* HE NAUTILUS

Volume 134, Numbers 3-4

December 18, 2020

ISSN 0028-1344

A quarterly devoted

to malacology.

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pti A poctenet =

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Volume 134, Number 3-4

December 18, 2020

ISSN 0028-1544

CONTENTS

Leonardo S. Souza A new species of Tritonoharpa Dall, 1908 (Gastropoda: Cancellariidae)

Renata S. Gomes from the southwestern Atlantic and an overview of other western Atlantic

Paulo Marcio S. Costa SPA Sc oo ei cre one ican ge oy green Sassie a ar Coe a as aes aa ea 95

Charles L. Powell, II Memorial to Warren O. Addicott—A remarkable student of Tertiary marine

Kevin J. Meade taollusiks. a1 the WOTNeAStern PACinG ..... sarki aiccsek anattanaaes edt eer ane 107

Shawn G. Wiedrick Six new species of Paciocinebrina (Gastropoda: Muricidae: Ocenebrinae)

Roland Houart IEC COPR ALTER CTCTO EL TSO lOO] OF) d/h] SURE aa oNGt Ate ne ENTE ns ote Retry Cre rege ae A wi

Kazutaka Amano A new species of Arca (Bivalvia: Arcidae) from the lower Miocene Asahi

Hiroshi Kurita Formation on the Japan Sea side of central Honshu, with remarks on the

westward faunal migration from the eastern Pacific .........cccceeeteeeteteeeneeee 132

RESEARCH NOTE

Justin N. Voss Imposex in the city: First evidence of female masculinization in Queen

Gabriel A. Delgado Conch Aliger gigas (Linnaeus, 1758) (Gastropoda: Strombidae) in Florida .... 138

Nancy J. Brown-Peterson

Author Tradex...230.2 ican Soak Peved ok oi Berries Se I eae ee ee ee nf ID Pe ick 143

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THE NAUTILUS 134(3-4):95-106, 2020

Page 95

A new species of Tritonoharpa Dall, 1908 (Gastropoda:

Cancellariidae) from the southwestern Atlantic and an overview of

other western Atlantic species

Leonardo S. Souza Renata S. Gomes

Malacologia, Departamento de

Invertebrados and

Museu Nacional, Rio de Janeiro,

BRAZIL

[email protected]

Instituto Mar Adentro, Rio de Janeiro, Brazil

Malacologia, Departamento de Invertebrados

Museu Nacional, Rio de Janeiro, BRAZIL

Paulo Marcio S. Costa

Fundacao Instituto de Pesca do Estado

do Rio de Janeiro

Escritério Regional Costa Verde, Angra

dos Reis, Brazil

and

Malacologia, Departamento de Invertebrados

Museu Nacional, Rio de Janeiro, BRAZIL

ABSTRACT

A new southwestern Atlantic species belonging to genus Tri-

tonoharpa is described from northeastern and southeastern

Brazil. Tritonoharpa curvapex new species has a strongly

deviated nucleus in the protoconch, which is the main feature

that distinguishes it from the remaining congeners from the

western Atlantic. Shell dimensions and number of cords in the

teleoconch also distinguish this new species from the other

Tritonoharpa. An overview of the remaining extant Tritono-

harpa from the western Atlantic is also presented. We provide

taxonomical remarks, illustrations and the known distribution of

Tritonoharpa lanceolata (Menke, 1828), Tritonoharpa cuba-

patriae (Sarastia, 1975), Tritonoharpa bayeri (Petuch, 1987),

Tritonoharpa leali Harasewych, Petit, and Verhecken, 1992 and

Tritonoharpa janowskyi Petuch and Sargent, 2011.

Additional Keywords: Neogastropoda, Volutoidea, Plesio-

tritoninae, taxonomy

INTRODUCTION

The family Cancellariidae Forbes and Hanley, 1851 consists

of marine gastropods and is known from all oceans (Lima

et al., 2007), but the majority of the species are tropical or

temperate (Harasewych and Petit, 2011). This family cur-

rently comprises about 350 extant species (MolluscaBase,

2020) and presents a great variety of shell features (Modica

et al., 2011b). Most recent classifications include three

subfamilies: Cancellariinae Forbes and Hanley, 1851,

Admetinae Troschel, 1865 and Plesiotritoninae Beu and

Maxwell, 1987 (Modica et al., 2011b; Bouchet et al., 2017).

Among the Plesiotritoninae, the genus Tritonoharpa

Dall, 1908 is one of the most diverse with 19 extant valid

species (MolluscaBase, 2018). Five species of Tritono-

harpa are known from the western Atlantic: T. lanceolata

(Menke, 1828), T. cubapatriae (Sarastia, 1975), T. bayeri

(Petuch, 1987), T. leali Harasewych, Petit and Verhecken,

1992 and T. janowskyi Petuch and Sargent, 2011. Tri-

tonoharpa lanceolata is currently known from several

localities in the western Atlantic, occurring from the

eastern USA to southeastern Brazil (Beu and Maxwell,

1987; Rios, 2009). Tritonoharpa cubapatriae is known

from Cuba and Panama (Caribbean coast) (Sarastia, 1975;

Lee, 2009). Tritonoharpa bayeri and T. janowskyi are

known only from their type localities off Caribbean

Colombia and Florida, USA, respectively. Tritonoharpa

leali is currently restricted to the states of Bahia, Espirito

Santo, and Rio de Janeiro, Brazil (Harasewych et al., 1992:

Absalao et al., 2006). Thus, until now T. lanceolata and T.

leali are the only species recorded in Brazil (Harasewych

et al., 1992: Absalao et al., 2006; Rios, 2009).

Verhecken (1991) commented on the scarcity of rec-

ords or descriptions of Cancellariidae in the western

Atlantic south of the Equator, but since then reports on

cancellariids from Brazil increased considerably, espe-

cially in deep-sea studies (Harasewych et al., 1992;

Simone and Birman, 2006; Barros and Lima, 2007; Barros

and Petit, 2007; Lima et al., 2007). Recent surveys of

malacological collections in Brazil revealed an unde-

scribed species of Tritonoharpa from the coast of Brazil

and the need for a revision of other known congeners in

the western Atlantic. We formally describe Tritonoharpa

curvapex new species and provide taxonomical com-

ments on the other western Atlantic Tritonoharpa.

MATERIALS AND METHODS

The material studied consists mostly of empty shells

deposited in malacological collections, collected by vari-

ous oceanographic expeditions and smaller collecting

events. The largest expeditions are: 1. REVIZEE Central:

“Programa de Avaliagéo do Potencial Sustentavel de

Recursos Vivos da Zona Econémica Exclusiva, Score

Central” (Program of Evaluation of the Sustainable

Page 96

Potential of Living Resources in the Economic Exclusive

Zone), carried out by “Ministério do Meio Ambiente”,

Brazilian government, samples collected by the research

vessel (R/V) ANTARES and the supply-boat AsTRO GAROUPA

between 1996-2002; 2. BPOT MR: “Projeto de Caracter-

izagao e Monitoramento Ambiental da Bacia Potiguar —

Malha Regional” (Project of Environmental Characterization

and Monitoring of Potiguar Basin — Regional Grid), carried

out by Petrobras SA, samples collected by the supply-

boat AsTRO Garoupa between 2002-2004; 3. AMBES:

“Caracterizagaéo Ambiental Marinha da Bacia do Espirito

Santo e porcaéo Norte da Bacia de Campos” (Marine envi-

ronmental characterization of the Espirito Santo Basin and

North part of Campos Basin), carried out by Petrobras SA,

samples collected by the R/V LUKE THoMas and R/V SEwarD

JouNson, between 2010-2013.

Most shells were photographed (using focus stacking)

by an AxioCam ICc 5 camera coupled to a stereomi-

croscope ZEISS Discovery.V20. A few shells were studied

under a scanning electron microscope (SEM) Jeol JSM-

6390LV for a detailed examination of sculpture. Shell

measurements are: shell length (SL), body whorl length

(BWL), aperture length (AL), shell width (SW), aperture

width (AW), protoconch height (PCH) and width (PCW).

The counting of protoconch whorls follows Verduin

(1982) and Verhecken (2007). Some shells were treated

with mineral oil to enhance the color pattern.

Collection acronyms are: ANSP: Academy of Natural

Sciences of Philadelphia, Drexel University, Philadelphia;

FLMNH: Florida Museum of Natural History, Gaines-

ville; IBUFRJ: Instituto de Biologia, Universidade Fed-

eral do Rio de Janeiro, Rio de Janeiro, Brazil; MNRJ:

Museu Nacional, Rio de Janeiro, Brazil; MORG: Museu

Oceanografico “Professor Eliézer de Carvalho Rios”,

Universidade Federal do Rio Grande, Rio Grande, Brazil:

MZSP: Museu de Zoologia, Universidade de Sao Paulo,

Sao Paulo, Brazil; SMF: Naturmuseum Senckenberg,

Frankfurt, Germany; USNM: National Museum of Nat-

ural History, Smithsonian Institution, Washington, DC.

Due to the recent fire at the Museu Nacional in Brazil

(Zamudio et al., 2018) most lots from this institution listed

in the material examined here, in addition to lots lent from

other institutions, were destroyed. Destroyed specimens

(spm) or shells (sh) are marked by a dagger (f) in the

material examined of each species.

SYSTEMATICS

Family Cancellariidae Forbes and Hanley, 1851

Subfamily Plesiotritoninae Beu and Maxwell, 1987

Genus Tritonoharpa Dall, 1908

Nivitriton Iredale, 1929. Type species: Triton antiquatus

Hinds in Reeve, 1844 (by original designation); Recent,

tropical Indo-Pacific.

Esbelta Sarasia, 1975. Type species: Ranella lanceolata

Menke, 1828 (by original designation); Miocene to Re-

cent, western Atlantic.

THE NAUTILUS, Vol. 134, Nos. 3-4

Type Species: Tritonoharpa vexillata Dall, 1908 (by

original designation); Recent, western America and the

Galapagos Island.

Description: Shell with weakly to moderately convex tel-

eoconch whorls, retaining prominent varices on early as well

as later whorls, well developed columellar collar, lacking

columellar plaits or with a single very low, narrow columellar

plait, and with a nematoglossan radula, consisting of a thin

membrane and one central row of rachidian teeth (adapted

from Beu and Maxwell, 1987 and Modica et al., 2009).

Tritonoharpa curvapex new species

(Figures 1-9)

Diagnosis: Shell up to 15.0 mm long, 5.5 mm wide, with 6-7

whorls, cream to light brown, with scattered brown blotches

and thin spiral bands. Whorls convex, with moderately

prominent varices (1-2 per whorl) from early whorls to

later whorls, sculptured with cancellate pattern. Protoconch

paucispiral, smooth, 1.0 whorl, nucleus strongly deviated.

Description: Shell about 15.0 mm long, 6.5 mm wide, spire

angle 35°, shouldered. Protoconch paucispiral, 1.0 whorl,

nucleus deviated about 30° from teleoconch axis.

Protoconch-teleoconch transition abrupt, marked by a thin

lip and onset of cancellate sculpture. Teleoconch with up to 7

whorls, moderately inflated, weakly shouldered; suture

deeply impressed, almost obscured by the axial coronation of

succeeding whorl. Spiral and axial cords present, producing

cancellate sculpture; strong nodules at the intersection of

spiral and axial cords; spiral cords more pronounced than axial

cords. Varices high, non-collabral, narrow, appearing peri-

odically in every 2/3 whorl. About 11 axial ribs between

varices in the last whorl; About 16 spiral cords from the suture

to the beginning of the siphonal canal in the last whorl (frontal

view). Aperture wide, elliptical, with well-defined posterior

notch. Inner lip smooth, with columellar collar reflected over,

but not adherent to, fasciole and pseudoumbilicus. Outer lip

with up to 8 pairs of denticles confined to flared region

beyond varix. Siphonal canal short, weakly deviated, partially

covered abaxially. Shell color cream to brownish, with

scattered brownish blotches and thin spiral bands.

Type Locality: Brazil: Northeast coast, imprecise locality

between Pernambuco and Bahia states, coll. 1977.

Type Material: Holotype: IBUFRJ 5430 (sh). Para-

types: Brazil: Bahia state: Camamu Basin, 13°28’29” S

38°48’41” W, 30 m (MNRJ 14925, 1 sht; MNRB]J 27973, 1

sht); Espirito Santo state: REVIZEE Central I stn. C65,

18°52’58" S 39°06’00" W, 50 m, coll. R/V Antares, 25/iv/

1996 (IBUFR] G520; seh)

Measurements: Holotype, IBUFRJ 5430: Whorls= 6.0;

SL= 12.7 mm; BWL= 8.5 mm; AL= 6.0 mm; SW=

9.2 mm; AW= 2.7 mm; PCH= 0.8 mm; PCW= 0.9 mm:

SL/SW= 2.4.

