cf. varians Laseron, 1956 (Niitzel,

1998: fig. 6D), one species of the complex of Cerithiopsis

tubercularis (Montagu, 1803) (Niitzel, 1998: fig. 1E) and

Tubercliopsis macalpinei (Laseron, 1951) (Laseron, 1951:

fig. 40), in addition to some species of the newtoniellid

genus Retilaskeya (e.g., Marshall, 1978: fig. 31).

The marginal teeth of C. eiseni fit the concept of the

“spaghetti- madaleg or “Cerithiopsis-group” radula defined

by Niitzel (1998), comprising several species (including

some of Cerithiopsis) that bear elongated teeth. We in-

clude the radula of the western Atlantic species Cer-

ithiopsis prieguei Rolan and Espinosa, 1996, Cerithiopsis

aimen Roldan and Espinosa, 1996 and Cerithiopsis gem-

mulosa (C. B. Adams, 1850)-illustrated in Rolan and

Espinosa (1996) and Rolan et al. (2007)—in the list pre-

sented by Niitzel (1998). In contrast, the “Synthopsis-

group” radula defined by Niitzel (1998) bears brush-like

teeth, with short and broad basal plates and numerous thin

cusps. As addressed by Niitzel (1998), the “Synthopsis-

group” includes the genus Joculator, and it is clearly

different from the radula of C. eiseni. A phylogeny is

necessary to elucidate the generic allocation of this

species, which has the shell more similar to Joculator, whereas

the radula fits the concept of Niitzel’s “Cerithiopsis-group’.

Geographic Distribution: Panama: Honda Bay (type

locality); Chile: Caldera (this study). The record from

Galapagos requires further confirmation.

Bathymetric Distribution: shallow subtidal (this study)

to 16 m (type locality).

Family Triphoridae Gray, 1547

Genus Monophorus Grillo, 1877

Type Species: Trochus perversus Linnaeus, 1758, by

monotypy. Recent, northeastern Atlantic and Mediterranean.

Monophorus monocelha new species

(Figures 19-43)

Triphorinae sp.: Romero and Valdebenito (2002: 498,

fig. 6).

Type Locality: 97°05'15" S, 70°51'29" W,

beach, under sunken rocks at very low tide,

Regi6n de Atacama, Chile.

Type Material: Holotype: MNRJ 23164, JE Araya and

M. Araya coll., 09/1/2016. Paratypes: MNRJ 2 3165, type

locality, J.F. Araya and M. Araya coll., 09/i/2016, 7

specimens, 4 d.

Calderilla

Caldera,

Additional Material Examined: MZSP_ 118802,

96°57'17" S, 70°48'05" W, Los Patos Sector, Caldera,

Region de Atacama, J. F. Araya coll., October 10, 2012, 12

she ‘ils, worn: MZSP. 118806, Calderilla Beach, in sand at

low tide, Caldera, Regi6n de Atacama, J. F. Araya and

M. Araya coll., August 12, 2012, 3 shells, worn.

Etymology: the spe cific name alludes to the brown mark

between the eyes of specimens, re »sembling a unibrow (in

Portuguese, * ‘monocelha”). Epithet as a noun in apposition,

Diagnosis: very thick shell, brown teleoconch, yellowish

protoconch; median spiral cord emerges be stween_ sixth/

seventh whorl: brown mark between the eyes; radula 4-1-

1-1-4.

Description: Shell: Sinistral, elongated, very thick,

cyrtoconoid, slightly concave profile, up to 4.7 mm long

(broken apex), 1.8 mm wide, ratio lengthAvidth 2.4 to 2 5.

Protoconch multispiral, conical/cylindrical, ().45—0.48 mm

long, 0.36—0.40 mm wide; 4.5 convex whorls, well-defined

transition between protoconch and teleoconch; embry-

onic shell dome-shaped, reticulate sculpture; larval shell

with two distinct spiral cords, situated at 40% and 67% of

last whorl height, entirely crossed by nearly orthocline

axial ribs, nUmBbe ring 32 in last whorl. Teleoconch with up

to eight whorls; two main spiral cords at beginning,

abapical one continuous to one on protoconch; “anata

spiral cord emerges very weakly on sixth or seventh whorl,

closely bordering adapical cord, gradually developing and

re -aching same size of abapical core (adapical one more

prominent on late whorls, 1.3 thicker than others at

body whorl) after about one whorl; at body whorl, mean

width of cords is equal or up to 1.4X higher than mean

distance between cords; 19 to 21 orthocline axial ribs on

seventh whorl; rounded nodules of large size; distinct and

well-developed suture, with very camel sutural cord:

nodulose subperipheral cord, two equally thick bas il

cords, adapical one slightly to moderately nodulose,

abapical one smooth; two soniell supranume srical cords may

appear, one between me dian and abapical cords, another

Page 6 THE NAUTILUS, Vol. 133, No. 1

Figures 19-27. Monophorus monocelha new species. 19. MNRJ 23164, holotype, 4.1 mm long. 20-22. MNRJ 23165, paratypes.

20. 4.1 mm long. 20-22. 4.4 mm long. 23-24. Body whorl, dorsal and frontal view, same shell as 22. 25-27. Holotype. 25, 27.

Protoconch, frontal and apical view. 26. Embryonic shell. Scale bars: 19-22 = 1 mm; 23-24 = 500 pm; 25, 27 = 100 pm; 26 = 50 wm.

M.R. Fernandes and J.F. Araya, 2019

between abapical and subperipheral cords; small and oval

aperture, 0.84— 0.95 mm long, 0.73—0.75 mm wide, ratio

lengthAvidth 1.2 to 1.3; anterior canal curved downward/

backward, open or partly closed by projection of outer

lip, 0.40-0.43° mm long, 0.20— (0.33 mm wide, ratio

lengthAvidth 1.2 to 2.1 . posterior « canal as deep sinus,

(.18—0.23 mm long. Te Jeowonchh brown to dark brown,

protoconch yellowish.

External Morphology of Soft Parts: Color cream-

white, with a brown and irregular stripe in the base of

cephalic tentacles and hemecn eyes, resembling a ‘uni-

brow’; some black fecal pellets present along nae stine,

with a vesicular shape, anteriorly rounded and posteriorly

acute, 87-115 pm long.

Operculum: Rounded/ovate, thin, semi-transparent, mem-

branous, multispiral, poorly distinct whorls, nucleus sub-

central, dislocated 15% to 22% from center toward

marg

opercular disc by 15% to 30%.

Jaw: With micro-pores on external side.

Radula: Formula 4-1-1-1-4: central, lateral and inner

marginal teeth (M1 and M2) claw-shaped, outer mar ginal

feelin (M3 and M4) somewhat hand-shaped; central foo

with five cusps, of which three central ones (cusps 2, 3,

and 4) are triangular and wide (cusp 3 is 1.2—2X shorter

than cusps 2 and 4), in addition to pair of small marginal

cusps (cusps 1 and 5) that attain maximum length of 35%

to 50% of cusps 2 and 4; lateral teeth with five (or six)

triangular cusps of similar size, cusp 4 (or 5) can be up to

1.7X longer than remaining cusps, all cusps of lateral

teeth narrower than three central cusps (2, 3, and 4) of

central tooth; M1 usually with four cusps, cusp 3 more

prominent (up to 1.3 longer than cusps 2 and 4), cusp 1

absent or up to 1.3 shorter than cusps 2 and 4; M2 with

five to six cusps, external ones (cusps 1 and 5 or 6) re-

duced; M3 with four cusps, cusps 2 and 3 1.82.3 X longer

than cusps 1 and 4; M4 with four (or even five) cusps,

external ones (cusps | and 4 or 5) reduced; central tooth

3.9-4.8 wm wide, lateral teeth 3.7-4.3 pm wide, M1

2.4-2.9 wm wide, M2 2.7-3.0 tm wide, M3 2.0-2.7 wm

wide, M4 1.5—2.4 wm wide.

Remarks: Despite the great variation in shell color in the

genus, many species ae Monophorus have brown shells

es g.., Bouchet, 1985: Roldn and Pefias, 2001); this is the

case of Monophorus monocelha new species. The new

species is distinct from any other described triphorid from

the eastern Pacific, showing some superficial similarity

with the dark shells of 7) iphora inconspicua C. B. Adams,

1852, from western Panama, and Triphora peninsularis

Bartsch, 1907, from Baja Califomia. Monophorus mon-

ocelha differs from the holotype of T. peninsularis

(USNM 106424) by having a much more developed su-

ture, darker brown color Gor pale brown as the holotype

of T. peninsularis) and later emergence of the median

spiral cord of teleoconch (in the sixth/seventh whorl of

M. monocelha, but fourth whorl in the holotype of

gin; diameter of operculum exceeds diameter of

T. peninsularis (M.R. Femandes, pers. obs.) or fifth whorl

(Bartsch, 1907). One worn paratype of T. peninsularis

(USNM 635558) more closely resembles M. monocelha,

although still showing a much reduced suture; types of

T. peninsularis will be illustrated in an oncoming study

about Triphoroidea from the eastern Pacific (Fernandes

et al., in prep.).

Monophorus monocelha corresponds to the larval form

designated as ee in Romero and Valdebenito

(2002: 498, fig. 6), from Punta de Lobos, ~330 kin

southern of thet type locality. Sculpture, number of whorls,

and length of protoconch (up to 0.48 mm in shells herein

studied vs. 0.51 mm in Punta de Lobos) are nearly

identical between them.

Monophorus monocelha has an unusual brown mark

between the eyes (Figures 28, 30, 32), a feature also

described (but not illustrated) for the eastern Atlantic

species Monophorus pantherinus Roldn and Penas, 2001,

although M. monocelha does not have reddish patches

along the head-foot as observed in most species of the

genus (e.g., Bouchet and Guillemot, 1978; Bouchet,

1985): Ghrel however, could be explained by discoloration

caused by storage in ethanol. The simple morphology of

the operculum ( (Figures 34-36) is similar to those de-

scribed for the genus (e.g., Marshall, 1983; Fernandes and

Roldan, 1988; Romani, 3015). The distinctive radula of

M. monocelha shows only four marginal teeth in each side

(Figure 40), the smallest number ae teeth per row so far

recorded in Monophorus. The previous smallest numbers

of teeth per row were those of Monophorus ateralbus

Roldan and Fermandez-Garcés, 1994 and M. pantherinus,

both with six marginal teeth on each side.

Geographic Distribution: Chile: Caldera.

Bathymetric Distribution: shallow subtidal.

Genus Inella Bayle, 1879

Type Species: Triforis (Ino) gigas Hinds, 1843, subsequent

designation by Jousseaume (1884). Recent, southwestern

Pacific.

Inella basalis (Odhner, 1922) new combination

(Figures 44-62)

Trifora basalis Odhner, 1922: 223, pl. 8, fig. 7; Rozbaczylo

and Castilla (1987: 176, species list); S Spates rg and Warén

(1993: 122, species list).