L.S. Souza et al., 2020

Page 97

Figures 1-9. Tritonoharpa curvapex new species. 1-4. Holotype, IBUFRJ 5430. 5. Paratype, MNRJ 27973. 6-9. Paratype,

MNB] 14925. 1, 5-7. Entire shell in frontal view, white square in 7 indicates detail in 8. 2. Entire shell in lateral view. 3. Entire shell in

dorsal view. 4, 9. Detail of protoconch, lateral and apical views, respectively. 8. Detail of teleoconch surface. Scale bars: 1-3, 5-7 =

5 mm; 4 = 500 pm; 8-9 = 200 pm.

Etymology: Curvus, Latin for bent; apex, Latin for tip;

referring to the strongly deviated nucleus of the protoconch.

Distribution: Brazil: From Pernambuco to Espirito

Santo. Empty shells from depths between 30 m to 50 m.

Remarks: The holotype IBUFR] 5430 (Figures 14) and

the paratype IBUFRJ 9525 of T. curvapex are safely

preserved in the IBUFRB]J collection. The other paratypes

(Figures 6-9) were destroyed by the fire. To avoid the

designation of a destroyed specimen as the holotype, the

shell of IBUFRJ 5430 was selected as the holotype despite

the imprecise locality of collection.

Tritonoharpa curvapex differs from all other western

Atlantic species of the genus in having a conspicuous

protoconch in which the nucleus is strongly deviated

(Figures 4, 9). This newly described species is most similar

to T. bayeri (Figures 23-27) based on the general shape and

on having an aperture that is not very constricted posteri-

orly. The protoconch of the holotype of T. bayeri (USNM

Page 98

859853) is partially broken (Figures 25, 26), but it is possible

to note that the nucleus is not deviated as in T. curvapex.

Tritonoharpa curvapex can also be distinguished from T.

bayeri by the relatively smaller dimensions of the shell

(12.7 mm long, 5.2 mm wide vs. 16.2 mm long, 7.6 mm

wide, respectively, both with 5 whorls) and by the smaller

number of axial ribs between varices of the last whorl (11 ws.

16-18). These species also differ by their color pattern, T.

curvapex has brownish spots like T. bayeri, but also some

spiral bands (Figure 5) that are not present in T. bayeri.

Tritonoharpa curvapex also resembles Tritonoharpa

ponderi Beu and Maxwell, 1987, from Australia, and Tri-

tonoharpa caunbonensis Pacaud, Ledon, and Loubry, 2015,

an extinct species from the Eocene of Paris Basin, because

these species also have a strongly deviated protoconch (Beu

and Maxwell, 1987: pl. 18, fig. E, I; Pacaud et al., 2015: pl. 10,

fig. LE). However, T. curvapex can be distinguished from T.

ponderi by the abrupt increase in diameter of the teleoconch,

which occurs more gradually in T. ponderi. Tritonoharpa

curvapex can be distinguished from T. caunbonensis by its

relatively larger size (holotype of T. curvapex, IBUFRJ 5430,

~6 whorls, 12.7 mm long, 5.2 mm wide, SL/SW= 2.4 vs.

holotype of T. caunbonensis, MNHN.F.A51472, ~7 whorls,

10 mm long, 3.9 mm wide, SL/SW= 2.6), and in having

thinner varices and a narrower inner lip.

Notes on Other Western Atlantic Tritonoharpa

Tritonoharpa lanceolata (Menke, 1828)

(Figures 10-20)

Ranella lanceolata Menke, 1828: 87.

For synonymy up to 1987, see Beu and Maxwell (1987: 40)

Colubraria lanceolata (Menke, 1828) — Perry and

Schwengel (1955: 158, pl. 31, fig. 222); Daccarett and

Bossio (2011: 94, fig. 444).

Tritonoharpa lanceolata (Menke, 1828) — Lyons (1989: 26,

pl. 9, fig. 1); Lyons (1998: 27 [annotated list]); Petit and

Harasewych (2005: 63[annotated list]); Wolfe (2008

[annotated list]); Tunnell Jr. et al. (2010: 234); Espinosa

et al. (2012: 291, fig. 525); Lamy and Pointier (2017: 528,

pl. 184, fig. 9A~B).

Type Locality: Puerto Rico (Menke, 1828).

Type Material: Probably lost. After K.T. Menke’s death his

collection was sold and dispersed (Zilch, 1967; Kohn, 1988).

The most probable place to find would be the SMF,

however there is no material that seems to belong to

Menke’s material of T. lanceolata (Sigrid Hof, pers. comm. ).

Other Material Examined: USA: Georgia: off Georgia

(30°54’18” N, 80°36'06” W, 34 m), coll. R/V Bacsy, 26/ii/

1980: USNM 824010 [3 sh]; Florida: 12 miles Northeast

Cape Canaveral (28°37' N, 80°30’ W): USNM 486176 [1

sh]; Oculina Reef (27°50' N, 79°58’ W, 91 m), coll.

Houbrick: USNM 798073 [1 sh]; W of Boca Grande

(26°44’ N, 84°09’ W, 55 m), coll. 1971: ANSP 395042 [6

THE NAUTILUS, Vol. 134, Nos. 3-4

sh]; Off Key West (182-213 m), coll. IX/1963: ANSP

294763 [1 sh]. Caribbean area: Jamaica: Montego Bay,

coll. A. L. Mhering, 17/xii/1954: USNM 712137 [1 sh].

Panama: Minas Bay, Payardi Island: USNM 743644 [2 sh].

Haiti: Jeremy: USNM 383239 [1 sh]. Dominican Re-

public: Las Galeras, Samana Bay (2 m), coll. 1994: ANSP

408384 [2 sh]. Puerto Rico: North of Mayaguez Docks:

USNM 662026 [2 sh]. Virgin Islands: St. Thomas, coll.

Brady, 1968: USNM 702792 [2 sh]. Anguilla: Sombrero

Island: USNM 92993 [1 sh]. Grenada: Grand Anse Bay

(12°01'46” N, 61°46'19” W, 3-6 m), coll. 08/x/2012: MZSP

108664 [1 sh]. Trinidad and Tobago: Off Scarborough,

coll. 30/xi/1989: FLMNH 281379 [7 shf].

Measurements: USNM 92993: Whorls= 8.0; ANSP

395042: SL= 29.1 mm; BWL= 17.0 mm; AL= 11.5 mm;

SW= 9.0 mm; AW= 6.0 mm; PCH= not measured:

PCW= not measured; SL/SW= 3.2. Whorls= 7.5: SL=

24.1 mm; BWL= 14.3 mm; AL= 10.1 mm; SW= 8.6 mm;

AW= 4.8 mm; PCH= 1.1 mm; PCW= 1.2 mm; SL/SW=

2.8. ANSP 294763: Whorls= 7.0; SL= 20.2 mm; BWL=

13.7 mm; AL= 9.7 mm; SW= 7.0 mm; AW= 4.4 mm:

PCH= 0.8 mm; PCW= 1.0 mm; SL/SW= 2.9.

Distribution: USA: New Jersey, North Carolina, Georgia,

Florida (Beu and Maxwell 1987), Texas (Rosenberg et al.,

2009; Tunnell Jr. et al., 2010). Bermuda (Lamy and Pointier,

2017); Bahamas (Beu and Maxwell, 1987; Redfern, 2013):

Cuba; Haiti (Beu and Maxwell, 1987); Mexico (Vokes and

Vokes, 1983; Beu and Maxwell, 1987); Jamaica (Beu and

Maxwell, 1987); Honduras (Lamy and Pointier, 2017); Costa

Rica (Lamy and Pointier, 2017); Panama (Lamy and

Pointier, 2017); Colombia (Daccarett and Bossio, 2011);

Puerto Rico (Menke, 1828); Virgin Islands (Mérch, 1877;

Beu and Maxwell, 1987); St. Martin (Lamy and Pointier,

2017); Anguilla (Lamy and Pointier, 2017); Guadeloupe

(Mérch, 1877; Lamy and Pointier, 2017); Martinique

(Mérch, 1877); Trinidad and Tobago (Beu and Maxwell,

1987); Venezuela (Beu and Maxwell, 1987); Suriname

(Altena, 1975): Brazil: from North to Southeast coast (?)

(Rios 2009). From 0 to 178 m (Rosenberg et al., 2009).

Remarks: The whereabouts of the type material of T.

lanceolata is unknown and as mentioned above, it is

probably lost. The original description is not very infor-

mative and the species was not figured by Menke (1828),

consequently the true identity of T. lanceolata is chal-

lenging to assess despite being frequently cited in the

literature. We have examined one shell from Puerto Rico,

area of the type locality which was not precisely defined,

but this specimen is severely worn (Figure 12). We follow

here the description of Beu and Maxwell (1987) and

Harasewych et al. (1992) to recognize T. lanceolata.

Tritonoharpa lanceolata usually has slightly distorted

teleoconch whorls (Figures 10-14, 16) in comparison to

its congeners and reaches a larger size. Beu and Maxwell

(1987) described a variation of the teleoconch sculpture in

T. lanceolata, in which axial and spiral cords vary from

narrow to thick, resulting in different degrees of

L.S. Souza et al., 2020

Page 99

Figures 10-22. Tritonoharpa species. 10-20. Tritonoharpa lanceolata (Menke, 1828). 10, 11, 18, 20. ANSP 395042. 12. USNM

662026. 13, 15. ANSP 294763. 14, 16, 17, 19. ANSP 408384. 21, 22. Tritonoharpa cubapatriae (Sarastia, 1975): holotype, Museo

Poey, Universidad de La Habana, Cuba. 10, 12-14, 16, 21. Entire shell in frontal view. 11. Entire shell in lateral view. 15, 17. Detail

of teleoconch sculpture. 18. Detail of protoconch in lateral view. 19, 20. Detail of protoconch in apical view. 22. Entire shell in dorsal

view. Scale bars: 10-14, 16, 21, 22 = 5 mm; 15 = 1 mm; 17 = 100 pm; 18 = 500 pm; 19, 20 = 200 pm. Credits: 19, 20, José

Espinosa.

nodulation. Harasewych et al. (1992: 45, fig. 8) depicted a

detail of the sculpture of T. lanceolata, showing spiral

cords more pronounced and thicker than axial cords.

Shells of T. lanceolata examined here also show well

developed spiral cords and weaker axial cords (Figure 17).

Tritonoharpa lanceolata also has a variable color pat-

tern in the protoconch and teleoconch (Beu and Maxwell,

1987). As currently known, the protoconch varies from

pale cream (Figures 12, 13) to purplish (Figures 10, 14,

18-20) and the teleoconch varies from homogeneous

pale cream with brownish spots and streaks (Figures 10,

13) to darker shells (Figure 14). Whitish shells are

usually beach worn specimens (Figure 12). Until more

evidence about the identity of T. lanceolata become

available (i.e., discovery of the type material, anatomy,

genetics, more material from Puerto Rico), it is difficult

to assess if there is more than one distinct species under

the same name.

In Brazil, the first record of T. lanceolata was reported

by Rios (1970) (as “Colubraria lanceolata”), who cited

localities from the Northeast and Southeast (Trindade

Island) regions. Rios (1975; 1985) added a record from

the North of Brazil. Later, Rios (1994) recorded T.

lanceolata only in Para, North Brazil and Rios (2009)

repeated again the records from North to Southeast

coast of Brazil. Rios’ illustrations (1975; 1985; 1994;

Page 100

THE NAUTILUS, Vol. 134, Nos. 3-4

Figures 23-27. Tritonoharpa bayeri (Petuch, 1987): holotype of Colubraria bayeri Petuch, 1987, USNM 859853. 23. Entire shell in

frontal view. 24. Entire shell in lateral view. 25, 26. Detail of protoconch in lateral and apical views, respectively. 27. Detail of

teleoconch sculpture. Scale bars: 23, 24 = 5 mm; 25-27 = 400 pm. Credits: 23, 24, USNM.

2009) do not permit the evaluation of the pattern of

sculpture in the teleoconch. Thus, a robust delimitation

of the species based on these figures is not possible.

Furthermore, despite being a catalogue of mollusks from

Brazil, the shells figured by Rios are not always from this

country (pers. obs.).

The malacological collections studied in the present

work usually housed vouchers from Brazil identified as T.

lanceolata. However, all the shells from Brazil studied

seems to present axial ribs thicker and slightly more

pronounced in comparison to the material from the

northern hemisphere, but a few shells were studied under

SEM. These features lead us to doubt whether T. lan-

ceolata does really occur in Brazil.

Tritonoharpa cubapatriae (Sarastia, 1975)

(Figures 21, 22)

Colubraria (Esbelta) cubapatriae Sarastia, 1975: 4, figs. 1-2.