Type Locality: Masatierra, 20-35 m, Juan Fernandez

Archipelago.

Type Material: The single shell described by Odhner was

not found at the Gothenburg Natural History Museum

(Dr. Kennet Lundin, pers. comm.), where other types of

Odhner are stored, nor at the Swedish Museum of Natural

History (Dr. Anna Persson, pers. comm. ).

Material Examined: Robinson Crusoe Island, Juan

Fernandez Archipelago: MNRJ 23161 [6 shells], MNR]

23162, 4 shells, Cumberland Bay, among coarse sand and

Page § THE NAUTILUS, Vol. 133, No. 1

Figures 28-36. Monophorus monocelha new species. 28-36. MNRJ 23165, paratypes. 28-33. External morphology of soft parts:

28-29, same individual, 30-31, other, 32-33, another. 34-36. Opercula. Scale bars: 28-31 = 1 mm; 32-33 = 500 jm; 34-36 = 100 pm.

pebbles, February 2004; SBMNH 457461, 33°40'13" S, ratio lengthAvidth 2.9 to 3.2. Protoconch paucispiral, very

78°56'11" W, EI Padre Bay, March 25, 2004, 5 shells. short, Iskernis 0.38—0.43 mm long, last whorl 0.45—0.66 mm

wide; 2.0-2.5 somewhat axially compressed whorls,

Description: Shell: Sinistral, elongated, cyrtoconoid, well-defined but gradual transition between protoconch

slightly concave profile, up to 9.9 mm long, 3. 4 mm wide, (without nodules ) and teleoconch (with nodules): first

M.R. Fernandes and J.F. Araya, 2019 Page 9

Figures 37-43. Monophorus monocelha new species 37-43. MNRJ 23165, paratypes. 37-38. Jaw, outer side. 39-43. Radula.

Seale bars: 37 = 20 pm: 38 = 1 pm; 39-40, 43 = 10 pm; 41, 42 = 2 wm. Letters: C, central tooth; L, lateral teeth; M1—M4, marginal

teeth.

Dare

Page 10

THE NAUTILUS, Vol. 133, No. 1

Figures 44-53.

45-46. SBMNH 457461. 45. 9.9 mm long. 46. 9.1 mm long. 47-50. MNRJ 2:

50. 6.0 mm long. 51-53. MNRJ 23162. 51. 4.3 mm long. 52-53. 4.1 mm long. Scale bars: 1 mm.

whorl small, de »pressed or slightly elevated; two strong,

smooth spiral cords, situate »d at 22-39% and 62-76% BE

last whorl height, with nearly same size or abapical cord

slightly more prominent toward end of protoconch;

stil sculpture absent except for occasional minute axial

marks above adapical cord. Teleoconch with up to 9.5

whorls; two spiral cords at beginning, both continuous

to equivalent cords on protoconch; adapical spiral cord

appearing on first or second whorl, gradually developing

and re aching nearly same size of other cords on

sixth/eighth whoul: at body whorl, mean width of cords

1.2-1.8X higher than mean distance between cords:

spiral microsculpture may occur between cords, espe-

cially on late whorls; 15 to 21 nearly orthocline

(sometimes slightly opisthocline) axial ribs on seventh

Inella basalis (Odhner, 1922) new combination. 44. Drawing of the holotype, from Odhner (1922: pl. 8, fig. 7).

3161. 47.6.8 mm long. 48. 6.5 mm ae 49.6.4 mm ae

whorl; rounded (especially adapical and median cords)

to slightly elliptical (especially abapical cord) nodules of

large! size; well-developed suture, with distinct sutural

Corl thick, smooth subperipheral cord, two to four

smooth basal cords, equally spaced; two small supra-

numerical cords may appear, one between median and

abapical cords, other between abapical and sub-

peripheral cords; ovate aperture, 0.81—2.10 mm long,

0.52-1.44 mm wide, ratio length/width 1.3 to 1.6; an-

terior canal very short, moderately wide, entirely open,

0.20-0.47 mm long, (.23-0.54 mm wide, ratio le ngth/

width 0.8 to 1.0; posterior canal as a moderately dee Pp,

triangular notch, 0.11—-0.31 mm long. Shell cream,

beige, rarely light brown, with light Brown internodular

spaces.

M.R. Fernandes and J.F. Araya, 2019

Page 11

Figures 54-62.

in the body whorl. 60-62. Protoconch, apical and frontal view. Scale bars: 54-58 = 500 xm; 59 =

100 ym.

Remarks: Despite Odhner’s (1922) statement that the

type of I. basalis has a broken apex, the described features

of the teleoconch agree with the material herein examined,

including the presence of four basal cords (two to four cords

in this study; Figures 54, 57), shell dimensions of 6.3 X

2.0 mm (up to 9.9 X 3.4 mm in this study) and adapical

spiral teleoconch cord reaching same size of other cords

since fifth whorl (but since sixth hou or later in this study).

Inella basalis shows considerable intraspecific variation in

Inella basalis (Odhner, 1922) new combination. 54-56, 60-61. MNRJ 23161. 57-59, 62. MNRJ 23162. 54-55,

57-58. Body whorl, ventral and dorsal view. 56. Developme nt of the adapical spiral cord of teleoconch. 59. Spiral microsculpture

50 pm; 60-61 = 250 pm; 62 =

some shell features, such as: adult size (3.5-9.9 mm); shape

of protoconch (more or less axially compressed, blunt and

wide); number of axial ribs in the teleoconch (seventh

whorl: 15 to 21); number of basal cords (two to four); and

presence of supranumerical cords.

We abstain from designating a neotype for I. basalis due

to an ongoing search ane attempts at retrieval of past-due

loans from the Gothenburg Natural History Museum to

other institutions. Such search may still yie Id the return of

Page 12

THE NAUTILUS, Vol. 133, No. 1

the holotype to that institution (Dr. Kennet Lundin, pers.

comm.).

Geographic Distribution: Chile: Juan Fernandez Ar-

chipelago (type locality).

Bathymetric Distribution: 20-35 m (type locality).

DISCUSSION

The present study records Inella basalis (Odhner, 1922)

new combination in the Juan Fernandez Archipelago

almost a century after its original description. Two specie S

from the northern continental shelf of Chile are also

studied, the cerithiopsid Cerithiopsis eiseni Strong and

Hertlein, 1939, previously known with certainty only from

western Panama, and the triphorid Monophorus mono-

celha new species. These are the only extant species of

both families from the continental shelf of Chile, in ad-

dition to Cerithiopsis magellanica. The reduced richness

of Triphoroidea therein might be explained by the low

water temperatures of the W arm Te apes Southeast-

ern Pacific province (Spalding et al., 2007) and/or merely

reflects inadequate sampling. The taxonomy of eastern

Pacific Triphoroidea has been never revised and is highly

problematic, owing to several species described from

worn material and ao illustrations in old works. An

extensive review of Triphoroidea types from the eastern

Pacific is being conducted (Fernandes et al., in prep.),

aiming to provide the backbone for future descriptions.

The two species from northern continental shelf of

Chile, C. eiseni and M. monocelha, have a similar shell

morphology regarding the brown color, thickness, and

narrow aperture. Very thick shells (with strong sculpture)

and narrow apertures probably reduce dhe predation

by crabs (Zipser and Vermeij, 1978; Bertness and

C ‘unningham, 1981), considering that the eastern Pacific

often presents gastropod shells thick and with narrow

apertures as suggestive of anti-predation mechanisms

(Vermeij, 1974). The existence of thicker shells of parasitic

or micropredator snails, such as those of Triphoroidea,

suggests fre quent switch of hosts or preys, as this may be

ache vantageous to avoid predation when outside of the

protective cover furnished by hosts/preys (Vermeij, 2015).

In addition, particular hydrodynamic conditions cannot be

ruled out as sources of selective pressures for differences

in shell thickness (e.g., Sitnikova and Maximova, 2016).

ACKNOWLEDGMENTS

We are grateful to: Dr. Kennet Lundin, Gothenburg

Natural History Museum, for his tireless search for the

type of Inella basalis; Drs. Alexander Niitzel, Bruce

Marshall, Daniel Geiger, and Emilio Rolan, for making

several important suggestions during the review of the

manuscript; Dr. Anders Warén, who helped with some

references; and Camila Messias, for operating the SEM

at the Department of Invertebrates (MNRJ). CNPq

(Conselho Nacional de Desenvolvimento Cientffico

e Tecnolégico, Brazil) provided a doctoral grant (number:

142371/2014- 9) to the first author during “inks study.

LITERATURE CITED

Adams, C.B. 1852. Catalogue of shells collected at Panama, with

notes on synonymy, station, and habitat. Annals of the

Lyceum of Natural History of New York 5: 229-549.

Albano, P.G., B. Sabelli, and P. Bouchet. 2011. The challenge of

small and rare species in marine biodiversity surveys:

microgastropod diversity in a complex tropical coastal en-

vironment. Biodiversity & Conservation 20: 3223-3237.

Baker, F. 1926. Expedition of the Califomia Academy of Sci-

ences to the Gulf of California in 1921. Mollusca of the

family Triphoridae. Proceedings of the California Academy

of Sciences 15: 223-239.

Baker, F., G.D. Hanna, and A.M. Strong. 1938. Some Mollusca

of the { families Cerithiopsidae, Ganthiidee and Cyclo-

strematidae from the Gulf of California and adjacent wa-

ters. Proceedings of the California Academy of Sciences 23:

217-244.

Bartsch, P. 1907. The West American mollusks of the genus

Triphoris. Proceedings of the United States National

Museum 33: 249-262.

Bartsch, P. 1911. The Recent and fossil mollusks of the genus

Cerithiopsis from the West Coast of America. Proceedings

of the United States National Museum 40: 327-367, pls.

36-41.

Bertness, M.D. and C. Cunningham. 1981. Crab shell-crushing

predation and gastropod architectural defense. Journal of

Experimental Marine Biology and Ecology 50: 213-230.

Bouchet, P. 1985. Les Triphoridae de Mediterranee et du

proche Atlantique (Mollusca, Gastropoda). Lavori S.I.M.

21: 5-58.

Bouchet, P. and H. Guillemot. 1978. The Triphora perversa-

complex in Western Europe. Journal of Molluscan Studies

44: 344-356.

Bouchet, P., P. Lozouet, P. Maestrati and V. Héros. 2002.

Assessing the magnitude of species richness in tropical

marine environments: exceptionally high numbers of

molluscs at a New Caledonia site. Biological Journal of the

Linnean Society 75: 421436.

Cardenas, J., C. Aldea, and C. Valdovinos. 2008. Chilean marine

Mollusca of northern Patagonia collected during the

CIMAR-10 Fjords Cruise. Gayana 72: 202-240.

Cecalupo, A. and I. Perugia. 2012. Family Cerithiopsidae H.