Tritonoharpa cubapatriae (Sarastia, 1975) — Petit and

Harasewych (1990: 17 [annotated list]; 2005: 42 [an-

notated list]); Lee (2009: 123); Rosenberg et al. (2009:

661 [annotated list]); Espinosa et al. (2012: 291, fig. 524

[reproduced from original illustration]).

Type Locality: Cuba: Habana: Marianao, 20 m.

Type Material: Holotype: Museo Poey, Universidad de

La Habana, Cuba (sh; here examined). Formerly at

“Instituto de Zoologia, Academia de Ciencias de Cuba”

and catalogued as type number 31 (sh; here examined by

photographs) (Sarastia 1975).

Distribution: Cuba (Sarastia, 1975); Panama (Lee,

2009). Known from 20 m (Sarastia, 1975).

Remarks: Beu and Maxwell (1987: 39) considered this

species as the possible largest specimen of T. lanceolata

due to the size of the holotype (45 mm long) (Sarastia,

1975) (Figures 21, 22). According to these authors: “ex-

amination of more extremely large western Atlantic

specimens will be necessary to be sure of the status of T.

cubapatriae”. Indeed, the differences pointed by Sarastia

(1975) in comparisons with T. lanceolata may be related to

the ontogeny phase of the species and we agree with Beu

and Maxwell (1987) in that the shell of T. cubapatriae does

not have more convex whorls than T. lanceolata.

The largest specimen of T. lanceolata we have examined

reaches about eight whorls, 29.1 mm long, collected in

Anguilla, Caribbean (USNM 92993). Lamy and Pointier

(2017: pl. 184, figs. 9A, B) illustrated a shell from Mar-

tinique with 38.4 mm long.

The only additional record of T. cubapatriae since its orig-

inal description was reported by Lee (2009: 123). He referred

to material from Caribbean Panama in a private collection,

with no information about depth and no illustration.

Tritonoharpa bayeri (Petuch, 1987)

(Figures 23-27)

Colubraria bayeri Petuch, 1987: 102, pl. 24, figs. 11-12.

Colubraria bayeri Petuch, 1987 — Daccarett and Bossio

(2011: 94, fig. 446).

Tritonoharpa bayeri (Petuch, 1987) — Lamy and Pointier

(2017: 528, pl. 184, fig. 8A-B).

Type Locality: Colombia: Guajira Peninsula, Off Cabo

La Vela, 35 m.

Type Material: Holotype: USNM 859853 (sh; here

examined).

Measurements: Holotype, USNM 859853: Whorls= 7.0;

SL= 16.2 mm; BWL= 11. mm; AL= 8.3 mm; SW=

7.6mm; AW= 3.7 mm; PCH= 1.0 mm; PCW= 1.2 mm; SL/

SW= 2.1.

L.S. Souza et al., 2020

Distribution: Guadeloupe (Lamy and Pointier, 2017),

Colombia (Petuch, 1987; Daccarett and Bossio, 2011).

From 8 to 35 m (Petuch, 1987; Lamy and Pointier, 2017).

Remarks: Tritonoharpa bayeri was \nown only from

Colombia until recently when Lamy and Pointier (2017)

recorded the species in Guadeloupe. The shell figured by

Lamy and Pointier (2017: pl. 184, fig. 8A-B) has a dark

brown coloration at the spire, differing from the holotype.

Petuch (1987) introduced the species in Colubraria

Schumarcher, 1817 and made no comparisons to other

species of Tritonoharpa. Tritonoharpa bayeri is very

similar to T. curvapex and was distinguished above.

Tritonoharpa bayeri differs from T. lanceolata and T.

leali by the faster increase in diameter, reaching a smaller

ratio of the SL/SW (2.1 vs. 2.8-3.2 and 2.6, respectively).

Furthermore, the aperture of T. bayeri is not so con-

stricted posteriorly as in T. lanceolata and T. leali.

Tritonoharpa bayeri has a similar ratio of SL/SW to

the initial whorls of T. cubapatriae, the latter species

cease a strong increase in diameter after the fifth or sixth

whorl and becomes more cylindrical in shape, reaching

a higher number of whorls and length. The presence of

thinner and lower varices in T. bayeri through all whorls

indicates that this species is not a young specimen of

T. cubapatriae.

Tritonoharpa leali Harasewych, Petit, and Ver-

hecken, 1992

(Figures 28-39)

Tritonoharpa leali Harasewych, Petit and Verhecken,

1992: 45, figures te path

Tritonoharpa leali Harasewych, Petit and Verhecken,

1992 — Rios (1994: 155, pl. Sly fig. 683A): Absalao et al.

(2006: 244): Rios (2009: 298, fig. TOU

Type Locality: Brazil: Vit6ria-Trindade Seamount

Chain, Davis Bank, MD55 stn. DC40, 20°40’ S, 34°41’ W,

60 m, coll. R/V MARION DUFRESNE, v/1987.

Type Material: Holotype: MORG 28659 (sh; here ex-

amined). Paratype: Brazil: Bahia state: off Itaparica Is-

land (USNM 860521, 1 sh; here examined).

Other Material examined: Brazil: Rio Grande do Norte

state: BPOT stn. MR45 (04°27'18" S, 37°04’41”" W, 47 m),

coll. 01/4/2010: MNRJ 34461 [2 sht]; BPOT stn. MR41

(04°49'39” S, 36°10/08” W, 59 m), coll. 27/x/2009: MNR]

34462 [1 sht]; Bahia state: Salvador, Barra (10-15 m), coll.

B. Linhares, xii/1992: MNRJ 21547 [6 shf]; (13°19'52” S,

38°52'52” W, 33 m): MNRJ 27933 bish tl (Sea 7/58" S,

38°44'38.40"” W).: MNRJ 28108 leo te (13927588;

38°46'26.40" W, 35 m): MNRJ 14931 [1 sht];

(13°28'01.20" S, 38°48’00"” W, 30 m): MNRJ 14926 [1

sht]; (13°28’30" S, 38°48'43.20” W, 30 m): MNRJ 14927

[1 sht]; (13°29'42” S, 38°48'18” W, 33 m): MNRJ 14928

Page 101

[1 sht], MNRJ 34464 [1 sht]; (13°30'43” S, 38°49'08.40"

W, 29 m): MNBJ 28848 [1 sht]; REVIZEE Central V stn.

5R (15°34’05"S, 38°49’48” W, 20 m), coll. 25/x/1997:

IBUFR] 12282 [1 sht]; REVIZEE Central I stn. C76

($5°5346° 9S. 38°31 05" WS 66 mm) coll. 30/v/1996:

IBUFR] 9186 [1 sh]; IBUFRJ 10115 [1 sh]; Off Nova

Vicosa, Abrolhos reef (17°57'58"S 38°42’18”W), coll. P.

Young and C. B. Castro, 19/ii/1993: MNRJ 10167 [1

spmt]; REVIZEE Central V stn. 16R (18°03'32"S

37°18'54"W, 100 m), coll. 29/i/2001: IBUFRJ 14437 [1

sh]; REVIZEE Central I stn. C66 (18°19'59” S, 38°55/01”

W, 41 m), coll. 26/iv/1996: IBUFRJ 9069 [1 sh]; Espirito

Santo state: REVIZEE Central I stn. VV38 (19°28'26’S,

38°22’30”" W, 71 m), coll. 29/1/1996: IBUFR]J 9809 [1 sh];

REVIZEE Central I stn. D39 (19°28’41” S 38°22'26"W,

84 m), coll. 29/1/1996: IBUFR] 7755 [1 sh]; REVIZEE

Central I stn. VV24 (20°00'18” S, 39°54’36” W, 45 m), coll.

27/i/1996: IBUFRJ 13027 [1 sh]; REVIZEE Central VI

stn. Y7 (20°50'56” S, 40°10'01” W, 75 m), coll. 28/vi/2002:

IBUFRJ 16324 [1 sh]; REVIZEE Central IT stn. 35R

(20°52'01" SS, - 4081001" WSS mm) coll: O8/xi/1997:

IBUFR] 12367 [1 sh]; AMBES 7 stn. A2 (21°03'29” S,

40°22'59" .\W, 40° m): MNRJ 34945 [1 spmt J; Rio de

Janeiro state: Arraial do Cabo, Forno beach: (22°57'58" S,

42°00'54"” W), coll. M.R. SA and G. Nunan, 18/xii/1983:

MNB]J 14521 [1 sht]; Arraial do Cabo, Prainha beach, coll.

P.M.S. Costa: MNRJ 22245 [1 sht]; (22°57'58" S, 42°00'39”

W, 6 m), coll. 18/ii/2005: MZSP 49326 [1 sh]; Sdo Paulo

state: Ilhabela, Vitéria Island (~23°44'59" S, 45°01'00" W.

3-10 m), coll. 06/xii/2012: MZSP 109352 [3 spm].

Measurements: Holotype, MORG 28659: Whorls= 5.5;

SL= 9.2 mm; BWL= 6.2 mm; AL= 4.4 mm; SW=

3.6 mm; AW= 2.1 mm; PCH= not measured; PCW= not

measured; SL/SW= 2.6; Paratype USNM 860521:

Whorls= 7.0; SL= 18.0 mm; BWL= 12.0 mm; AL=

7.8 mm; SW= 6.9 mm; AW= 4.0 mm; PCH= 0.9 mm;

PCW= 1.0; SL/SW= 2.6.

Distribution: Brazil: Rio Grande do Norte (present

study); Bahia (Harasewych et al., 1992; this study); Vitoria-

Trindade Seamount Chain (Harasewych et al., 1992); Rio

de Janeiro (present study); Sao Paulo (present study).

From 3 m to 100 m.

Remarks: Harasewych et al. (1992) pointed out some

differences between T. leali and T. lanceolata, mentioning

a “more inflated and thinner shell” in the former. The

holotype of T. leali (Figures 28, 29) is a relatively young

individual, with a stocky appearance, however the species

reaches a larger size (MNRJ 21548: one shell reaching

21.2 mm long) and in this case the general shape becomes

more elongated and more similar to the shape of T.

lanceolata. Small specimens of T. lanceolata also have a

more inflated appearance as noted by Beu and Maxwell

(1987: 40). Another difference pointed by Harasewych

et al. (1992) was the presence of axial cords more pro-

nounced than the spiral cords in contrast to T. lanceolata.

All the shells of Tritonoharpa from Brazil examined in the

Page 102

THE NAUTILUS, Vol. 134, Nos. 3-4

Figures 28-34. Tritonoharpa leali Harasewych, Petit and Verhecken, 1992. 28, 29. Holotype, MORG 28659; 30-34. Paratype,

USNM 860521. 35, 38. MNRJ 34462. 36, 37. MNRJ 34461. 28, 30, 35. Entire shell in frontal view. 29, 31. Entire shell in lateral view.

32, 38. Detail of teleoconch sculpture. 33, 36. Detail of protoconch in lateral view. 34, 37. Detail of protoconch in apical view. Scale

bars: 28-31, 35 = 5 mm; 32, 33 = 500 um; 38 = 100 um.

present study, except for the newly described species, are

more similar to the pattern of sculpture described by

Harasewych et al. (1992) for T. leali, despite being slightly

thinner (Figure 38) than in the holotype (Harasewych

et al., 1992: fig. 6). Thus, the occurrence of T. lanceolata in

Brazil is considered dubious and previous records are

probably misidentifications of T. leali.

Tritonoharpa leali was known from the central coast of

Brazil from Bahia to Espirito Santo states, and also at the

Vitoria-Trindade seamount chain (Leal, 1991; Harasewych

et al., 1992; Absalao et al., 2006; Rios, 2009). In the present

study, the geographic distribution of T. leali is extended

northwards to the state of Rio Grande do Norte (~04°S)

and southwards to the state of SA0 Paulo (~23°S), both in

Brazil.

One live specimen photographed at Arraial do Cabo,

Rio de Janeiro, Brazil, and here identified as T. leali is

densely spotted of orange in the head-foot area and the

tentacles are almost completely orange (Figure 39). This

color pattern was also described by Modica et al. (2009)

for a shell identified as “T. antiquata (Hinds, 1844)” but it

actually belongs to a species complex of T. angasi (Braziel,

1877) (Modica et al., 2011a: 121, 2011b: 692). The head-

foot of specimens identified as T. lanceolata from Gua-

deloupe, illustrated by Lamy and Pointier (2017: 528,

text-fig), and from areas near Peanut Island, Florida,

figured at the website of Bill Frank (www.jaxshells.org/

tlance5.htm), also have a very similar color pattern. This

color pattern seems common in the genus. It is also

possible to observe a large and flattened penis in the

Brazilian specimen (Figure 39), similar to the penis of “T.

antiquata” (Modica et al., 2009: fig. 7P). A more detailed

comparison of the penial morphology is limited by the

condition of each specimen (alive vs. preserved).