Adams & A. Aghuins, 1853 in the central Philippines

(Caenogastropoda: Triphoroidea). Quademi della Civica

Stazione Idrobiologica di Milano 30: 1-262.

Cecalupo, A. and I. Perugia. 2013. The Cerithiopsidae (Cae-

nogastropoda: “ Triphoroidea) of Espiritu Santo -Vanuatu-

(South Pacific Ocean). Published by the authors, Milan,

255 pp.

Cecalupo, A. and I. Perugia. 2014. Cerithiopsidae and New-

toniellidae (Gastropoda: Triphoroidea Gray) from French

Polynesia area (South Pacific Ocean). Novapex 15: 1-22.

Cecalupo, A and E. Robba. 2010. The identity of Murex

tubercularis Montagu, 1803 and description of one new

genus and two new species of the Cerithiopsidae (Gas-

tropoda: Triphoroidea). Bollettino Malacologico 46: 45-64.

DuShane, H. and B.C. Draper. 1975. The genus Seila in the

eastern Pacific. The Veliger 17: 335-345.

M.R. Fernandes and J.F. Araya, 2019

DY, > C

Page 13

Femandes, F. and E. Roldan. 1988. A familia Triphoridae

(Mollusca: Gastropoda) no Arquipélago de Cabo Verde.

Publicacdes Ocasionais da Sociedade Portuguesa de Mal-

acologia 11: 17-32, pls. I-IV.

Femandes, M.R. and A.D. Pimenta. 2015. Five new species and

two records of Triphorinae (C Sa Triphoridae)

from Brazil. Zootaxa 4012: 493-5

Femandes, M.R. and A.D. Pimenta. Bolts Synopsis of the

deep-sea groups of Triphoroidea (Gastropoda). Journal of

Natural History 51: 853-865.

Femandes, M.R. and A.D. Pimenta. 2017b. A study of triphorid

larvae and post-larvae at the Campos Basin deep- sea floor,

southeastern Brazil (Gastropoda: Triphoroidea). Journal of

Natural History 51: 867-881.

Geiger, D.L., B.A. Marshall, W.F. Ponder, T. Sasaki and A.

Warén. 2007. Techniques for collecting, handling,

preparing, storing, and examining small molluscan

specimens. Molluscan Research 27: 1-50.

Guzman, N., S. Sad, and L. Ortlieb. 1998. Descriptive catalogue

of aero molluscs (Gastropoda and Pelecypoda) eosin

Antofagasta area, 23°S (Chile). Estudios Oceanolégicos 17:

] 7-86.

Hertlein, L.G. and A.M. Strong. 1955. Marine mollusks col-

lected at the Galapagos Islands during the voyage of the

Velero HI, 1931-1932. In: Essays in the natural sciences in

honor of Captain Allan Hancock. University of Southem

California Press, Los Angeles, pp. 111-145.

Houston, R.S. 1985. Genital ducts of the Cerithiacea (Gastro-

poda: Mesogastropoda) from the Gulf of California. Journal

of Molluscan Studies 51: 183-189.

Jay, M. and J. Drivas. 2002. The Cerithiopsidae (Gastropoda) of

Reunion Island (Indian Ocean). Novapex 3: 1-45.

Jousseaume, F. 1884. Monographie des Triforidae. Bulletins de

la Société Malacologique de France 1: 217-270.

Laseron, C.F. 1951. Revision of the New South Wales Cer-

ithiopsidae. Australian Zoologist 11: 351-368.

Marincovich, L. 1973. Intertidal mollusks of Iquique, Chile.

Science Bulletin, Natural History Museum of Los Angeles

County 16: 1-49.

Marshall, B.A. 1978. Cerithiopsidae (Mollusca: Gastropoda) of

New Zealand and a provisional classification of the family.

New Zealand Journal of Zoology 5: 47-120.

Marshall, B.A. 1983. A revision of the recent Triphoridae of

Southem Australia (Mollusca: Gastropoda). Records of the

Australian Museum, supplement 2: 1-119.

Modica, M.V., P. Mariottini, J. Prkic, and M. Oliverio. 2013.

DNA- barcoding of sympatric species of ectoparasitic

gastropods of the genus Cerithiopsis (Mollusca: Gas-

tropoda: Cerithiopsidae) from Croatia. Journal of the

Marine Biological Association of the United Kingdom

93: 1059-1065.

Niitzel, A. 1998. Uber die Stammesgeschichte der Ptenoglossa

(Gastropoda). Berliner Geowissenschaftliche Abhandlungen,

ser. E (Palaeobiologie) 26: 1-229.

Odhner, N.H. 1922. Mollusca from Juan Fernandez and Easter

Island. In Skottsberg C. (ed). The Natural History of Juan

Fernandez and Easter Island. ele & Wiksells Bok-

tryckeri, Uppsala, pp. 219-253, pls. 8—9.

Raines, B.K. 2002. Conisanntions & the knowledge of Easter

Island Mollusca. La Conchiglia 34 (304): 11-40.

Rehder, H.A. 1980. The marine mollusks of Easter Island (Isla

de Pascua) and Sala y Gomez. Smithsonian Contributions to

Zoology 289: 1-167.

Rolan, E. and J. Espinosa. 1996. La familia Cerithiopsidae

(Mollusca: Gastropoda) en Cuba. 3. El género Cerithiopsis

s. l., especies con concha castafia. Iberian 13: 129-147.

Rolan, E. and A. Pefias. 2001. Two new species of the genus

Monophorus (Gastropoda, Triphoridae) in the east Atlantic

and Mediterranean Sea. Iberus 19: 31—40.

Rolan, E., J. Espinosa and R. Femandez-Garcés. 2007. The

family Cerithiopsidae in Cuba. 4. The genus Cerithiopsis s.

2 the banded and the variably Boloured species. Neptunea

: 1-29.

ee L. 2015. A new Mediterranean Monophorus species

(Gastropoda: Triphoridae). Bolletino Malacologico 51: 3-5.

Romero, M.S. and E.L. Valdebenito. 2002. Larvas veliger de

gastropodos Prosobranchia provenientes de Punta de

Lobos, Cuarta Region, Chile. Revista Chilena de Historia

Natural 75: 491-514.

Rozbaczylo, N. and J.C. Castilla. 1987. Invertebrados marinos

del archipiclago de Juan Fernandez. In: J.C. Castilla (ed.).

Islas ocednicas chilenas: conocimiento cientifico y neces-

idades de investigaciones. Ediciones Universidad Catolica

de Chile, Santiago, PP: 167-189.

Sandberg, L. and A. Warén. 1993. Molluscan taxa introduced by

Nils Hjalmar Odhner. Malakologische Abhandlungen 16:

117-134.

Sitnikova, T. and N. Maximova. 2016. On morphological and

ecological evidence of adaptive differentiation among stony

cliff littoral Baikal gastropods. Journal of Natural History

50: 263-280.

Spalding, M.D., H.E. Fox, G.R. Allen, N. Davidson, Z.A.

Ferdana, M. Finlayson, B.S. Halpem, M.A. Jorge, A.

Lombana, i A. Lourie, K.D. Martin, E. McManus,

J. Molnar, C.A. Recchia, and J. Robertson. 2007. Marine

ecoregions oF the world: a bioregionalization of coastal and

shelf areas. BioScience 57: 573-583.

Strebel, H. 1905. Beitriige zur Kenntnis der Molluskenfauna der

Magalhaen-Provinz. 3. Zoologische Jahrbiicher — 22:

575-666.

Strong, A.M. and L.G. Hertlein. 1939. Marine mollusks from

Panama collected by the Allan Hancock Expedition to the

Galapagos. Islands, 1931-1932. Allan Hancock Pacific Ex-

peditions 2: 177-245.

Vermeij, G_]. 1974. Marine faunal dominance and molluscan

shell form. Evolution 28: 656-664.

Vermeij, G.J. 2015. Gastropod skeletal defences: land, fresh-

water, and sea compared. Vita Malacologica 13: 1-25.

Wells, F.E. 1998. Superfamily Triphoroidea. In: P.L. Beesley,

G.J.B. Ross and A. Wells (eds.). Mollusca: the Southern

synthesis. Part B. CSIRO Publishing, Melboume, pp. 808-811.

Zamudio, K.R., A. Kellner, C. Serejo, M.R. Britto, C.B. Castro,

P.A. Buckup, D.O. Pires, M. Coun, A.B. Kury, L.A. Car-

doso, M.L. Monné, J. Pombal Jr., C.M. Patiu, V. Padula,

A.D. Pimenta, C.R.R. Ventura, E. Hajdu, J. Zanol, E.M.

Bruna, J. Fitzpatrick, and L.A. Rocha. 2018. Lack of Sci-

ence support fails Brazil. Science 361: 1322-1323.

Zipser, E. and G.J. Vermeij. 1978. Crushing behavior of

tropical and temperate crabs. Journal of Experimental

Marine Biology and Ecology 31: 155-172.

THE NAUTILUS 133(1):14-21, 2019

Page 14

New species and record of Diplommatina Benson, 1849

(Gastropoda: Diplommatinidae) from Java, Indonesia

Kristine Greke

Darza iela 10, Stopinu novads

Dzidrinas, LV-2130 LATVIA

[email protected]

ABSTRACT

Diplommatina (sensu lato) majapahit new species from Java is

described and illustrated. A redescription and a new record for

D. (sensu lato) halimunensis Nurinsiyah et Hausdorf, 2017 is

provided.

Additional Keywords: Taxonomy, morphology, endemism, Greater

Sunda Islands

INTRODUCTION

The Diplommatinidae of Java were recently revised

(Nurinsiyah and Hausdorf, 2017) with 23 species recor-

ded. Diplommatina (sensu lato) Benson, 1849 is the

most diverse diplommatinid genus in Java (17 species)

(Nurinsiyah and Hausdorf, 2017) and neighbouring Bali

(6 of 10 species) (Vermeulen and Whitten, 1998). Recent

sampling from Java provided new data on one in-

sufficiently known species as well as a new species, which

is described here.

MATERIALS AND METHODS

Recently collected material was stored in 99% ethanol.

Te -rminology of morphological terms generally follows Liew

at al. (2014), with modifications by G Sidke ( (2017). The au-

thor’s explanatory comments are placed in square brackets.

Specimens were examined under a Leica S6D stereoscopic

microscope. Photographs were taken with a Canon EOS

77D SLR camera attached to that microscope. Multiple

photographs were taken at different focal planes and reas-

sembled using CombineZP ® software. Scanning electron

microscope (SE M) images were taken using SEM provided

by the Latvian University’s Faculty of Biology. Shells were

cleaned mechanically with a brush and with KOH solution,

air dried, and placed onto conductive tape on aluminum

stubs. Shells were imaged using a Hitachi TM-3000 (Hitachi

High Technologies ®) SEM following methods as described

by Geiger et al. (2007). Abbreviations of depository

insitutions: KGC: Collection Kristine Greke, Riga, Latvia;

ZMHB: Museum fiir Naturkunde, Leibniz-Institut ftir

Evolutions- und Biodiversitatsforschung [formerly Museum

fiir Naturkunde der Humboldt- Univers, Berlin, Ger-

many; NME: Naturkundemuseum Erfurt, Germany;

NMNL: National Museum of Natural History “Natu-

ralis’, Leiden, the Netherlands.

RESULTS

L. Pfeiffer, 1856

Diplommatinidae

Diplommatina (sensu lato) Benson, 1849

Diplommatina (sensu lato) majapahit new species

http://zoobank.org/CF925 BOC-7329-462B-B46E-

CF9OFI6CT7TBDB

(Figures 1-6, 23-27, 32-33)

Description: Shell large, pale yellowish to pale reddish,

dextral, broadly conical spa a convex apex. Holotype shell

height (H) 4.5 mm, height of the ultimate whorl (HW)

2.6 mm, shell diameter (D) 2.5 mm, maximum diameter

of peristome (PD) 2.1 mm. Adult paratypes:

Paratype No H, mm D, mm PD, mm

] 48 Did 2]

») 47 2.8 2

3 4.6 2.4 2

4 4.5 D5) 1.9

Shell with 6.5-7 convex whorls, the 1.5 embryonic whorl is

microscopically pitted. The suture is rather deeply

impressed. The ultimate whorl narrows in umbilical view

(not visible in apical view), hardly wider than the penultimate

whorl in apertural view. The umbilicus is closed in adults.

The constriction is poorly defined, suture somewhat deeper

at the constriction. The position of the constriction aligns

with the parietal side of the aperture. The teleoconch is

sculptured with coarse high and widely spaced axial ribs,

which become slightly more widely spaced on the last half

whorl. The ribs are generally straight. There are no abrupt

changes in ribbing pattern. The ane are not aligned with

K. Greke, 2019

Page 15

Figures 1-6. Diplommatina (sensu stricto) majapahit new species, holotype: 1

Lateral view. 5. Apical view. 6. Operculum, outer surface.

those on previous whorls and are straight on earlier whorls,

becoming slightly oblique to the coiling axis on the penul-

timate al the ultimate whorls. There are about 6 ribs per

1 mm on the penultimate whorl in apertural view. Spiral

striations absent (Figure 32), delicate radial growth lines

present on the ultimate whorl only (Figure 33). The aperture

is slightly tilted in relation to the coiling axis (in lateral view;

Figure 3), is circular, and has an entire apertural rim. The

position of the aperture is more or less central against the

coiling axis in apertural view. The peristome is double,

obtuse angular at the transition from columellar to basal side.

The outer peristome is broader than and expanding beyond

Apertural view. 2. Abapertural view. 3, 4.

the inner peristome, broadly discontinued parietally. The

inner peristome forms a continuous polished cz allus. The

parietal margin of the peristome is attached to the ultimate

whorl. The palatal and columellar margins of the outer

peristome are slightly sinuous. There are two strong acute

parie stalis ( Figures 26-27), one long and strong radial (ver-

tical) pi alatalis ( (position of this lamella is almost central with

regard to the parietal side of the peristome; Figure 1) and

two long and strong longitudinal (horizontal) palatalis: the

lower one >-shaped, somewhat shorter, level with columellar

side of the peristome and upper one longer, straight situated

somewhat beyond the columellar side of the peristome

Page 16 THE NAUTILUS, Vol. 133, No. 1

Figures 7-14. Diplommatina (sensu stricto) diplostoma B. Rensch, 1931. 7-9. Specimen from Mt. Adeng, Bali Island. 7. Apertural

view. 8, 9. Lateral view. 10-14. Holotype. 10. Apertural view. 11. Abapertural view. 12. Lateral view. 13. Apical view. 14. Aperture,

latero-ventral view.

kK. Greke, 2019

Page 17

(Figure 27). One strong obtuse columellar denticle (lamella)

present, not directed darmamers ( Figure 1). The columella

is broad, glossy, obliquely twisted ancl forming a very strong

acute umbrella-like obliq ue columellar lamella in its lower

third and with an radlethaet knob just above the columellar

lamella (Figures 23-25). The operculum is simple (primitive,

of Greke 30) 7) - irregularly circular and translucent, pau-

cispiral, without ridges (Figure 6).

Type Material: Holotype: NME unnumbered. Para-

types: Five specimens [3 adults and 1 juvenile KGC

unnumbered, | adult NME unnumbered]. All August 20,

2017, leg. K. Greke, from type locality.

Type Locality; INDONESIA, E Java, ~46 km SE

Malang, Tumpak Sewu waterfall, 8°12'54" S, 112°56'39" E,

~600— 610 m, primary lowland rainforest vegetation on

limestone cliffs, under large fallen leaves.

Ecology: Found in karst area of Tumpak Sewu waterfall.

Specimens inhabit leaf litter accumulated limestone

cavities, observed at very wet place near the waterfall.

Etymology: The new species is named after the

Majapahit Empire, a thalassocracy in Southeast Asia,

based on the eastern part of Java (part of modern-day

Indonesia), existed 1293-1500 and consisted of present-

day Indonesia, Brunei, Malaysia, Singapore, Sulu Ar-

chipe lago (the Philippines), southern T ibe uiland, and East

Timor. Named as a noun in apposition.

Distribution: Greater Sunda Islands: eastern part of Java

(hitherto only known from type locality)

Differential Diagnosis: This is one of the largest Javan

Diplommatina and is most similar to D. (sensu lato) ) diplo-

stoma B. Rensch, 1931 sensu Vermeulen and Whitten

Figures 15-20. Diplommatina (sensu stricto) duplicilabra van Benthem Jutting, 1948 (D. diplostoma of authors), holotype. 15.

Apertural view. 16. Abapertural view. 17, 18. Lateral view. 19. Apical view. 20. Operculum, outer surface.

Page 18

THE NAUTILUS, Vol. 133, No.

(1998), from Bali and Java. Diplommatina diplostoma is

highly variable in shell shape and size, density of axial ribs.

mie following are significant features of D. majapahit new

species: The inner peristome, weaker than the outer one,

forms a continuous broad polished callus in apertural view

(the inner peristome is more prominent and does not form

a polished callus in D. diplostoma); the columellar lamella

is strong and thick (not acute as in D. diplostoma) and

Figures 21-30. Internal lamellae of the Javan and Balinese Diplommatina species. 21, 22. Diplommatina (sensu stricto) diplostoma

B. Rensch, 1931, specimen from Mt. Adeng, Bali Island. 21. Columella and columellar lamellae. 22. Parietalis. 23-27. Diplommatina

(sensu stricto) majapahit new species, paratype. 23-25. Columella (arrows indicate a knob) and columellar lamella. 26. Parietalis.

27. All lamellae. 28-30. Diplommatina (sensu stricto) halimunensis Nurinsiyah et Hausdorf, 2017, specimen from Mt. Salak, Java

Island. 28. Columella. 29. Parietalis. 30. Longitudinal palatalis. Not reproduce -d to the same scale.

K. Greke, 2019 Page 19

Mh Oa

We he't

Figures 31-35. Micrographs of the Javan and Balinese Diplommatina species. 31. D. (sensu stricto) diplostoma B. Rensch, 1931,

specimen from Mt. Adeng, Bali Island, penultimate whorl (visible numerous pores are result of shell degradation, not the actual

microsculpture). 32-35. D. (sensu stricto) majapahit new species paratype. 32. Penultimate whorl. 33. Last whorl (note delicate growth

lines). 34, 35. D. (sensu stricto) halimunensis Nurinsiyah et Hausdorf, 2017 specimen from Mt. Salak, Java Island, penultimate whorl.

Page 20

THE NAUTILUS, Vol. 133, No. 1

provided with an indistinct knob above (columellar la-

mella acute, knob absent in D. diplostoma, including its

type series). Palatal and parietal teeth are generally

stronger in new species than in D. diplostoma (including

types). The new species appears slightly larger (H

4.5-4.8 mm compared to 2.6-4.3 mm in D. diplostoma)

and is one-half whorl bigger (6.5-7 whorls compared to

5.25-6.5 whorls in D. diplostoma, according to Nurinsiyah

and Hausdorf (2017) and Greke (personal observations)).

Material Examined of D. diplostoma: Holotype

[ZMHB] (Figures 7-14), Sunda=Expedition Rensch

Fundort: [printe d| Batoeriti (800m), Bali [handwritten]

Datum: [printed] 3.-4.8.27 [handwritten] No: [printed] /

Zoolog. Museum Berlin [printed] Diplommatina (Met-

adiancta ?) diplostoma Batoeriti Bali 800m 75427

Rensch leg [handwritten] / Zoolog. Museum Berlin

printed] Diplommatina (Metadiancta ?) diplostoma

Rensch Zool. Jahrb. Syst., 61, p. 389, fig. 14, 193]

handwritten] [label red]. Paratype [ZMHB}], Sunda-

=Expedition Rensch Fundort: [printed] Batoeriti

800m), Bali [handwritten] Datum: [printed] 3.-4.8.27

[handwritten] No: [printed] / Zoolog. Museum Berlin

printed] Diplommatina (Metadiancta ?) diplostoma

Rensch Paratypen. Batoeriti, Bali 75428 Rensch

[handwritten]; additional material [8 adults KGC]:

INDONESIA, Bali Is., Gunung Adeng mt., 08°19.402' S,

115°08.545' E, 1590 m, April , 2011, primary mid-montane

rainforest, soil sample No 5, leg. A.Riedel.

Measurements D. diplostoma: Holotype H 3.5 mm,

D 2.1 mm, HW 2mm, PD 1.4 mm, 8-9 full ribs per 1 mm

on the ultimate whorl in apertural view. Paratype

D. diplostoma: H 3.8 mm, D 2.2. mm, HW 2 mm, PD

1.6 mm, 10 full ribs per 1 mm. Selected specimens from

Bali: H 4.3 mm, D 2.5 mm, HW 2.3 mm, PD 1.9 mm:

H 4.3 mm, D 2.4 mm, HW 2.3 mm, PD 1.7 mm:

H 4.3 mm, D 2.4 mm, HW 2.3 mm, PD 1.8 mm.

Material Examined of D. duplicilabra: Holotype

.| (Figures 15-20), Kawah Idjen. Merapi Wild-

reservaat 1600-1700 m 8.VIII.1931 C.Th.Cribb [hand-

written] / type geteekend ex [handwritten] / type en

paratypen [handwritten] Zodlog. Museum. Amsterdam.

[printed] Diplommatina duplicilabra V.B.Jutting Kawah

Idjen. Merapi Wildreservaat 1600-1700 m 8 Aug. 193]

leg.C.Th. Cribb [handwritten] STADTSDRUKKERI]

AMSTERDAM [printed] / holotype 3.48.005 1 dr

ZOOLOGISCH MUSEUM AMSTERDAM ZMA Moll.