L.S. Souza et al., 2020

alr) . na

Figure 39. Tritonoharpa leali Harasewych, Petit and Ver-

hecken, 1992: live specimen photographed at Prainha, Arraial do

Cabo, RJ, Brazil (Photo by P.M.S. Costa), observed under a rock.

Black arrow indicates the penis.

Tritonoharpa janowskyi Petuch and Sargent, 2011

(Figures 40-44)

Tritonoharpa janowskyi Petuch and Sargent, 2011: 177,

pl. 5, fig. D.

Tritonoharpa janowskyi Petuch and Sargent, 2011 —

Petuch (2013: 42, fig. 3.7F [reproduced from original

illustration] )

Type Locality: USA: Florida: Palm Beach County, Off

Palm Beach Island, 120 m.

Type Material: Holotype: USNM 1152535 (sh; here

examined).

Measurements: Holotype, USNM 1152335: Whorls= 7.5;

SL= 16.0 mm; BWL= 11.0 mm; AL= 8.0; SW= 7.3 mm;

AW= 4.0 mm; PCH= 1.1 mm; PCW= 1.1] mm; SL/

SW= 2.2.

Distribution: Only known from the type locality.

Remarks: Petuch and Sargent (2011) referred exclusively

to the holotype (Figures 40-44) and just a “few other

specimens” from the type locality. No other additional

record of the species was reported.

Tritonoharpa janowskyi is similar in shape to T. bayeri

(Figures 23-27), as both have a stocky appearance. The

former can be distinguished from T. bayeri by the pro-

portionally smaller length and width of the teleoconch, by

the thicker varices and by the higher number of axial cords

between varices (30 vs. 16-18) (Petuch, 1987; Petuch and

Sargent, 2011). Tritonoharpa janowskyi has a violet color

in the protoconch (Figures 42, 43) while T. bayeri has a

colorless protoconch (Figures 25, 26), but this difference

may be due to the conservation status of the shells. The

teleoconch of both species has a similar color pattern of

irregular brownish patches (Figures 23, 24, 27, 40,

Al AAy,

Tritonoharpa janowsky differs from T. leali in having

proportionally bigger dimensions comparing the holo-

types of both species. Furthermore, the spiral cords are

Page 103

more pronounced than axial cords in T. janowsky in

contrast to T. leali, and the former reaches a higher

number of axial cords between varices (30 us. 22).

DISCUSSION

The present overview of Tritonoharpa from the western

Atlantic demonstrates that there are doubts in the delim-

itation of most species and a more complete taxonomic re-

vision is necessary. The newly described species, T. curvapex,

is the most easily recognizable species presenting a

distinctive protoconch shape (Figures 4, 9). The other

species were rarely reported in the literature, except T.

lanceolata, and were described based on a few or unique

specimens. As commented by Verhecken (2011), who

studied species of Tritonoharpa from the Philippines,

the identification at species level in Tritonoharpa is rather

complicated. Modica et al. (2011b: 692) also pointed about

the difficulties in the taxonomy of Tritonoharpa species

from shallow and deeper waters of the Pacific, Indian and

Atlantic Oceans.

Although we recognized some diagnostic shell features

of most of the Tritonoharpa from the western Atlantic, the

scarcity of individuals hinders description of the variability

of these features. Tritonoharpa lanceolata is supposedly

the most common species, despite uncertainties about its

taxonomy, and shows a variable teleoconch sculpture and

color pattern (Beu and Maxwell, 1987). More recently,

Lamy and Pointier (2017: 528, pl. 184, figs. 8-14) iden-

tified two nominal species (T. bayeri and T. lanceolata)

and five morphotypes of Tritonoharpa sp. from Guadeloupe

(Caribbean). These authors did not describe the diagnostic

features of each morphotype, but based on the illustrations

they possibly considered the different color patterns in the

shells. At present, distinction of species by color pattern is a

dubious approach.

Knowledge about the anatomy of Tritonoharpa is even

more scarce, in the present study a few specimens of T.

leali with soft parts were originally available for study.

Unfortunately, part of these specimens was destroyed in

the fire of “Museu Nacional” prior to anatomical studies. A

Tritonoharpa species from the Philippines, erroneously

cited as T. antiquata (Modica et al., 2009, 201 1a, 2011b),

is the only species of the genus of which the anatomy is

known. More investigations on the anatomy of Tritono-

harpa may help in the species delimitation.

Thus, based on the material examined here we can

conclude that two species of Tritonoharpa certainly occur

in Brazil: T. curvapex and T. leali. Tritonoharpa leali has a

wider range of distribution in the Brazilian coast than

previously reported in the literature, with occurrences in

most of the Tropical Southwestern Atlantic province.

However, it is possible that we were unable to recognize

more than one species under the name T. leali in the

present overview. The occurrence of T. lanceolata in

Brazil needs clarifications.

The protoconch of all species of Tritonoharpa from the

western Atlantic indicates a lecitotrophic development by

Page 104

THE NAUTILUS, Vol. 134, Nos. 3-4

Figures 40-44. Tritonoharpa janowski Petuch and Sargent, 2011: holotype, USNM 1152535. 40. Entire shell in frontal view. 41.

Entire shell in lateral view. 42, 43. Detail of protoconch in lateral and apical views, respectively. 44. Detail of teleoconch sculpture. Scale

bars: 40, 41 = 5 mm; 42, 43 = 500 wm; 44 = 400 pm.

its diameter and number of whorls, which may explain the

restricted geographic distribution of most species. Tri-

tonoharpa lanceolata difters from the remaining species

by its wider distribution, but as discussed above, this

distribution is possibly over estimated based on the doubts

about the identity of this species.

We hope that the illustrations of specimens, including

type material whenever possible, and the gathering

of data about western Atlantic Tritonoharpa will be

useful for future studies about this genus, despite some

of the tentative identifications in a confusing taxonomic

scenario.

ACKNOWLEDGMENTS

We are grateful to Gary Rosenberg, Paul Callomon, and

Nasreen Phillips (ANSP), Amanda Bemis and John

Slapcinsky (FLMNH), Cleo Oliveira (IBUFR]), José

Espinosa (Instituto de Oceanologia, Cuba), Alexandre

Pimenta (MNRJ), Paula Spotorno-Oliveira and the late

Eliézer de C. Rios (MORG), Luiz Simone (MZSP), Sigrid

Hof (SMF), Ellen Strong and Yolanda Villacampa

(USNM) for access to collections, loan of specimens,

photographs and/or information regarding type material.

We thank André Verhecken (RBINS), José H. Leal

(BMSM), Jean-Francois Lesport and Jean-Michel Pacaud

(MNHN), Rodrigo Salvador (MNZ, Te Papa Tongarewa)

for providing important literature concerning Cancellar-

iidae. Many thanks to M.G. Harasewych (USNM) and

André Verhecken for providing critiques that improved

the manuscript. We appreciate the support of C. Messias

(MN-UFRJ) for SEM operation. Thanks to Petrobras SA

for the material collected during BPOT MR and AMBES

projects. The senior author very much appreciates the

support of his friends Carine Goncalves and Gustavo

Miranda during a research visit to the USA.

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THE NAUTILUS 134(3-4):107-116, 2020

Page 107

Memorial to Warren O. Addicott—A remarkable student of Tertiary

marine mollusks of the northeastern Pacific

Charles L. Powell, II

U.S. Geological Survey, Menlo Park, CA 94025

[email protected]

Kevin J. Meade

California State University, Fullerton

Fullerton, CA 92831 USA

[email protected]

ABSTRACT

Warren Oliver Addicott (1930-2009) was a prolific molluscan

paleontologist who produced in excess of one hundred publi-

cations, most of which deal with Tertiary fossil mollusks from

western North America. In these publications he is credited with

naming more than 80 new molluscan taxa, and recognizing and

illustrating an additional 17 taxa that lacked sufficient material to

be described as new. In addition, he is honored in having nine

species named for him (eight mollusks and one arthropod).

"Warren Addicott was an exemplary paleontologist, far

ahead of his peers, and willing to move into new areas and

issues. He helped to supervise the Circum-Pacific Map

Project, with many maps being put together and published

under his tenure." (Ellen J. Moore, 2010, pers. comm.,

October 2010)

“ .Warren Oliver Addicott (Ph.D. 1956), whose career with

the Paleontology and Stratigraphy Branch of the U.S.

Geological Survey in Menlo Park set new standards for

monographic systematics, faunal and_biostratigraphic

documentation, and photographic illustration as well as

initiating new insights and integration of molluscan research

with active margin tectonics, refined climate curves, lat-

itudinal distributions based on marine mollusks, increased

radiometric age control on molluscan faunas, extension of

molluscan paleontological interpretations into Alaska, and

fostering exchange and collaboration with Japanese mol-

luscan paleontologists.” (Hickman, 2009.)

MEMORIAL

Warren Addicott was born in Fresno, California on

February 17, 1930 to Irwin and Astrid Addicott. He was

the eldest of two brothers, the younger being Alan

Addicott. His formative years were spent in California

where he attended Fresno High School in Fresno. At

Fresno High School he played basketball making the

local “Fresno Bee’s [newspaper] All Yosemite League

Basketball Team” (The Fresno Bee newspaper, Feb-

ruary 27, 1946: 11; https:/Avww.newspapers.com/clip/

§987288/the-fresno-bee-the-republican/). Afterward he

attended Pomona College, where he was a student of

the founding director of the geology department,

Alfred O. Woodford. Warren graduated in 1951 witha

BA in geology. A year later he received his M.A. from

Stanford University’s Hopkins Marine Station in Pa-

cific Grove, California. Subsequently he attended the

University of California at Berkeley where he received

his Ph.D. from the Museum of Paleontology in 1956.

While there, Warren completed his pioneering work

on the Neogene Kern River deposits (1956) under

J. Wyatt Durham, Charles W. Gilbert, and Ralph L.

Langenheim.

In 1962, Warren joined the then-expanding Paleon-

tology and Stratigraphy Branch of the U.S. Geological

Survey (USGS) in Menlo Park, California. There he had a

long and illustrious career focused on molluscan fossils of

the eastern Pacific collaborating with colleagues in the

Russian Far East, Japan, and later Spain. His first paper

published by the USGS was with Jack G. Vedder, a re-

nowned geologist in his own right, where they discussed

the relationship between paleotemperature and Miocene

mollusks in the Kern River area. Probably his most

prominent publication is “Miocene Gastropods and

Biostratigraphy of the Kern River Area, California”

(Addicott, 1970d), which, 50 years later, remains a classic

and an outstanding publication. In addition to his pub-

lications (listed below) Warren authored hundreds of

Internal Reports on Referred Fossils (known as E&R’s

within the USGS) for other federal scientists and scholars

around the world. John Barron (USGS) related a memory

of Warren to us: when he first started his career at the

USGS, he asked Warren for advice on how he should

work. Warren suggested he divide his time into thirds, one

third on service work (E&R’s), one third on collaboration,

and one third on his own research. This division of labor

has similarly worked well for many USGS paleontologists

over the years.

Warren was always willing to aid students and fellow

scientists in their work. He sponsored the USGS postdoc

of Judith Terry Smith (Smithsonian) for a year, although

the two did not interact much. He helped Clifford M.

Nelson with his dissertation on the gastropod genus

Page 108 THE NAUTILUS, Vol. 134, Nos. 3-4

Figure 1. Warren O. Addicott. Photograph courtesy of Eric Addicott.

Neptunea. Nelson would later write a letter to Warren that when speaking with Warren in 1973 about his un-

thanking him for his help; this letter is now preserved in dergraduate thesis, he offered to loan him some fossils

the archives at the Museum of Paleontology, University of that Warren and Joe Clark (Indiana University of Penn-

California, Berkeley. Richard G. Stanley (USGS) recalls sylvania) had collected from Richard’s field site. Warren

C.L. Powell, II and K.J. Meade, 2020

was also interested in Richard’s 1987 poster on right-

lateral displacement of the San Andreas fault, which many

other scientists disapproved of at the time. “I remember

Warren as an exemplary and thoughtful scientist, and as a

generous and kind human being,” he related to us. Warren

always answered letters that Lindsey Groves (Natural

History Museum of Los Angeles County) sent him, as this

latter worked on his masters thesis at California State

University, Northridge. In addition, whenever possible he

joined Robert J. McLaughlin and Dave Durham (both

USGS) on field trips to Salinas Valley localities, and was

always willing to look at “the most scruffy material.” James

C. Ingle, Jr. (Stanford University) tells of how he worked

with Warren on several US-Japan projects dealing with

Neogene paleontology around the Pacific Rim and that he

later helped Jim obtain funding for field work on the

Burica Formation in Panama. “He was a premier pale-

ontologist, a wonderful colleague, and simply a great

human being in every respect,” he recalled.

In 1971, Warren became a consulting Associate Pro-

fessor of Geology at Stanford University where he served

as counsel for graduate students and supervised the

University’s Cenozoic and modern mollusk collections.