135940 Diplommatinidae 080.0 Diplommatina duplici-

labra Van Benthem Jutting, 1948 INDOENSIA Jawa

Kawah Idjen Merapi Game Reserve, 1600-1700 m 1931

08 08 sta. Leg.C.T. Cribb Ex. coll. Original publication:

Vn Benthem Jutting, W.S.S., 1948. Sy siomeatsie studies on

the non-marine Mollusca of Indo-Australian Archipelago

I. Families Hybrocenidae, Helicinidae, Cyclophoridae,

Pupinidae ... [partly unreadable] Autoref: Van Benthem

Jutting, 1948c: 597-599, fig. 54 Add.publ: Notes: Cur-

rent Scientific Name: Diplommatina duplicilabra Van

Benthem Jutting, 1948 Det. Operculum is glued on a

separate paper card with handwritten text "operc.".

Measurements D. duplicilabra: Holotype H 3 mm, D

1.7 mm, HW 1.6 mm, PD 1.2 mm, 15-16 full ribs per

1 mm on the ultimate whorl in apertural view. Spiral

striations not indicated.

After careful examination of types and additional spec-

imens of D. diplostoma, as well as the holotype of D.

duplicilabra van Benthem Jutting, 1948, it seems unlikely

that D. duplicilabra should be considered a junior synonym

of D. diplostoma as stated by Vermeulen and Whitten

(1998) and followed by Nurinsiyah and Hausdorf (2017).

Diplommatina (sensu lato) halimunensis Nurinsiyah

et Hausdorf, 2017

(Figures 28-30, 34-35)

Material Examined: INDONESIA, W Java, Halimun-

Salak NP, Mt. Salak, N of Kawah Ratu crater, 6°42'12" S,

106°42'10" E, August 30, 2017, 1225 m, primary lower

montane rainforest, leaf litter, leg. D.Telnov [10 adult

specimens KGC and 5 NME].

Notes: I provide illustrations of the internal lamellae of

this species for the first time. Parietal lamella is weakly

developed as a flat hump (Figure 29). Palatal lamella as in

Figure 30. Columella slossy and thick, widened basally

(Figure 28). In all orndtied| specimens, spiral striations are

distinct and dense, clearly visible under 80 magnifica-

tion (Figure 34). There are 3-4 rows of spiral tier per

50 wm measured on the penultimate whorl above the

constriction under 300 magnification) (Figure 35).

This species was originally ‘Gesoalxed from three speci-

mens ee .d at Mt. Halimun (Nurinsiyah and Hausdorf,

2017). New material comes from dense and very wet leaf

litter ot Mt. Salak volcano’s primary rainforest, that is

19-20 km NNE from the type locality in same geographical

and geological area.

ACKNOWLEDGMENTS

It is my ple asant duty to express gr. atitude to Dmitry Telnov

(The Entomological Society of Latvia, Riga) for assistance

with preparing photogr aphs used in this publication. I am

also deeply indebted to Bram van der Bijl (NMNL) and

Christine Zorn (ZMHB) for providing access to the types of

D. duplicilabra and D. ee The author is thankful to

Valters Gobin’ and Ugis Kagainis (both University of Latvia

Faculty of Biology, Department of Zoology and Animal

Ecology, Riga) fio producing the scanning dlecton micro-

graphs. Anonymous referees are thanked for ve iluable

comments on the final version of the manuscript. I am

indebted to Fred Naggs (Natural History Museum, London,

U.K.) for improving “the English.

LITERATURE CITED

Geiger, D.L., B.A. Marshall, W.F. Ponder, T. Sasaki, and

A. Warén. 2007. Techniques for collecting, handling,

K. Greke, 2019

Page 2]

preparing, storing and examining small molluscan speci-

mens. Molluscan Research 27: 1-50.

Greke, K. 2017. Taxonomic review of Diplommatinidae

(Caenogastropoda: Cyclophoroidea) from Wallacea and the

Papuan Region: 151-316, pls 19-47. In: Telnov, D., Barclay,

M. V. L. and Pauwels, O. S. G. (eds) Biodiversity, bio-

geography and nature conservation in Wallacea and New

Guinea. Volume III. The Entomological Society of Latvia,

Riga, 658 pp., 172 pls.

Liew, T.-S., J.J. Vermeulen, M.E. bin Marzuki, and M.

Schilthuizen. 2014. A cybertaxonomic revision of the

micro-landsnail genus Plectostoma Adam (Mollusca,

Caenogastropoda, Diplommatinidae), from Peninsular

Malaysia, Sumatra and Indochina. ZooKeys 393: 1-107.

Nurinsiyah, A.S. and B. Hausdorf. 2017. Revision of the Dip-

lommatinidae (Gastropoda: Cyclophoroidea) from Java.

Zootaxa 4312: 201-245.

Rensch, B. 1931. Die Molluskenfauna der Kleinen Sunda-Inseln

Bali, Lombok, Sumbawa, Flores und Sumba. I. Zoologische

Jahrbiicher. Abteilung fiir Systematik, Okologie und Geo-

graphie der Tiere 61: 263-396.

Vermeulen, J.J. and A.J. Whitten. 1998. Fauna Malesiana. Guide

to the Land Snails of Bali. Backhuys Publishers, Leiden,

ix + 164 pp.

THE NAUTILUS 133(1):22-25, 2019

Page 22

A Paleocene species of Procardia (Bivalvia: Anomalodesmata:

Parilimyidae) from the Katsuhira Formation in Urahoro Town,

eastern Hokkaido, Japan

Kazutaka Amano

Department of Geoscience

Joetsu University of Education

1 Yamayashiki

Joetsu 943-8512, JAPAN

[email protected].]p

ABSTRACT

A new species of anomalodesmatan bivalve, Procardia inouei, is

described from the Paleocene (upper Selandian to lowest

Thanetian) Katsuhira Formation in eastern Hokkaido, Japan.

The genus Procardia flourished during the late Jurassic to

Cretaceous, and this new species is the frst Paleocene record of

Procardia. Procardia clearly survived the end-Cretaceous mass

extinction, and includes the late Eocene to early Miocene

species P. dolicha (Suter, 1917) from New Zealand. The Recent

species of Panacca Dall, 1905 can be distinguished from

Procardia easily, but possibly were evolutionally derived from

Procardia.

Additional Keywords: Paleocene, bivalve, Procardia, new spe-

cies, Panacca

INTRODUCTION

The parilimyid genus Procardia Meek, 1871 is one of the

characteristic Mesozoic anomalodesmatan bivalves. Fos-

sils of this genus have been found from the Upper Jurassic

in Spain, the Lower Cretaceous in England, the Cen-

omanian in the United States Western Interior Sea and

the Maastrichtian in Germany, Poland, Ukraine and

Russia (Woods, 1909: Cox and Newell in Cox et al., 1969:

Runnegar, 1974; Jablonski and Raup, 1999; Ol6riz et al.,

2003). Only one younger species has been recorded, the

early Eocene to early Miocene species Procardia dolicha

(Suter, 1917), from the South Island of New Zealand (Beu

and Mexawlll 1990, p. 132, pl. 9h, i). Beu and Maxwell

(1990) considered this species to be a relict from the

Mesozoic Era in New Zealand.

On the other hand, Procardia is very similar to Panacca

Dall, 1905, which now lives in lower sublittoral to abyssal

depths. Eight species of Panacca are known, from off

West Africa, Meteor Seamounts, Massachusetts, Chile,

Indonesia, southwestern Japan and Tasmania (Coan,

2000; Krylova, 2006; Sasaki and Okutani, 2007). They are

carnivores or scavengers, burrowing in soft mud, and do

not prete r cold waters (Morton, 1981, 1982; Huber,

2010). Probably because of their ecology and their fragile

shell, their occurrence is so scarce that Panacca africana

(Locard, 1898), P. chilensis Coan, 2000, P. montana

Krylova, 2006, and P. trigona Sasaki and Okutani, 2007

were each based on a single specimen. However, there is

no fossil record of ans genus. Panacca differs from

Procardia mainly by lacking a lunule.

One articulated bivalve specimen of Procardia has been

collected from the Paleocene Katsuhira Formation (late

Selandian to earliest Thanetian; see Amano et al., 2018) in

eastern Hokkaido by Mr. K. Inoue (Obihiro City). I de-

scribe this species as new and discuss the relationship

between Procardia and Panacca.

MATERIALS AND METHODS

One articulated specimen of Procardia inouei new

species was obtained from the dark gray mudstone of the

Katsuhira Formation exposed along the Urahoro River

30 m north of the mouth of the olextcnlbitrerencra River,

Urahoro Town, eastern Hokkaido (Figure 1; 42°59'12" N,

143°37'35" E). The new species was associated with

Meganuculana alleni Amano and Jenkins, 2017, Thyasira?

sp., Neverita majimai Amano and Jenkins, 2018,

Kangilioptera inouei Amano and Jenkins, 2014 and

Biplica paleocenica Amano and Jenkins, 2018. As ob-

served by Amano et al. (2018), it is estimated that the

Katsuhira Formation was deposited at a depth of 200 to

500 m.

The holotype of the new species is catalogued in the

University Museum of the University of Tokyo (UMUT).

The associated fauna is stored at College of Science and

Engineering, Kanazawa University.

SYSTEMATIC PALEONTOLOGY

Superfamily Pholadomyoidea King, 1844

K. Amano, 2019

Okhotsk Sea =

Hokkaido Island |

Study area |" ( nS, — \

Pacific Ocean

F eee y, Katsuhira

(Locality iC Sees

Kokatsuhirazawa River

200m : i | Zi

=i

Figure 1. Map showing the type locality of Procardia inouei

new species. Base map om “Katsuhira,” original scale 1:25,000:

topographical map published by the C Geospatial Information

Authority of Japan.

Family Parilimyidae Morton, 1951]

Remarks: This family was separated from Pholado-

myidae King, 1844 based mainly on anatomical char-

acters, by Morton (1981, 1982). Judging from the fossil

record (Runnegar, 1974), Pholadomyoidea should be

allocated on the basal part of the bivalve phylogenetic

tree. Although the molecular sphylogeny of the

MMerlolocnats has been examined by Dreyer et al.

(2003) and Harper et al. (2006), Pholadomyoidea have

not been included, probably because living specimens

are difficult to obtain. Coan (2000), Krylova (2006), and

Sasaki and Okutani (2007) mistakenly attributed the

family Parilimyidae to Morton (1982). Morton (1981)

had already proposed this family, as indicated by Huber

(2010). When Coan (2000) included two genera, Par-

ilimya Melvill and Standen, 1899 and Panacca Dall,

1905, he considered Nipponopanacca Habe, 1977 to be

a synonym of Parilimya. However, as Matsukuma

(1989) discussed, by comparing it with the type ae

of Pholadomya, P. candida G. B. Sowerby I, 1823,

Nipponopanacca should be treated as a subgenus of

Pholadomya G. B. Sowerby I, 1823 (see also Olutemni,

2017). Runnegar (1974) suggested that Panacca is a

junior synonym of the Mesozoic genus Procardia Meek,

1871. As discussed be low, the genus Panacca is separated

from Procardia, which is also included in Parilimyidae.