Around 1980, Warren left the Branch for Paleontology

and Stratigraphy, joining the Circum-Pacific Map Project

(C-P M Project), because he was tired of having to ask for

money from projects he helped. The goal of the C-P M

Project was to gather together scientists from different

parts of the world toward developing geologically signif-

icant maps throughout the Pacific region. Warren was

appointed General Chairman of the C-P M Project in

1983, taking over from John A. Reinemund, who had been

appointed Director in 1982. A few years later, in 1986,

Warren retired from the USGS. That same year, he and

his second wife Suzanne Aubin moved to Ashland, Ore-

gon. After moving, Warren taught at the University of

Southern Oregon from 1986 until his final retirement in

1990.

Throughout his career Warren was heavily involved in

the broader scientific community. He became a fellow of

the California Academy of Sciences, gave lectures and

lead field trips for the American Association of Petroleum

Geologists (AAPG) along with being editor of the Pacific

Petroleum Geologists, a newsletter of the Pacific Section

AAPG, and from 1971 to 1981 was a Trustee of Bulletins

of American Paleontology (Paleontology Research Insti-

tute, Ithaca, New York). He was a founding member of

the Western Society of Malacologists (1968) and is be-

lieved to have regularly attended meetings until at least

1975. In 1970, Warren was elected Secretary of the Pa-

leontological Society, a position he held until 1976. Later,

he was elected President of the Paleontological Society

(1980). He also served on the editorial board for The

Veliger, a quarterly published by the California Mala-

cozoological Society, Inc.

When not working on research, Warren loved to gar-

den. He was vice-president of the Peninsula Camelia

Society in 1964 after moving to Menlo Park for his job at

the USGS. In 1964 and again in 1968 he won several

Page 109

camellia growing contests run by the local camellia society

(Southern California Camellia Society, 1964: 22; 1968:

9-18), and wrote about the subject in the specialized

literature (Addicott, 1967d: 27.)

He was also skilled at ceramics and a talented poet.

Warren loved music, especially blues and jazz, and

loved to compose his own music that he would play on

the piano. He was married twice, first to Susanne Smith

(1955-1972), and later to Suzanne Aubin (1976-2009).

He left two children, Eric Addicott and Carol Kral, as

well as two stepchildren, David Le Drew and Christine

LeDrew-Johnson. Eric recounted to us: “Dad made it a

priority to spend lots of time with my sister and me, and

in many ways the divorce enhanced his relationship

with us. He was always so loving and caring, and he

always trusted my judgment and never really tried to

guide or advise me too much. He introduced me to

fishing, backpacking, cycling, and skiing, activities which I

still do.”

Warren was a world traveler and took an avid interest in

other cultures. Not only did he travel for research-related

reasons, but also for pleasure. He would even take time

during overseas scientific conferences to visit scenic lo-

cations. For example during the 1971 Mediterranean

Neogene Congress in Lyon, France, he visited the Rhone

and Bordeaux basins in France.

After his retirement he moved to Ashland, where he

continued to travel. In 1997, he wrote a letter to the Nu

Alpha Phi fraternity at Pomona College (of which he was a

member during his time there) noting that he and his

second wife Suzanne had traveled to India in 1996 and

stayed there for three months. He also worked as a realtor

while in Ashland to keep busy.

Warren O. Addicott passed away July 11, 2009, with his

wife Suzanne by his side, from complications of cancer. A

hard worker and kind man, he named over 80 new

molluscan taxa during his lifetime. His contributions to

molluscan paleontology, and paleontology in general,

solidified Warren Addicott’s name as one of the great

paleontologists of the late 20th Century.

NEW SPECIES NAMED IN HONOR OF WARREN

O. ADDICOTT

(names presented as in original combinations)

Mollusca

Bivalvia

Macoma addicotti Nikas (1977)

Mactromeris addicotti Kanno (1971)

Modiolus addicotti Moore (1984)

Mytilus (Tumidimytilus) addicotti Kafanov (1985) —

Holotype figured by Allison and Addicott (1976), pl. 3,

fig. 9 [= My. addicotti|

Pitar addicotti Zinsmeister (1983)

Cephalopoda

Moroteuthis addicotti Clarke and Fitch (1979)

Gastropoda

Nassarius grammatus addicotti Adegoke (1969)

Scaphopoda

Page 110

Cadulus (Platyschides?) addicotti Emerson (1957)

Arthropoda

Crustacea

Balanus addicotti Zullo (1979)

NEW MOLLUSCAN NAMES PROPOSED BY

WARREN O. ADDICOTT

Bivalvia

Nucula salmonensis Addicott, 1966c

Platyodon colobus fowleri Addicott, 1966c

Propeamussium (Propeamussium) leohertleini Addicott,

1971la

Gastropoda

Acteon (Rictaxis) weaveri Addicott, 1970d

Aforia clallamensis tricarinata Addicott, 1966c

Antillophos woodringi Addicott, 1970d

Austrotrophon kernensis medialis (Addicott, 1970d)

Balcis lutzi Addicott, 1970d

Balcis petrolia Addicott, 1970d

Barkeria Addicott, 1970d — as a subgenus of Cancellaria

Calicantharus rancherianus Addicott, 1970d

Calicantharus woodfordi Addicott, 1970d

Calliostoma carsoni Addicott, 1970d

Calyptraea coreyi Addicott, 1970d

Cancellaria (Crawfordina) kernensis Addicott, 1970d

Cancellaria (Euclia) circumspinosa Addicott, 1970d

Cancellaria (Euclia) ocoyana Addicott, 1970d

Cancellaria (Narona) birchi Addicott, 1970d

Cancellaria galei Addicott, 1970d

Cancellaria keenae Addicott, 1970d

Catilon Addicott, 1965c — a subgenus of Nassarius

Crassispira olcesensis Addicott, 1970d

Crepidula bractea Addicott, 1970d

Crucibulum (Dispotaea) papulum Addicott, 1970d

Demondia Addicott, 1965c — a subgenus of Nassarius

Epitoonium (Gyroscala) barkerianum Addicott, 1970d

Epitonium (Nitidscala) tedfordi Addicott, 1970d

Forreria emersoni Addicott, 1970d

Gibbula (Tumulus?) baileyi Addicott, 1970d

Glyphostoma carinata Addicott, 1970d

Haminoea articensis Addicott, 1966c

Kelletia lorata Addicott, 1970d

Knefastia grarcesana Addicott, 1970d

Mangelia (Notocytharellar) hartensis Addicott, 1970d

Mitra (Atrimitra) andersoni Addicott, 1970d

Mitrella (Columbellopsis) alta Addicott, 1970d

Morula (Morunella) granti Addicott, 1970d

Nassarius (Catilon) smooti Addicott, 1965c

Nassarius hoquiamensis Addicott, 1966c

Nassarius (Phrontis) harrellensis Addicott, 1970d

Nassarius (Phorntis?) posoensis Addicott, 1970d

Nassarius (Catilon?) salinasensis Addicott, 1965c

Natica vokesi Addicott, 1966c

Nerita (Theliostyla?) joaquinensis Addicott, 1970d

Niso cottonwoodensis Addicott, 1970d

Ocenebrina clarki (Addicott, 1970d)

Odostomia (Chrysallida?) sequoiana Addicott, 1970d

THE NAUTILUS, Vol. 134, Nos. 3-4

Odontomia (Menestho) repenningi Addicott, 1970d

Polystira englishi Addicott, 1970d

Scaphander dollaris Addicott, 1966c

Sulcoretusa? israelski Addicott, 1970d

Tectionatica? satsopensis Addicott, 1966c

Tegula ellenae Addicott, 1966c

Tegula (Omphalius) dalli arnoldi Addicott, 1970d

Tegula (Omphalius) laevis Addicott, 1970d

Terebra (Fusoterebra?) adelaidana Addicott, 1970d

Terebra (Strioterebrum) stirtoni Addicott, 1970d

Trochotropis tricarinata Addicott, 1970d

Trochotropis (Iphinoe) goweri Addicott, 1966c

Turbonilla (Chemnitzia) hannali Addicott, 1970d

Turbonilla hormigacuesta Addicott, 1970d

Turbonilla (Tragula) greenhomensis Addicott, 1970d

Turbonilla (Ptycheulimella) edisonensis Addicott, 1970d

Turritella kernensis Addicott, 1970d

Turritella (Idaella) vaquinana Addicott, 1966c

Volvuella joaquinensis Addicott, 1970d

In addition to the species named by Warren (above) he

recognized and illustrated a number of other potential

new species. Unfortunately, these possible new species

were not represented by sufficient material for him to

name them and are listed in his publications simply as new

species (n. sp.), or new species? (n. sp.?).

Bivalvia

Macoma n. sp. Addicott, 1969a |= M. addicotti Nikas,

1977]

Gastropoda

Astraea (Pomaulax) n. sp. Addicott (1970d)

Cancellaria n. sp. Addicott, 1969a |= C. fergusoni Carson,

1926]

Cancellaria n. sp.P aff. C. alaskensis Clark (1932),

Addicott, 1980

Cancellaria (Euclia?) n. sp. Addicott, 1966c

Cymatium n. sp. Addicott, 1970d

Diodora (Diodora) n. sp. Addicott, 1970d

Diodora? n. sp. Addicott, 1970d

Neverita (Glossaulax) n. sp.? Addicott, 1970d

Odostomia (Chrysallida?) n. sp. Addicott, 1970d

Odostomia (Evalea?) n. sp. Addicott, 1970d

Oliva (Oliva) n. sp.? Addicott, 1970d

Terebra (Strioterebrum) n. sp.? Addicott, 1970d

Terebra (Terebra) n. sp. Addicott, 1970d

Triumphis? n. sp. Addicott, 1970d

Turbonilla (Chemnitzia) n. sp. Addicott, 1970d

Turbonilla (Chemnitzia?) n. sp. Addicott, 1970d

Turbonilla (Pyrgiscus) n. sp. Addicott, 1970d

PUBLICATIONS OF WARREN O. ADDICOTT

1953

Emerson, W.K., and W.O. Addicott. 1953. A Pleisto-

cene invertebrate fauna from the southwest corner of

San Diego County, California. San Diego Society of

Natural History Transactions 11(17): 429-444.

C.L. Powell, IJ and K.J. Meade, 2020

1956

Addicott, W.O. 1956. Miocene stratigraphy northeast

of Bakersfield, California. Berkeley. University of

California, Ph.D. dissertation.

1958

Emerson, W.K., and W.O. Addicott. 1958. Pleistocene

invertebrates from Punta Baja, Baja California, Mexico.

American Museum Novitates 1909: 1-11.

1959

Addicott, W.O., and W.K. Emerson. 1959. Late Pleis-

tocene invertebrates from Punta Cabras, Baja California,

Mexico. American Museum Novitates 1925: 1-33.

1963

a. Addicott, W.O. 1963. An unusual occurrence of

Tresus nuttallii (Conrad, 1837) (Mollusca: Pelecy-

poda). The Veliger 5(4): 143-144.

b. Addicott, W.O. 1963. Interpretation of the inverte-

brate fauna from the Upper Pleistocene Battery

Formation near Crescent City, California. Pro-

ceedings of the California Academy of Sciences,

Fourth Series 31(13): 341-347.

. Addicott, W.O., and J.G. Vedder. 1963. Paleo-

temperature inferences from late Miocene mollusks

in the San Luis Obispo—Bakersfield area, California.

U.S. Geological Survey Professional Paper 475C:

C63—C68.

1964

a. Addicott, W.O. 1964. A Late Pleistocene inverte-

brate fauna from southwestern Oregon. Journal of

Paleontology 38(4): 650-661.

b. Addicott, W.O. 1964. Pleistocene invertebrates from

the Dune Terrace, western Santa Monica Mountains,

California. Bulletin of the Southern California

Academy of Sciences 63(3): 141-150.

©)

1965

a. Addicott, W.O. 1965. On the identification of

Schizopyga californiana Conrad, a California Plio-

cene gastropod. Proceedings of the California

Academy of Sciences, Fourth Series 33(2): 47-58.

b. Addicott, W.O. 1965. The enigmatic late Cenozoic

gastropod Schizopyga californiana |abs.]. American

Malacological Union, Annual Report for 1964: 44.

c. Addicott, W.O. 1965. Some western American Ce-

nozoic gastropods of the genus Nassarius. U.S.

Geological Survey Professional Paper 503B: B1—B24.

d. Addicott, W.O. 1965. Miocene macrofossils of the

southeastern Jan Joaquin Valley, Calif. U.S. Geo-

logical Survey Professional Paper 525C: C101-C109.

e. Durham, D.L., and W.O. Addicott. 1965. Pancho

Rico Formation, Salinas Valley, California. U.S.

Geological Survey Professional Paper 524A: Al—A22.

f. Durham, D.L., and W.O. Addicott. 1965. Upper

Miocene and Pliocene marine stratigraphy in

southern Salinas Valley, California. U.S. Geological

Survey Bulletin 1194: E1—E7.

Page 111

1966

a. Addicott, W.O. 1966. Late Pleistocene marine pa-

leoecology and zoogeography in central California.

U.S. Geological Survey Professional Paper 523C:

CIC |

b. Addicott, W.O. 1966. Miocene macrofossils of the

southeastern San Joaquin Valley, California. U.S. Geo-

logical Survey Professional Paper 525C: C101—C109.

c. Addicott, W.O. 1966. New Tertiary marine mollusks

from Oregon and Washington. Journal of Paleon-

tology 40(3): 635-646.

d. Repenning, C.A., D.L. Jones, and W.O. Addicott.

1966. Geology of the Great Valley. Mineral Infor-

mation Service 22(1): 3-6.

1967

a. Addicott, W.O. 1967. Age of the Skooner Gulch

Formation, Mendocino County, California. U.S.

Geological Survey Bulletin 1254C: C1-C11.

b. Addicott, W.O. 1967. Zoogeographic evidence for

late Tertiary lateral slip on the San Andreas fault,

California. U.S. Geological Survey Professional Pa-

per 593D: D1-D12.

c. Addicott, W.O. 1967. Paleontologic evidence for

large post-early Miocene lateral slips on the San

Andreas fault, California [abs.]. Geological Society of

America, Program, 63rd annual meeting, Santa

Barbara, March 22-25, 1967: 17.

1968

a. Addicott, W.O. 1968. Mid—Tertiary zoogeography

and paleogeographic discontinuities across the San

Andreas fault, California. Pp. 144-165 [in:] W.R.

Dickinson and A. Grantz, Proceedings of conference

on geologic problems of San Andreas fault system.

Stanford University Publications, Geological Sci-

ences 11: 1 —387.

b. Addicott, W.O. 1968. Neogene molluscan zoogeog-

raphy and climatic change in the northeastern Pacific

Ocean [abs.]. Geological Society of America, 1968

Annual Meetings, Program with abstracts: 2-3.

c. Bradley, W.C., and W.O. Addicott. 1968. Age of first

marine terrace near Santa Cruz, California. Geo-

logical Society of America Bulletin 79(9): 1203-1210.

1969

a. Addicott, W.O. 1969. Late Pliocene mollusks from

San Francisco Peninsula, California, and their pa-

leogeographic significance. Proceedings of the Cal-

ifornia Academy of Sciences, Fourth Series 37(3):

DTH,

b. Addicott, W.O. 1969. Late Pliocene marine mollusks

from the northeastern Santa Cruz Mountains, Cal-

ifornia [abs.]. Abstracts with Program for 1969 (Cor-

dilleran Section), Geological Society of America, 3: 1.

c. Addicott, W.O. 1969. Tertiary climatic change in the

marginal northeastern Pacific Ocean. Science

165(3893): 583-586.

Page 112

d. Addicott, W.O. and S. Kanno. 1969. Current pale-

ontological investigations on Cenozoic marine mol-

lusks of the west coast of North America. The Veliger

121) 1852130:

. Hopkins, D.M., D.W.Scholl, W.O. Addicott, R.L.

Pierce, P.B. Smith, J.A. Wolf, D. Gershanovich, B.

Kotenev, K.E. Lohman, J.H. Lipps, and

J. Obradovich. 1969. Cretaceous, Tertiary, and early

Pleistocene rocks from the continental margin in the

Bering Sea. Geological Society of America Bulletin

80(8): 1471 —1480.

f. Scholl, D.W., H.G. Greene, W.O. Addicott, W.R.

Evitt, R.L. Pierce, S.H. Mamay, M.S. Marlow. 1969.

Adak “Paleozoic” site, Aleutians—in fact of Eocene

age [abs.]. American Association of Petroleum Ge-

ologists Bulletin 53(2): 459.

1970

a. Addicott, W.O. 1970. Biogeographical sketch of Leo

b.

OQ

Hertlein. The Nautilus 84(2): 37-41.

Addicott, W.O. 1970. Bibliography of Leo George

Hertlein for the period of 1925 to 1970. The Nautilus

849). 43352,

. Addicott, W.O. 1970. Latitudinal gradients in Ter-

tiary molluscan faunas of the Pacific coast. Paleo-

geography, Palaeoclimatology, Palaeoecology 8(4):

237-32:

. Addicott, W.O. 1970. Miocene gastropods and bio-

stratigraphy of the Kern River area, California. U.S.

Geological Survey Professional Paper 642: 1-174.

https://pubs.usgs.gov/pp/0642/report.pdf

. Addicott, W.O. 1970. Tertiary climatic change in San

Joaquin Basin, California — evidence from shallow—

water mollusks [abs.]. American Association of Pe-

troleum Geologists Bulletin 54(3): 561.

f. Addicott, W.O. 1970. Tertiary paleoclimatic trends in

ry

the San Joaquin Basin, California. U.S. Geological

Survey Professional Paper 644D: D1-D19.

. Addicott, W.O. 1970. Additional remarks on studies

of Cenozoic marine mollusks of the Pacific Coast.

The Veliger 13(1): 110.

. Addicott, W.O. 1970. Paleoclimatic history of the

northeastern Pacific margin — a paleontologic ap-

plication of modern molluscan zoogeographic data

[abs.]. The Echo, Western Society of Malacologists,

Annual meeting, Stanford Calif: 15 -16.

. Musuda, K., and Addicott, W.O. 1970. On Pecten

(Amusium) condoni Hertlein from the west coast of

North America. The Veliger 13(2): 153-156.

1971

a.

b.

Addicott, W.O. 1971. Some Paleogene mud pectens

of the genus Propeamussium from Alaska and Cal-

ifornia. The Veliger 13(3): 226-230.

Addicott, W.O. 1971. Tertiary marine mollusks of

Alaska: an annotated bibliography. U.S. Geological

Survey Bulletin 1343: 1-30.

THE NAUTILUS, Vol. 134, Nos. 3-4

c. Addicott, W.O. 1971. Some trends in Neogene

marine molluscan paleontology on the west coast of

North America. Abstracts with Program, Geological

Society of America 3(2): 70.

1972

a. Addicott, W.O. 1972. Clark’s Tertiary molluscan

types from the Yakataga District, Gulf of Alaska. U.S.

Geological Survey Professional Paper 750C:

(Gite Sols

. Addicott, W.O. 1972. Neogene molluscan paleon-

tology along the west coast of North America,

1840-1969 — trends and status. Journal of Paleon-

tology 46(5): 627-636.

. Addicott, W.O. 1972. Provincial middle and late

Tertiary molluscan stages, Temblor Range, Cal-

ifornia. Pp. 1 —26 [in:] Proceedings of the Pacific

Coast Miocene biostratigraphic symposium. Pacific

Section, S.E.P.M., Bakersfield. 364 pp.

d. Addicott, W.O., and G. Plafker. 1972. Paleocene

mollusks from the Gulf of Alaska Tertiary province —

a significant new occurrence on the North Pacific

Rim. U.S. Geological Survey Professional Paper

750B: B48—B52.

1973

a. Addicott, W.O. 1973. A historical resume of research

CO

in Neogene marine molluscan paleontology and

biostratigraphy in western North America. Science

Reports of the Tohoku University. Series 2: Geology

— Tohoku Daigaku Rika Hokoku. Dai 2: Shu

Chishitsugaku 6(6): 7-14.

. Addicott, W.O. 1973. Neogene marine mollusks of

the Pacific coast of North America — an annotated

bibliography, 1797-1969 — a compilation of reports

for the period 1797-1969 dealing with marine

mollusks of Miocene and Pliocene age. U.S. Geo-

logical Survey Bulletin 1362: 1 —-201.

. Addicott, W.O. 1973. Oligocene molluscan biostra-

tigraphy and paleontology of the lower part of the

type Temblor Formation, California. U.S. Geological

Survey Professional Paper 791: 1 —-48.

. Addicott, W.O. 1973. Oligocene and Miocene mol-

luscan stages, Temblor Range, California [abs.].

American Association of Petroleum Geologists Bul-

letin 57(2): 429.

. Addicott, W.O. 1973. Giant Neogene pectinids of

eastern North Pacific — chronostratigraphic and

zoogeograpahic significance [abs.]. American Asso-

ciation of Petroleum Geologists Bulletin 57(4): 766.

f. Addicott, W.O., and J.S. Galehouse. 1973. Pliocene

fe:

marine fossils in the Paso Robles Formation, Cal-

ifornia. U.S. Geological Survey Journal of Research

1(5): 509-514.

Bartow, J.A., A.M. Sarna-Wojcicki, W.O. Addicott,

and K.R. Lajoie. 1973. Correlation of marine and

continental Pliocene deposits in northern California

C.L. Powell, II and K.J. Meade, 2020

—"

by tephrochronology [abs.]. American Association of

Petroleum Geologists Bulletin 57(4): 769.

. Galehouse, J.S., and W.O. Addicott. 1973. Paleo-

geographic significance of Pliocene marine inverte-

brates from the Paso Robles Formation, southern

Coast Ranges, California [abs.]. Abstracts with Pro-

grams, Geological Society of America 5(1): 46-47.

. Nilsen, T. H., T.W. Dibblee, Jr., and W.O. Addicott.

1973. Lower and middle Tertiary stratigraphic units

of the San Emigdio and western Tehachapi Moun-

tains, California. U.S. Geological Survey Bulletin

IBiZae ee.

1974

a.

Addicott, W.O. 1974. Recognition and distribution of

Mytilus condoni Dall, a unique Pliocene and Pleis-

tocene bivalve from the Pacific Coast. The Veliger

16(40): 354-348.

. Addicott, W.O. 1974. Giant Pectinids of the eastern

North Pacific margin: significance in Neogene zoo-

geography and chronostratigraphy. Journal of Pale-

ontology 48(1): 180-194.

. Addicott, W.O., and H.G. Greene. 1974. Zoogeo-

graphic significance of a late Quaternary occurrence

of the bivalve Astarte off the central California coast.

The Veliger 16(3): 249-252.

1975

a. Addicott, W.O. 1975. Early Miocene age of the

O

Clallam Formation, western Washington [abs.]. U.S.

Geological Survey Bulletin 1405A: A26.

. Addicott, W.O. 1975. Miocene biostratigraphy,

western Washington, in Paleontology—Cenozoic of

the United States [abs.]. U.S. Geological Survey

Professional Paper 975: 176.

. Addicott, W.O. 1975. Provincial age and correlation

of the Clallam Formation, northwestern Washington

[abs.]. Abstracts with Programs, Geological Society of

America 7(3): 289.

Snavely, Jr., P.D., N.S. MacLeod, W.W. Rau, W.O.

Addicott, and J.E. Pear. 1975. Alsea Formation — an

Oligocene marine sedimentary sequence in the

Oregon Coast Range. U.S. Geological Survey Bul-

letin 1395F: 1 —21.

1976

a.

Addicott, W.O. 1976. Molluscan paleontology of the

lower Miocene Clallam Formation, northwestern

Washington. U.S. Geological Survey Professional

Paper 976: 1 —-44.

. Addicott, W.O. 1976. New molluscan assemblages

from the upper member of the Twin River Forma-

tion, western Washington: significance in Neogene

chronostratigraphy. U.S. Geological Survey Journal

of Research 4(4): 437-447.

. Addicott, W.O. 1976. Neogene molluscan stages of

Oregon and Washington. Pp. 95-115, in A.E. Frit-

sche, H. TerBest, Jr., and W.W. Wornardt [in:]

Page 113

Neogene Symposium. Pacific Section, $.E.P.M., San

Francisco. 160 pp.

d. Addicott, W.O. 1976. On the significance of the

bivalve Acila gettysburgensis (Reagan) in middle

Tertiary chronostratigraphy of the Pacific Coast. The

Veliger 19(2): 121-124.

e. Addicott, W.O., and P.D. Snavely, Jr. 1976. Re-

connaissance of mollusk—bearing Neogene rocks,

Almeria Province, eastern Andalusia, Spain [abs.].

Western Society of Malacologists, Annual Report 9:

49-50.

f. Allison, R.C., and W.O. Addicott. 1976. The north

Pacific Miocene record of Mytilus (Plicatomytilus) a

new subgenus of Bivalvia. U.S. Geological Survey

Professional Paper 962: 1 —22.

g. Plafker, G., and W.O. Addicott. 1976. Glaciomarine

deposits of Miocene through Holocene age in the

Yakataga Formation along the Gulf of Alaska margin,

Alaska. U.S. Geological Survey Open-File Report

(0-04; 1336.

h. Plafker, G., and W.O. Addicott. 1976. Marine glacial

deposits of Miocene through Holocene age along the

Gulf of Alaska margin. Pp. 1 —23 [in:] T.P. Miller,

Recent and ancient sedimentary environments in

Alaska. Alaska Geological Society, Anchorage 313 pp.