Moreover, judging from its shape and shell sculpture,

Kanakimya Campbell and Grant-Mackie, 1995, from

Middle Jurassic rocks of New Caledonia, should be in-

cluded in this family.

Genus Procardia Meek, 1871

Type Species: Isocardia? hodgei Meek, 1871 by original

designation.

Remarks: Procardia is characterized by having an ante-

riorly truncated triangular shell, slightly highe r th: an long or

with he ight nearly equal to length, radial abs present over

the ole surface apart from fie anterior flat area, some

commarginal ribs or distinct growth lines, and a distinct

lunule on the flat area. The Recent genus Panacca Dall,

1905 differs from Procardia by its longer shell and by

having no radial ribs on the poste rior antl Moreover, as

pointe “a by Beu and Maxwell (1990), species of Panacca

lack a Junule on the anterior flat area. Kanakimya was

proposed by Campbell and Grant-Mackie (1995) as a

subgenus of Pholadomya from Middle Jurassic rocks in

New Calec lonia, based on only one species, Pholadomya

(Kanakimya) uitoe Campbell and Grant-Mackie, 1995.

Kanakimya differs from Procardia by having a sub-

truncated posterior margin and no radial “lbs on the

posterior surface, although it is similar to Procardia in

having a high shell with an anterior flat area and a lunule.

Procardia inouei new species

(Figures 2-5)

Diagnosis: Small, trigonal Procardia sculptured with 13

radial ribs. Radial ribs with central crest and gently sloping

sides; interspaces each wider than one rib. Anterior flat area

sculptured only with fine growth lines; with distinct lunule.

Description: Shell fragile, nacreous inside, small (24.9 mm

in length), trigonal, slightly higher than long (height

25.2 mm), equivalve, earemely inequilateral. Posterior

dorsal margin long, broadly arched, gradually merging into

subcircular posterior margin; ventral margin ne arly "straight;

anterior dorsal margin Sone anterior margin slightly con-

cave, truncated. Unie well inflated, locate ak at anterior one-

eighth (12.4 %) of shell. Whole surface apart from anterior

flat area sculptured with 13 radial ribs, each with central

crest, gently sloping sides, and fine growth riblets; in-

terspaces each wider than rib width. Anterior flat area

sculptured with distinct fine growth lines. Lunule rather

wide, moderately depressed, bounded by rough ridge.

Escutcheon de pressed, lanceolate, between strong radial

ribs of both valves. Ligament external, occupying Shell? of

postero-dorsal margin. Interior characters unknown.

Type Material: Holotype: UMUT CM 33112 (length

24.9 mm; height, 25.2 mm; depth, 21.5 mm).

Type Locality: Cliff along Urahoro River, 30 m north of

the mouth of Kokatsuhirazawa River, Urahoro Town,

eastern Hokkaido, Japan.

Remarks: Procardia inouei new species is similar to

Procardia decussata (Mantell, 1822) from the Cretaceous

Chalk in southeastern England. Both have a triangular

shell with radial ribs and growth riblets and a wide anterior

flat area. However, the present new species is smaller

(P. decussata attains 98mm in length; Woods, 1909) and

has fewer radial ribs than P. decussata (P. decussata has

18 radial ribs, which are fine on the posterior area).

Page 24 THE NAUTILUS, Vol. 133, No. 1

lunule

ligament

Figures 2-5. Procardia inouei new species (holotype, UMUT CM 33112) from the Katsuhira Formation. 2. Lateral view of right

valve. 3. Dorsal view. 4. Anterior view. 5. Lateral view of left valve.

Procardia hodgei (Meek,

Procardia, from the Campanian Pierre Shale in the In-

terior Sea Way area in the USA, differs from the present

new species by having many rounded radial ribs and

distinct commarginal grow th ribs (see Meek, 1876:

Runnegar, 1974).

Marwick (1944) orn d Cardium (Fragum) doli-

chum Suter, 1917 and C. (F.) maorinum Suter, 1917, both

with type specimens from The lower to middle Oligocene

in the South Island of New Zealand, and referred both to

Procardia dolicha. Beu and Maxwell (1990, p. 132, pl. 9h,

i) illustrated this species and recorded it from late Eocene

to early Miocene rocks in New Zealand. Procardia dolicha

differs from P. inouei new species by having a larger shell

(55 mm in length), more numerous radial ribs (‘ 25 to 28)

and a narrower flat area with a more distinctly demarcated

lunule.

Etymology: The new species is named for Kiyokazu

Inoue (Obihiro City), the collector of the holotype.

Distribution: Known only from the type locality, in the

upper Selandian to lowest Thanetian Katsuhira Forma-

tion, Urahoro Town, eastern Hokkaido.

DISCUSSION

This is the first record of Procardia in the Paleocene and

from the northern Pacific region. As already mentioned

by Beu and Maxwell (1990), hfs genus survived the end-

Cretaceous mass extinction in New Zealand. Nine

species or genera from the Katsuhira Formation were

able to survive this event, partly because they lived in

dee p water (Amano et al., 2018). Procardia inouei new

species is added to the relict forms from the Mesozoic

fauna.

The oldest Procardia sp. is recorded from an Upper

Jurassic carbonate with siliceous sponges in southern

1871), the type species of

Spain (Ol6riz et al., 2003). In the Cretaceous, species of

Procardia are widespread in Europe and US Interior Sea

Way (Woods, 1909; Jablonski and Raup, 1999). How-

ever, there is no record of this genus from the Creta-

ceous in Japan (Nagao, 1943; Hayami, 1975; Tanaka and

Toshimitsu, 2003). The migration route of this genus to

Japan during the Paleocene is uncertain.

On the other hand, the present-day genus Panacca has a

world-wide distribution in warm and relatively deep

waters, but has no fossil record (Coan, 2000; Krylova,

2006: Sasaki and Okutani, 2007). The New Zealand relict

species Procardia dolicha (Suter, 1917) is unusual

having a rather elongate shell similar to Panacca. How-

ever, the presence of radial ribs on whole surface and a

lunule, which are characters of Procardia, were described

and illustrated in P. dolicha by Marwick (1944) and Beu

and Maxwell (1990). It is uncertain, but the elongate

shape of P. dolicha suggests the possibility that Panacca

evolved from Procardia.

ACKNOWLEDGMENTS

I acknowledge Alan Beu (GNS Science) and Eugene V.

Coan (Santa Barbara Museum of Natural History) for

reviewing the manuscript. We also express many thanks to

Kiy koma Inoue (Obihoro in eastern Hokkaido) who

collected the holotype and kindly donated the specimen.

This study was supported by a Grant-in-aid for Scientific

Research from the Japan Society for Promotion of Science

(C, 17K05691, 2017-2019).

LITERATURE CITED

Amano, K. and R.G. Jenkins. 2014. A new Paleocene species of

Aporrhaidae (Gastropoda) from eastern Hokkaido, Japan.

Paleontological Research 18: 33-39.

Amano, K. and R.G. Jenkins. 2017. Paleocene protobranch bi-

valves from Urahoro Town in eastern Hokkaido, Northem

Japan. Journal of Conchology 42: 283-297.

K. Amano, 2019

Page 25

Amano, K., R.G. Jenkins, and H. Kurita. 2018. New and

Mesozoic-relict mollusks from Paleocene wood-fall com-

munities in Urahoro Town, eastern Hokkaido, northern

Japan. Journal of Paleontology 22: 105-111.

Beu, A.G. and P.A. Maxwell. 2000. Cenozoic Mollusca of New

Zealand. Drawings by R.C. Brazier. New Zealand Geo-

logical Survey Paleontological Bulletin 58: 1-518.

Campbe Ml, Vel and J.A. Gran Mackie. 1995. Jurassic Pholado-

myidae (Bivalvia) from New Zealand and New Caledonia.

New Zealand Journal of Geology and Geophysics 38: 47-59.

Coan, E.V. 2000. A new species of pannnen from Chile (Bivalvia:

Pholadomyoidea: Paralimyidae). Malacologia 42: 165-170.

Cox, L.R. and N.D. Newell. 1969. Family Pivelhvlonadhre @ aray,

1847. In: Cox, L. R. et al., eds., Treatise on Invertebrate

Paleontology, Part N (Mollusca 6) (2): 491-952. Geological

Society of nenoren and University of Kansas, Laymenee.

Dall, W.H. 1889. On the hinge of pelecypods and its devel-

opment, with an attempt toward a better subdivision of the

group. American Journal of Science 38: 445-462.

Dall, W.H. 1905. Note on the genus Aporema Dall. The Nautilus

18: a

Dreyer, G. Steiner, and E.M. Harper. 2003. Molecular

co es of Anomalodesmata (Mollusca: Bivalvia) infer-

red from 18S rRNA sequences. Zoological Journal of the

Linnean Society 139: 229-246.

Habe, T. 1977. Sy. stematics of Mollusca in Japan. pe and

canes Zukan-no-Hokuryukan, Tokyo, 372 pp. (In

Japanese. )

Harper E.M., H. Dreyer and G. Steiner. 2006. Reconstructing the

Anomalodesmata (Mollusca: Bivalvia): morphology and mol-

ecules. Zoological Journal of the Linnean Society 148: 395-420.

Hayami, I. 1975. A systematic survey of the Mesozoic Bivalvia

from Japan. Bulletin of the University Museum, the Uni-

versity of Tokyo 10: 1-249.

Huber, M. 2010. Compendium of bivalves. Conchbooks,

Harxheim, 901 pp.

Jablonski, D. and D.M. Raup. 1999. Unpublished data on

Cretaceous—Paleocene molluscan assemblages. http://

fossilworks.org/?a=displayReference&reference_no=40.

King, W. 1844. On a new genus of Palaeozoic shells. Annals and

Magazine of Natural ‘History 147 Bl3—317.

Krylova, im M. 2006. Bivalves of seamounts of the north-eastern

Atlantic. Part I. In: Mironov, A.N., A.V. Gebruk, and

A.J. Southward eds., Biogeography of the North Atlantic

Seamounts: 76-95, KMK Scientific Press, Moscow.

Linnaeus, C. 1758. Systema Naturae (tenth edition). $24 pp.,

Holmiae, Laurentii Salvii.

Locard, A. 1898. Mollusques testacés. vol. 2.515 pp. Expéditions

scientifiques du Travailleur et du Talisman pendant les

années 1880, 1881, 1882, 1883. Masson, Paris.

Mantell, G. 1822. The fossils of the South Downs: or illustrations

of the geology of Sussex. 323 pp., Lupton Relfe, London.

Marwick, J. 1944. New Zealand fossil and Recent Cardiidae

(Mollusca). Transactions of the Royal Society of New

Zealand 74: 255-272.