1977

a. Addicott, W.O. 1977. Neogene chronostratigraphy of

nearshore marine basins of the eastern north Pacific.

Proceedings of the First International Congress on

Pacific Neogene Stratigraphy, Science Council, Ja-

pan. Tokyo: 151-175.

b. Addicott, W.O. 1977. Significance of pectinids in

Tertiary biochronology of the Pacific Northwest

States [abs.]. Abstracts with Programs, Geological

Society of America 9(7): 874.

. Addicott, W.O., P.D. Snavely, Jr., D. Bukry, and R.Z.

Poore. 1977. Neogene stratigraphy and paleontology

of southern Almeria Province, Spain: an overview.

U.S. Geological Survey Open—File Report 77-716: 1

—69.

d. Wehmiller, J.F., K.R. Lajoie, K.A. Kvenvolden, E.

Peterson, D.F. Belknap, G.L. Kennedy, W.O.

Addicott, J.G. Vedder, and R.W. Wright. 1977.

Correlation and chronology of Pacific coast marine

terrace deposits of continental United States by fossil

amino acid stereochemistry — technique evaluation,

relative ages, kinetic model ages, and geologic im-

plications. U.S. Geological Survey Open-File Report

tio: L196.

o)

1978

a. Addicott, W.O. 1978. Late Miocene mollusks from

the Queen Charlotte Islands, British Columbia,

Canada. U.S. Geological Survey Journal of Research

6(5): 677-690.

b. Addicott, W.O. 1978. Marine paleogeography and

paleontology of the Salinas basin during the latest

Page 114

‘o)

part of the Miocene with notes on macrofossils from

near San Lucas, California. Pp. 83 —90 [in:] Addicott,

W.O., Neogene biostratigraphy of selected areas in

the California Coast Ranges. U.S. Geological Survey

Open-File Report 78-446 109 pp.

. Addicott, W.O (ed.). 1978. Neogene biostratigraphy

of selected areas in the California Coast Ranges. U.S.

Geological Survey Open-File Report 78—446:1 —109.

. Addicott, W.O. 1978. Notes on the geology of Point

Lobos State Reserve, Monterey County, California.

Pp. 91 —96 [in:] Addicott, W.O., Neogene biostra-

tigraphy of selected areas in the California Coast

Ranges. U.S. Geological Survey Open-File Report

78-446 109 pp..

. Addicott, W.O. 1978. Pectinids as biochronologic

indices in the Neogene of the eastern North Pacific.

Pp. 11 —23 [in:] Proceedings of the second working

group meeting biostratigraphic datum—planes of the

Pacific Neogene. IGCP Project 114. Geological

Research and Development Centre (Bandung,

Indonesia), Special Publication 1 364 pp.

f. Addicott, W.O. 1978. Revision of the age of the

g.

ia;

="

ed

Pancho Rico Formation, central Coast Ranges,

California. U.S. Geological Survey Bulletin 1457A:

88-89.

Addicott, W.O., J.A. Barron, and J.W. Miller. 1978.

Marine late Neogene sequence near Santa Cruz,

California. Pp. 97 —109 [in:] Addicott, W.O., Neo-

gene biostratigraphy of selected areas in the Cal-

ifornia Coast Ranges. U.S. Geological Survey

Open-File Report 78-446 109 pp.

Addicott, W.O., R.Z. Poore, J.A. Barron, H.D.

Gower, and K. McDougall. 1978. Neogene biostra-

tigraphy of the Indian Creek—-Shell Creek area,

northern La Panza Range, California. Pp. 11-38 [in:]

Addicott, W.O., Neogene biostratigraphy of selected

areas in the California Coast Ranges. U.S. Geological

Survey Open-File Report 78-446 109 pp. [also

available in Blake, G.H., 1980, Pacific Section,

S.E.P.M 11-38],

. Addicott, W.O., P.D. Snavely, Jr., D. Bukry, and R.Z.

Poore., 1978. Neogene stratigraphy and paleontology

of southern Almeria Province, Spain; an overview.

U.S. Geological Survey Bulletin 1454: 1 —49.

. Addicott, W.O., G.R. Winkler, and G. Plafker. 1978.

Preliminary megafossil biostratigraphy and correla-

tions of selected stratigraphic sections in the Gulf of

Alaska Tertiary province. U.S. Geological Survey

Open-File Report 78-491 2 oversized sheets.

. Armstrong, A.K., P.D. Snavely, Jr., and W.O.

Addicott. 1978. Porosity evolution, late Miocene

reefs, Almeria Province, southern Spain. U.S. Geo-

logical Survey Open-File Report 78-940: 1 —20.

1979

a. Addicott, W.O. 1979. Oligocene molluscan biostra-

tigraphy of the Indians, Santa Lucia Range, central

California. Pp. 45 49 [in:] S.A. Graham, Tertiary and

THE NAUTILUS, Vol. 134, Nos. 3-4

Quaternary geology of the Salinas Valley and Santa

Lucia Range, Monterey County, California. Pacific

Coast Paleogeography Field Guide 4 (Pacific Section,

S.E.P.M.) 148 pp.

. Addicott, W.O. 1979. The marine Cenozoic of the

Pacific Coast states; an overview of the growth and

development of biostratigraphic research. Pp. 325

[in:] J.M. Armentrout, M.R. Cole, and H. TerBest,

Jr., Cenozoic paleogeography of the western United

States: Pacific Coast Paleogeography Symposium

(Pacific Section, S.E.P.M, Los Angeles, CA) 335 pp.

. Addicott, W.O., P.D. Snavely, Jr., R.Z. Poore, and D.

Bukry. 1979. La secuencia Nedégena marina de los

Campos de Dalias y de Nijar (Almeria). Estudios

geoldgicos 35(1/2):609-631.

. Armentrout, J.M., W.O. Addicott, J.A. Barron, and

K. McDougall. 1979. Pacific Coast Neogene corre-

lations; a progress report |[abs.]. Abstracts with

Programs, Geological Society of America 11(7): 381.

. Clark, J.C., E.E. Brabb, and W.O. Addicott. 1979.

Tertiary paleontology and stratigraphy of the central

Santa Cruz Mountains, California coast ranges.

Guidebook for Geological Society of America, Cor-

dilleran Section, 75‘" annual meeting, April 1979: 23.

1980

a. Addicott, W.O. 1980. Biostratigraphy of the marine

@

Neogene sequence at Cape Blanco, southwestern

Oregon. U.S. Geological Survey Professional Paper

Pi AG aod 320)

. Addicott, W.O. 1980. Biostratigraphy of the marine

Miocene sequence at Cape Blanco, southwest Ore-

gon [abs.]. Abstracts with Programs, Geological So-

ciety of America 12(3): 93.

. Addicott, W.O. 1980. Highlights in the 130-year

history of marine Cenozoic stratigraphic paleontol-

ogy on the Pacific Coast of North America. Pp. 1 -19

lin:] H. Igo, Professor Saburo Kanno Memorial

Volume, Tsukuba University Publications 177 pp.

. Addicott, W.O. 1980. Miocene stratigraphy and

fossils, Cape Blanco, Oregon. Oregon Geology 42(5):

87-97.

. Addicott, W.O. 1980. Status of the circum—Pacific

map project: a summary of the Circum—Pacific map

project meeting in Menlo Park, California, May 6-8,

1980. U.S. Geological Survey Open-File Report

S0-1050) hk 377.

f. Addicott, W.O. 1980. Project 114, biostratigraphic

datum-—planes of the Pacific Neogene. U. S. contri-

butions to the International Geological Correlation

Program; a record of the activities and scientific

contributions of United States participants,

1974-1979: 15-17,

. Addicott, W.O., and R.Z. Poore. 1980. Paleogene/

Neogene boundary in western North American se-

quences [abs.]: International Geological Congress,

Abstracts—Congres Geologique Internationale,

Resumes 26(1): 194.

C.L. Powell, II and K.J. Meade, 2020

h. Addicott, W.O., and R.Z. Poore. 1980. The

Paleogene-Neogene boundary in eastern North

Pacific marine sequences |[abs.]. Abstracts of

Papers—Pacific Science Association, XIV Pacific

Science Congress 2: 5-6.

. Armstrong, A.K., P.D. Snavely, and W.O. Addicott.

1980. Porosity evolution of upper Miocene reefs,

Almeria Province, southern Spain. American Asso-

ciation of Petroleum Geologists Bulletin 64(2):

188-208.

. Reinemund, J.A., W.O. Addicott, and P.W. Richards.

1980. Circum—Pacific Map Project [abs.]. U.S.

Geological Survey Professional Paper 1175: 335.

k. Stanley, S.M., W.O. Addicott and K. Chinzei. 1980.

Lyellian curves in paleontology: possibilities and

limitations. Geology 8(9): 422-426.

—"

md ©

1981

a. Addicott, W.O. 1981. Brief history of Cenozoic

marine biostratigraphy of the Pacific Northwest. Pp.

3 —16 [in:] J.M. Armentrout Pacific Northwest Ce-

nozoic Biostratigraphy. Geological Society of

America, Special Paper 184 172 pp.

b. Addicott, W.O. 1981. Significance of pectinids in

Tertiary biochronology of the Pacific Northwest. Pp.

17 -38 [in:] J.M. Armentrout Pacific Northwest

Cenozoic Biostratigraphy. Geological Society of

America, Special Paper 184 172 pp.

. Addicott, W.O. 1981. Mapping half the world;

Circum—Pacific map project; 7” annual meeting,

May 27-30, 1981, Menlo Park, California, U.S.A.

Episodes 1981(2): 48-49.

d. Nolan, T.B.., J.A. Reinemund, P.W. Guild, P.L.

Bateman, A.L. Clark, J.R. Hein, W.D. Carter, W.D.,

W.O. Addicott, R.G. Coleman, G.K. Czamanske, J.E.

Gair, R.B. Numan, G.M. Richmond, and R.P.

Sheidon. 1981. International Geological Correlation

Program. U.S. Geological Survey Professional Paper

L27ae 277-2.

e. Poore, R.S., J.A. Barron, and W.O. Addicott. 1981.

Biochronology of the northern Pacific Miocene. Pp.

91 -97 [in:| N. Ikebe, M. Chiji, R. Tsuchi, -Y.

Morozumi, T. Kawata, Proceedings of IGCP 114

international workshop on Pacific Neogene biostra-

tigraphy. 6th International working group meeting,

Osaka, Japan, Nov. 25-29, 1981 142 pp.

©)

1982

Reinemund, J.A., and W.O. Addicott. 1982. Circum-

Pacific map project: framework for international re-

sources assessment [abs.]. American Association of

Petroleum Geologists Bulletin 66(7): 982.

1985

Addicott, W.O., coordinator. 1985. Manganese nod-

ules, seafloor sediment, and sedimentation rates of the

Circum-Pacific region/Circum-Pacific Council for En-

ergy and Mineral Resources, Circum Pacific Map

Page 115

Project. American Association of Petroleum Geologists:

oversize map, scale 1:17,000,000.

1987

Moore, E.J., and W.O. Addicott. 1987. The Miocene

Pillarian and Newportian (Molluscan) stages of

Washington and Oregon and their usefulness in cor-

relations from Alaska to California. U.S. Geological

Survey Bulletin 1664: 1-13.

1988

Addicott, W.O., and Gryc, G. 1988. Scope and status of

the Circum-Pacific map project: a programmatic

overview including a resume of project activities during

1986 and 1987. U.S. Geological Survey Open-File

Report 88-215: 1 -110.

ACKNOWLEDGMENTS

Many people are thanked for sharing their memories of

Warren including Gene Coan, George Kennedy, Lindsey

Groves, Robert McLaughlin, Ellen Moore, Andrei Sarna-

Wojcicki, Judy Terry Smith, Richard Squires, Carol Sta-

dum, Richard Stanley, Paul Valentich-Scott, and John

Wehmiller. Special thanks are given to Eric Addicott,

Warren’s son, who shared the picture of Warren and of

growing up with him. Ashley Dineen (University of

California, Museum of Paleontology) is thanked for

providing down information about Warren’s time at

UCMP. Also thanked are Mary McGann and Richard

Stanley (USGS) for their helpful reviews.

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McDougall. 1980. Neogene biostratigraphy of the Indian

Creek-Shell Creek area, northern La Panza Range, Cal-

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Nikas, A.J., III. 1977. Description of a new bivalve of the genus

Macoma from the Pliocene of central California. The Ve-

liger 19(4): 434 -437.