Matsukuma, A. 1989. Studies on the Kawamura Collection

(Mollusca) stored in the National Science Museum, Tokyo

-VI. Living Pholadomyid bivalves from the northwestern

Pacific, wah de scription of a new species. Venus 48:

207-221.

Meek, F.B. 1871. Descriptions of new species of fossils from

Ohio and other western states and territories. Proceedings

of the Academy of natural Sciences of Philadelphia, for

1871, 23: 159-184.

Meek, F.B. 1876. A report on the invertebrate Cretaceous and

Tertiary fossils of the upper Missouri Country. Report of

the United States Geological Survey of the Territories 9:

1-629.

Melvill, J.C. and R. Standen. 1899. Report on the marine Mollusca

obtained during the first expedition of Prof. A. C. Haddon

to the Torres Straits, in 1888-1889. Journal of the Linnean

Society of London, Zoology 2 27: 150-206.

Morton, B. 1981. The Avnonnellorls esmata. Malacologia 21: 35-60.

Morton, B. 1982. The functional morphology of Parilimya

fragilis (Bivalvia: Parilimyidae nov. fam.) with a discussion

on the origin and ev olution of the carnivorous se ptibranchs

anda re dhastRextion of the Anomalodesmata. Transactions

of Zoological Society of London 36: 153-216.

Nagao, T. 1943. Pholadomya from Japan. Journal of the Geo-

logical Society of Japan 50: 153-160.

Olan, T. 2017. Anomalodesmata. In: Okutani, T. ed., Marine

Mollusks in Japan, second edition: 1206-1217. Tokai Uni-

versity Press, Hiratsuka. (in Japanese and English)

Ol6riz, F., M. Leolid, and F.]. Rodriguez-Tovar. 2003. A Late

Jurassic carbonate ramp colonized by sponges and benthic

microbial communities (External Prebetic, Southern

Spain). Palaios 18: 528-545.

Runnegar, B. 1974. Evolutionary history of the bivalve class

Anomalodesmata. Journal of Paleontology 48: 904-939.

Sasaki, T. and T. Okutani. 2007. A new Spe cies of Panacca

(Bivalvia: Pholadomyoidea: Parilimyidae) from Kanesunose

Bank, off Central Japan. Venus 65: 372-374.

Sowerby, G.B., I. 1823. [Genus Pholadomya]. The genera of

Recent and fossil shells, for the use of students in con-

chology and geology, with original plates 1: 2 pp., Sowerby,

London.

Suter, H. 1917. Description of new Tertiary Mollusca occurring

in New Zealand, accompanied by a few notes on necessary

changes in nomenclature. Part 1. New Zealand Ge ological

Survey Palaeontological Bulletin 5: 1-93.

Tanaka, H. and S. Toshimitsu. 2003. Cretaceous Bivalvia. In:

Ikeya, N., H. Hirano and kK. Ogasawara eds., The database

of Japane se fossil type specimens described during the qth

Century (Part 3). Palaeontological Society of Japan, Special

Papers 41: 192-276.

Woods, H.M.A. 1909. Solenidae, Saxicavidae, Teredinidae,

Anatinidae, Pholadomyidae, Pleuromyidae, Poromyidae, and

Cuspidariidae. A Monograph of the Cretaceous Lamelli-

branchia of England, vol. 2, part 6: 217-260, Palaeonto-

graphical Society, London.

THE NAUTILUS 133(1):26-30, 2019

Page 26

A large new Wareniconcha (Bivalvia: Vesicomyidae) from a Pliocene

methane seep deposit in Leyte, Philippines

Tomoki Kase

National Museum of

Nature and Science, Tokyo

Department of Geology

and Paleontology

Tsukuba, Ibaraki 305-0005, JAPAN

[email protected]

Shinji Isaji

and Institute, Chiba

[email protected]

Natural History Museum

Chiba 260-8682, JAPAN

Steffen Kiel

Swedish Museum of

Natural History

Department of Palaeobiology

10405 Stockholm, SWEDEN

[email protected]

Yolanda M. Aguilar

Mines and Geosciences

Bureau

Marine Geological

Survey Division

Quezon City, PHILIPPINES

[email protected]

ABSTRACT

A new species of the genus Wareniconcha, W. mercenarioides,

belonging to the che aicembiene bivalve subfamily Pliocardiinae

(family V jesicomyidae), is described from a Pliocene methane-see p

deposit at Liog- Liog on Leyte Island, Philippines. With a length of

almost 12 cm, this species is significantly larger than the six extant

species currently considered as be longing to Wareniconcha. In

addition to being very large, W. me ween naiae >s is more inflated and

has aconside rably more rounds d shell outline compared to the oval

shells of other Wareniconcha species. This is the first fossil record

of Wareniconcha. Considering the overall similarity of the Leyte

seep fauna to species living at vents and seeps in the vicinity of

southem Japan, we anticipate that similarly large, and close ly re-

lated species, might still be extant in the Indo-West Pacific region.

Additional Keywords: Indo-Pacific Ocean, hydrothermal vents,

shell size, deep-sea

INTRODUCTION

Vesicomyid bivalves are a major component of faunal

communities around many deep-sea hydrothermal vents,

methane-seeps, and sunken whale carcasses (Boss and

Turner, 1980; Paull et al., 1954; Smith et al., 1989; Kojima

et al., 2004; Krylova and Sahling, 2010; Johnson et al.,

2017). They thrive in these environments because they

live in sy mbiosis with sulfur -oxidizing bacteria from which

they obtain most, if not all, of their nutrients (Arp et al.,

1984: Fisher, 1990). In addition to being abundant,

several vesicomyid species reach very large sizes, such as

‘Ec tenagena exte nta from methane- -seeps in Monterey

Canyon (up to 24.5 cm; Krylova and Moskalev, 1996), the

iconic ‘Calyptogena’ magnifica from vents on the East

Pacific Rise and Galapagos Ridge that reaches 26.3 cm in

length (Boss and Turmer 1980), rand Abyssogena novacula

anew a, Sahling, and Janssen, 2010 reaching 27.7 cm in

length (Krylova et al. 2010).

In comparison to these large species, members of the

vesicomyid genus Wareniconcha reach only moderate

sizes of up to 7.6 cm, as in the case of the type species

Wareniconcha guineensis (Thiele and Jaeckel, 1931) from

methane seeps in the eastern Atlantic Ocean. Other

species subsequently considered as belonging to Ware-

niconcha reach similar sizes (e.g., both W. winckworthi

(Prashad, 1932) and W. solidissima (Prashad, 1932) from

the Bali Sea in Indonesia reach about 7 cm), whereas

others are somewhat smaller (e.g., W. ovalis (Dall 1895)

from the eastern Pacific, and W. cretacea (Smith, 1906)

from West of Sri Lanka in the Indian Ocean, reach 5—6 cm

in length). Here we describe a new species of Ware-

niconcha from Pliocene seep deposits in Leyte Island,

Philippines that reaches nearly 12 cm in length.

Chemosymbiotic bivalves of the mytilid subfamily

Bathymodiolinae show a remarkable increase in size

through geologic time that can be linked to the acquisition

of diftorent types of symbiotic associations (Lorion et t all,

2013). In addition, vesicomyid bivalves showed an in-

crease in shell size since their first appearance in the

middle Eocene (Kanie and Kuramochi, 2001; Amano and

Kiel, 2007, 2011; Amano et al., 2014). Although ves-

icomyid bivalves do not show the diversity of symbiotic

associations as bathymodiolins do, vesicomyid clades show

a diversity of sulfide- binding affinities that enable (or

restrict) them to colonize dlistitaet niches within methane

seep sites related to sulfide flux and availability (Barry and

Kochevar, 1998; Goffredi and Barry, 2002; Decker et al.,

2014). Future studies of shell size and sulfide-binding

affinities could provide new insights into the long-term

evolution of these traits through Earth’s history.

MATERIALS AND METHODS

Several methane-seep deposits have been discovered in

deep-water strata of the Visayan back-are basin along the

northwestern coast of the island of Leyte in the Philippines

(Majima et al., 2007; Majima et al., 2010). These massive

mudstones were mapped as Bata Formation and initially

considered late Miocene in age (Corby and al., 1951; Porth

et al., 1989: Mines and Geosciences Bureau, 2010). The

T. Kase et al., 2019

Page 27

[o)

Figures 1-5. Wareniconcha mercenarioides new species; holotype (NMNS PM 28168). 1. Left valve. 2. Right valve. 3. Dorsal

view. 4. Hinge and ligament nymph of left valve, micro-CT scan. 5. Hinge and ligament nymph of right valve, micro-CT scan.

specimens reported here were obtained from a giant cal-

careous concretion packed with chemosynthetic bivalves,

found exposed at Liog-Liog Point between Tabango and

Campopo bays, ate IIe 17 977 FN 24c9 55 (Majima

et al., 2007). Ongoing stratigraphic and micropaleontologic

work indicates that the Bata Formation exposed around the

Liog-Liog Point ranges in age from late Pliocene to early

Pleistocene, and the giant concretion is likely to have been

derived from the upper Pliocene part of the Bata For-

mation. The type material is deposited at the National

Museum of Nature and Science, Tsukuba, Japan (NMNS

PM 28168, PM 28169) and the National Museum, Manila,

Philippines (NM P-2148).

To observe the hinge structure, we performed non-

destructive analysis using an X-ray microfocus CT system

(TESCO TXS320-ACTIS) at the National Museum of

Nature and Science, Tokyo, at experimental conditions of

247 kV and 240 pA. The resolution of the square slice CT

was 97 mm per 1024 pixels, and the spacing between each

CT slice was 0.1 mm. Analysis and surface rendering were

performed using the software OsiriX version 3.9.2 32-bit

(an open-source DICOM viewer for Macintosh).

SYSTEMATICS

Family Vesicomyidae Dall and Simpson, 1901

Genus Wareniconcha Cosel and Olu, 2009

Type Species: Vesicomya guineensis Thiele and

Jaeckel, 1931; Recent, from ca. 2500 to 4000 m depth

on the West African continental margin (Cosel and Olu,

2009).

Remarks: Cosel and Olu (2009) included only the type

species in Wareniconcha. In a subsequent review of

vesicomyid species, Krylova and Sahling (2010) con-

sidered also the following species as belonging to

Wareniconcha: Vesicomya compressa Prashad, 1932,

Vesicomya cretacea Smith, 1906, Vesicomya lepta Dall,

age I

Page 28

THE NAUTILUS, Vol. 133, No. 1

Figures 6-9. Wareniconcha mercenarioides new species; paratype | (NMP-2145). 6. Left valve. 7. View on posterodorsal margin.

8. Right valve. 9. View on anterior side.

1895, Vesicomya ovalis Dall, 1895, and Vesicomya

winckworthi Prashad, 1932.

Wareniconcha mercenarioides new species

(Figures 1-9)

Diagnosis: Large and inflated Wareniconcha species that

is about as high as wide, with a broad ligament nymph

plate.

Description: Large, inflated shell, up to 12 cm long,

slightly longer dhe high; umbones large, blunt, strongly

prosogyrate; shell suninee sculpture by fine, re gular ¢ ono

increments; anterior part short, somewhat pointed: post-

erodorsal margin broadly and evenly rounded, with angular

transition Folie equally broadly and eve nly rounded ventral

margin. Right valve with strong cardinal 1 radiating ante-

riorly, positioned anterior of Senet cardinal 3a thin and

short, positioned just anterior of umbo, cardinal 3b short,

moderately strong, pointing posteriorly. Left valve with

strong and elongate cardinal 2a subparallel to dorsal

margin, cardinal 2b strong, quadrate, just beneath umbo,

cardinal 4 thin, elongate, pointing posteriorly; nymph plate

long and broad.

Type Locality: The Liog-Liog seep deposit.

Type Material: Holotype: NMNS PM 28168, articu-

lated specimen (length 11.3 em, height 9.7 cm, width

6.5 cm); Paratype 1: NMP-2148, pe eulated specimen

(length 11.3 cm, height 10.1 cm, width 5.7 cm); Paratype

2: NMNS PM28169, disarticulated specimen (length ca.

12 cm, height, ca. 10 cm) consisting of incomplete right

valve.

Distribution and Habitat: Pliocene methane-seep

carbonates at Liog-Liog Point, Tabango municipality in

Leyte, Philippines.

T. Kase et al., 2019

Page 29

Etymology: For its shell shape resembling the venerid

genus Mercenaria.

Remarks: Wareniconcha mercenarioides differs from all

species assigned to Wareniconcha (see list above in the

remarks about Wareniconcha.) by being larger, more

inflated, and being roughly as wide as high, whereas all

other Wareniconcha species are oval in an anterior-

posterior direction. Furthermore, W. guineensis, W.

compressa, and W. lepta have a narrower hinge plate than

W. mercenarioides.

Many of the accompanying bivalve species at the type

locality belong, or are very similar to, extant species from

vents and se eps in Japanese waters and especially in the

Okinawa Trough (including, for example, Bathymodiolus

securiformis (Okutani et al., 2004), Archivesica soyoae

(Okutani, 1957), Archivesica similaris (Okutani et al.,

1997), Archivesica kawamurai (Kuroda, 1943), and

Pliocardia’ kuroshimana (Okutani et al., 2000); TK and SK,

unpublished). Therefore, we anticipate that Wareniconcha

mercenarioides or a very closely related species may still be

extant somewhere around the Indo-West Pacific.

ACKNOWLEDGMENTS

We would like to thank L. L. Jasareno (Director, Mines

and Geosciences Bureau, Philippines) for a research

permit, W. Mago, F. Kanoda, and E. Azurin (Mines and

Geosciences Bureau of the Philippines) for their assis-

tance in fieldwork, and H. Hayashi (Shimane University),

A. Gil S. Fernando (the University of Philippines) for the

microfossil analyses, C.

Nature and Science, Tsukuba) for running the CT scans,

and Elena Krylova (P.P. Shirshov Institute of Oceanology,

Moscow) for her kind review of the manuscript. imancrall

support to TK was provided by the Japan Society for the

Promotion of Science, Japan through Grant-in-Aid for

Scientific Research 16K05600, and to SK by the Swedish

Science Foundation (Vetenskapsradet) through grant

2016-03920.

LITERATURE CITED

Amano, K., R.G. Jenkins, M. Ohara, and S. Kiel. 2014. Miocene

vesicomyid species (Bivalvia) from Wakayama in southern

Honshu, Japan. The Nautilus 128: 9-17.

Amano, K. and S. Kiel. 2007. Fossil vesicomyid bivalves from the

North Pacific region. The Veliger 49: 270-293.

Amano, K. and S. Kiel. 2011. Fossil Adulomya (Vesicomyidae,

Bivalvia) from Japan. The Veliger 51: 76-90.

Arp, A.J., J.J. Childress, and C.R. Fisher. 1984. Metabolic and

blood gas transport characteristics of the hydrothermal vent

biv alve, Calyptogena magnifica. Physiological Zoology 57:

648-662.

Barry, J.P., and R.E. Kochevar. 1998. A tale of two clams:

differing chemosynthetic life styles among vesicomyids in

Monterey Bay cold seeps. Cahiers de Biologie Marine 39:

329-331.

Sakata (National Museum of

Boss, K.J. and R.D. Turner. 1980. The giant white clam from the

Galapagos Rift, Calyptogena magnifica species novum.

eee 20: 161-194.

Corby, G.W. et. al. 1951. Geology and oil possibilities of the

Philippine s. Republic of the Philippine s, Department of Ag-

riculture and Natural Resources, Technical Bulletin 21: 1-363.

Cosel, R.v., and kK. Olu. 2009. Large Vesicomyidae (Mollusca:

Bivalvia) from cold seeps in fhe Gulf of Guinea off the

coasts of Gabon, Congo and northern Angola. Deep-Sea

Research II 56; 2350-2379.

Dall, W.H. 1895. Diagnoses of new species of mollusks from the

west coast of America. Proceedings of the U.S. National

Museum of Natural History 18: 7-20.

Decker, C., N. Zorn, N. Potier, E. Leize-Wagner, K. Olu, and

A.C. Andersen. 2014. Globin’s structure and function in

vesicomyid bivalves from the Gulf of Guinea cold seeps as

an adaptation to life in reduced sediments. Physiological

and Biochemical Zoology 87: 855-869.

Fisher, C.R. 1990. Chemoautotrophic and methanotrophic

symbioses in marine invertebrates. Reviews in Aquatic

Sciences 2: 399-436.

Goffredi, S.K. and J. P. Barry. 2002. Species-specific variation in

sulfide physiology between closely related Vesicomyid

clams. Marine Ecology ee Series 255: 227-238.

Johnson, S.B., E.M. Krylova, A. Audzijonyte, H. Sahling, and

R.C. Vrijenhoek. 2017. Phylogeny and origins of chemo-

synthetic vesicomyid clans. Systematics and Biodiversity

15: 346-360.

Kanie, Y. and T. Kuramochi. 2001. Two new species of the

Vesicomyidae (Bivalvia: Mollusca) from the Pliocene

Shiramazu Formation of the Chikura Group in the Boso

Peninsula, Japan. Science Report of the Yokosuka City

Museum 48: 1-9.

Kojima, S., K. Fujikura, and T. Okutani. 2004. Multiple trans-

Pacific migrations of deep-sea vent/seep-endemic bivalves

in the family Vesicomyidae. Molecular Phylogenetics and

Evolution 32: 396-406.

Krylova, E.M. and L.I. Moskalev. 1996. Ectenagena extenta, a

new species of vesicomyid bivalve from Monterey Bay,

California. Ruthenica 6: 1—10.

Krylova, E.M. and H. Sahling. 2010. Vesicomyidae (Bivalvia):

current taxonomy and distribution. PLoS ONE 5: e9957.

Krylova, E.M., H. Sahling, and R. Janssen. 2010. Abyssogena: a

new genus of the family Vesicomyidae (Bivalvia) from deep

water vents and seeps. Journal ‘of Molluscan Studies 76:

107-132.

Kuroda, T. 1943. Akebiconcha, a new pelecypod genus. Venus

13: 14-18.

Lorion, J., S. Kiel, B.M. Faure, M. Kawato, S.Y.W. Ho, B.A.

Marshall, S. Tsuchida, J.-I. Miyazaki, and Y. Fujiwara. 2013.

Adaptive radiation of che mosymbiotic deep-sea mussels.

Proceedings of the Royal Society B 280: doi.org/10.1098/

rspb.201 3.1243

Majima, R., R.G. Jenkins, T. Kase, Y.M. Aguilar, T. Nanjo, R.

Wani, H. Wada, A.G.S. Fernando, and H. Hayashi. 2010. In

situ Calyptogena colonies from Pliocene back-arc basin fills

in Leyte Island, Philippines. Journal of the Geological

Society of Jee 116: XV-XVI.

Majima, R., T. Kase, S. Kawagata, Y.M. Aguilar, K. Hagino, and

M. Maeda. 2007. Fossil eal seep assemblages from I seyte

Island, Philippines. Journal of Geography 116: 643— 652

(with pictorial figures).

Mines and Geosciences Bureau. 2010. Geology of the Philip-

pines. Second Edition. Department of Environment a

Page 30

THE NAUTILUS, Vol. 133, No. 1

Natural Resources, Mines and Geosciences Bureau,

Quezon City, 532 pp.

Okutani, T. 1957. Two new species of bivalves from the deep

water in Sagami Bay collected by the R.V. Soyo-Maru.

Bulletin of ‘alle Regional Fisheries Research Laboratory

17: 27-31.

Okutani, T., K. Fujikura, and S. Kojima. 2000. New taxa and

review “at vesicomyid bivalves collected from the Northwest

Pacific by deep sea research systems of Japan Marine

Science & Technology Center. Venus 59: 83-101.

Okutani, T., K. Fujikura, and T. Sasaki. 2004. Two new species of

Bathymodiolus (Bivalvia: Mytilidae) from methane seeps on

the Kuroshima Knoll off the Yaeyama Island, southwestern

Japan. Venus 62: 97-110.

Okutani, T., S. Kojima, and J. Ashi. 1997. Further discovery of a

new taxon of vesicomyid clam from the Nankai Trough, off

Honshu, Japan. Venus 56: 185-188.

Paull, C.K., B. Hecker, R. Commeau, R.P. Freeman-Lynde,

Neumann, W.P. Corso, S. Golubic, ].E. Hook, E. Sikes, es

Notice

J. Curray. 1984. Biological communities at the Florida

Escarpment resemble hydrothermal vent taxa. Science 226:

965-967.

Porth, H., C. Miller, and C.H. von Daniel. 1989. The sedi-

mentary formations of the Visayan Basin, Philippines.

Geologisches Jahrbuch B70: 29-87.

Prashad, B. 1932. The Lamellibranchia of the Siboga expedition.

Systematic part II (exclusive of the eratinantcle)). Siboga

Expeditie 53¢c: 1-351.

Smith, C.R., H. Kukert, R.A. Wheatcroft, P.A. Jumars, and J.W.

Deming. 1989. Vent fauna on whale remains. Nature 341:

27-28.

Smith, E.A. 1906. Natural history notes from R.I.M.S. ‘In-

vestigator. -Series II., No.10. On Mollusca from the Bay of

Bengal and the Arabian Sea. Annals and Magazine of

Natural eee 18: 157-175 and 245-264.

Thiele, J. and S. Jaeckel. 1931. Muscheln der Deutschen

Tiefsee-Expedition. II Teil. Deutsche Tiefsee-Expedition

1898-1899 21: 159-268.

THE 2019 R. TUCKER ABBOTT VISITING CURATORSHIP

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