Zinsmeister, W.]. 1983. New late Paleocene molluscs from the

Simi Hills, Ventura County, California. Journal of Pale-

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THE NAUTILUS 134(3—4):117-131, 2020

Page 117

Six new species of Paciocinebrina (Gastropoda: Muricidae:

Ocenebrinae) from the northeast Pacific

Shawn G. Wiedrick'

Roland Houart”

Natural History Museum of Los Angeles County | Royal Belgian Institute of Natural Sciences, Rue Vautier 29,

900 Exposition Blvd.

Los Angeles, California, 90007, USA

[email protected] and

1000 Brussels, Belgium and Institute of Systematics, Evolution,

Biodiversity (ISYEB)

Muséum national d’Histoire naturelle (MNHN), CNRS, SU,

EPHE, UA, CP 51, 57 rue Cuvier, 75005 Paris, France

roland. [email protected]

ADS TRACT

The western North American genus, Paciocinebrina Houart,

Vermeij, and Wiedrick 2019, is both impressively speciose and

phenotypically plastic, challenging the views of both lumpers and

splitters. A recent study has revealed that the northeastern

Pacific species of Paciocinebrina are distinct from the European

type species, Ocinebrina aciculata (Lamarck, 1822) and other

eastern Atlantic species, including Ocenebra Gray, 1847. Based

on newly collected specimens and material already available,

mostly from California, a spiral morphology comparison of six

new species are herein described, including habitat and distri-

bution records.

Additional Keywords: Gastropoda, Muricidae, Ocenebrinae,

Paciocinebrina, new species, northeastern Pacific Ocean

INTRODUCTION

The predatory snail genus Paciocinebrina Houart, Ver-

meij, and Wiedrick, 2019 has long been a grouping of

temperate snails chiefly restricted to the northeast Pacific

and similar in appearance to northeastern Atlantic and

Mediterranean species of Ocinebrina Jousseaume, 1880.

Molecular studies (Barco et al., 2017) support these

genera as distinct lineages, both having radiations of di-

versity. Houart et al. (2019) used morphological and

biogeographic comparisons to provide a faunal list and to

describe several new species from the southern range of

Paciocinebrina. Paciocinebrina is one of about 33 rec-

ognized genera within the muricid subfamily Ocenebrinae

Cossmann, 1903 (Molluscabase eds., 2020). Voucher

material examined range from intertidal to subtidal (to

686 m) habitats, and were reported from as far north as

the Kenai Peninsula, Kenai-Cook County, Alaska (59° N)

and as far south as Bahia Magdalena, Baja California Sur,

' Museum Associate

? Research Associate at both institutions listed

Mexico (24° N). Assuming that the LACM collections

examined accurately reflect distributional patterns, these

snails are much more abundant in the northern portion of

the range but are still present at moderate densities south

of San Luis Obispo County, California.

The diversity of this group has long been debated,

highlighted by the conflicted views of Radwin and

D’Attilio (1976) and Fair (1976). Shell and radular

morphology have been the basis of Paciocinebrina de-

scriptions (Radwin and D7’Attilio, 1971; 1976), with

morphological characters having a historical perspective,

embraced by malacologists in the diagnosis of molluscan

species and higher taxa in systematic studies (Ponder and

Lindberg, 1997).

The reproductive life histories of this group are poorly

understood and known from very few studies (Merle et al.,

2011: 23). Paleobiologists (Jablonski and Lutz, 1983) have

found that morphological features in the protoconch

sculpture of marine benthic gastropods are a result of

their mode of development (Bouchet and Strong, 2010:

65; Merle et al., 2011: 23). The use of this feature to

determine species delineation, biodiversity, and life his-

tory patterns has been broadly used for neogastropods

(Shuto, 1974; Bouchet, 1990), including studies on

muricids (Radwin and D’Attilio, 1976; D’Attilio, 1980;

1981; Myers and D’Attilio, 1986; Merle et al., 2011: 17,

23) and conoideans (Powell, 1966: 6; Bouchet, 1990). The

uniform possession of paucispiral protoconchs in eastern

Pacific Paciocinebrina species was noted by McLean

(1996: 80) and indicates that the mode of reproduction

across the taxon is by intracapsular development. Hansen

(1980), Jablonski and Lutz (1983), and Jablonski (1986)

found higher speciation and extinction rates in this mode

of development, which seems to parallel the radiation and

fossil record of this group (Marshall et al., 2012; Niitzel,

2014). Fluctuating sea level and temperature changes

likely contributed to higher periods of diversification,

whether in warm periods (Mayhew et al., 2012) or cold

periods (Davis et al., 2016), as reflected in the Neogene

Page 118

and Quaternary fossil record of California. Modes of larval

dispersal may have been under selection by the strong

' currents seen in the eastern Pacific, a model discussed by

Brown (2014).

Several contributing factors are likely the cause of

species within this genus to inhabit rocky substrate,

whether intertidal or subtidal. Ecological and physiolog-

ical factors such as reproduction, feeding, and ecological

niches are likely influenced by habitat selection by these

gastropods. Abbott (1968: 126) illustrated western At-

lantic muricid egg capsules, which appear to be similar in

appearance to northeastern Pacific Paciocinebrina spe-

cies. Laboratory studies by Spight et al. (1974: 234) de-

tailed egg capsule laying on rocks by Paciocinebrina

interfossa (Carpenter, 1864), with Griffith (1967: 69)

noting some eastern Pacific muricids known to guard eggs

during incubation. This reproductive process appears to

limit connectivity between populations, as long spanning,

intertidal and subtidal sandy shorelines tend to isolate

rocky reefs in the eastern Pacific, which could restrict

migration between these populations, isolating them to

rocky habitats. Diet, feeding preference, and food

abundance likely play a role in habitat selection. Species

within this group are carnivorous and feed by means of

radula and an accessory boring organ, which in combi-

nation, are capable of drilling through a prey’s hard flesh

or exoskeleton (Vermeij, 1993: 105, 108; McLean, 2007:

740; Bertsch and Rosas, 2016: 216-217). The preferred

diet of Paciocinebrina species is primarily comprised of

sessile barnacles, mollusks, and other invertebrates re-

stricted to rocky habitats.

Paciocinebrina species have been reported as feeding

or associated with the mantle of the gumboot chiton,

Cryptochiton stelleri (Middendorff, 1846) (Talmadge,

1975; Abbott and Haderlie in Morris et al., 1980: 277),

barnacles (Rice, 1971: 48; Abbott and Haderlie, 1980:

277; Palmer, 1988; McLean, 2007: 740; McLean, 1996:

78; Bertsch and Rosas, 2016: 216; Merle et al., 2011: 24),

limpets (Palmer, 1988; Bertsch and Rosas, 2016: 216),

bivalves (Palmer, 1988; McLean, 2007: 740; McLean,

1996: 78; Merle et al., 2011: 24), other mollusks (Palmer,

1988; Abbott and Haderlie, 1980: 277; Merle et al., 2011:

24) and the shelled polychaete worm, Spirorbis (Palmer,

1988: 192).

New species descriptions are based on the analysis of

e material from the California Academy of Sciences

(CASIZ), Natural History Museum of Los Angeles

County (LACM), San Diego Natural History Museum

(SDNHM), National Museum of Natural History,

Smithsonian Institution (USNM), University of Cal-

ifornia, Museum of Paleontology (UCMP), several private

collections, and the collection of subtidal and intertidal

specimens ranging from Devil’s Gate, Humboldt Co.

(40°N) to Laguna Beach, Orange Co., California (33°N).

A revision of Paciocinebrina is presented in Wiedrick

(thesis) including the chresonymy, synonymy, biodiver-

sity, biogeography, and ecology of known nominal taxa

based on existing literature, the aforementioned speci-

mens, and molecular data.

THE NAUTILUS, Vol. 134, Nos. 3-4

MATERIALS AND METHODS

INSTITUTIONAL ABBREVIATIONS

CASIZ: California Academy of Sciences, Invertebrate

Zoology, San Francisco, California, U.S.A

LACM: Natural History Museum of Los Angeles County,

Malacology Department, California, U.S.A.

NHMUK: Natural History Museum of the United

Kingdom, London, England, U.K.

SDNHM: San Diego Natural History Museum, San

Diego, California, U.S.A.

SGW: collection of Shawn G. Wiedrick

UCMP: University of California, Museum of Paleontol-

ogy, Berkeley, California, U.S.A.

USNM: National Museum of National History, Smith-

sonian Institution,Washington, D. C., U.S.A.

ZIN: Zoological Institute of Russian Academy of Sciences,

St. Petersburg, Russia

SPIRAL MORPHOLOGY ABBREVIATIONS

Terminology used to describe the spiral cords and the

internal denticles of the outer lip (based on Merle, 1999;

2001; 2005; Merle and Houart, 2003):

ab: abapical (or abapertural); abis: abapical infrasutural

secondary cord (on subsutural ramp); ABP: abapertural

primary cord on the siphonal canal; abs: abapertural

secondary cord on the siphonal canal; ad: adapical (or

adapertural); adis: adapical infrasutural secondary cord

(on subsutural ramp); ADP: adapertural primary cord on

the siphonal canal; ads: adapertural secondary cord on the

siphonal canal; IP: infrasutural primary cord (primary cord

on subsutural ramp); MP: median primary cord on the

siphonal canal; ms: median secondary cord on the siphonal

canal; P: primary cord; P1: shoulder cord; P2—P6: primary

cords of the convex part of the teleoconch whorl; s: sec-

ondary cord; s1-s6: secondary cords of the convex part of

the teleoconch whorl; SP: subsutural cord; t: tertiary cords;

D1-D6: abapical denticles; ID: infrasutural denticle.

RESULTS

SYSTEMATICS

Muricidae Rafinesque, 1815

Ocenebrinae Cossmann, 1903

Genus Paciocinebrina Houart, Vermeij and Wiedrick,

2019

Type Species: Tritonium (Fusus) luridum Middendorff,

1848: 244, by original designation, Recent, Sitka, Alaska

(Syntype, ZISP 62131).

Remarks: The complete faunal list and description was

provided by Houart et al. (2019: 209-210). Additional new

species are proposed below.

Paciocinebrina bormannae new species

(Figures 8-12)

S.G. Wiedrick and R. Houart, 2020 Page 119

11

Figures 1-12. Paciocinebrina species. 1-7. Paciocinebrina grandilurida new species. 1-2. Shelter Cove, California, intertidal on

rock, length 26.2 mm, width 13.5 mm (SGW 115). 3-4. Holotype, Shelter Cove, California, intertidal, length 27.0 mm, width 15.0 mm

(LACM 3761). 5. Spiral cords morphology. 6. Apertural denticles morphology. 7. Protoconch, scale bar: 1.0 mm (SGW 116). 8-12.

Paciocinebrina bormannae new species. 8—9. Holotype, Cayucos, California, on rocks at low tide, length 22.4 mm, width 12.3 mm,

(LACM 3760). 10. Spiral cords morphology. 11. Protoconch, scale bar: 1.0 mm (SGW 117). 12. Apertural denticles morphology.

Page 120

CHRESONYMY

Ocenebra lurida aspera.-Smith and Gordon, 1948: 189.

Ocenebra lurida.-_Abbott and Haderlie, 1980: 277-278,

ng, 13.76.

Ocinebrina foveolata.—Liff-Grieff, 2006: 4, lower right

figures.

Description: Large for genus, holotype 22.4 mm in

length, width 12.3 mm. Shell robust, thick, shape broad,

elongate-ovate, spire angle variable, siphonal canal nar-

row, short, acutely tapered, color rustic brown, with dark

brown incised lines between cords, aperture white, blue

overtones, siphonal canal creamy tan. Protoconch bulbous

with subtle carination, shoulder and median cord abruptly

formed, converged by vertical ribs of first teleoconch

whorl, clathrate in appearance, later cords strong, over-

riding ribs, fimbriations vaulted, close set, extremely fine

and low profile in interspaces. First teleoconch whorl with

P1, P2 spiral cords, second with P1 and P2 cords, starting

IP and-sl..cords,. third whorl swith IP: Pl: s1..P2 cords:

starting adis and s2 cords, fourth with adis, IP, P1, sl, P2,

s2 cords, starting P3 cord, final whorl with SP, adis, IP,

amis Ply SIL Pe S24 Peso PA S45 so, PG ADP..ads:

ms, MP, ABP and abs spiral cords, nine faint axial ribs on

final whorl. Apertural denticles morphology with ID and

six primary denticles, ID denticle extremely indistinct, D1

denticle weak, D2—D4 denticles stronger, final denticles

slightly weaker. Aperture elongate ovate, moderate in

size, lip of aperture thick, edge sharp, strongly projecting,

parietal wall weak posteriorly, thicker anteriorly of colu-

mellar wall, siphonal canal comparatively thin, tapered

towards anterior end.

Type Material: Holotype LACM 3760, length 22.4 mm,

width 12.3 mm.

Type Locality: Cayucos, San Luis Obispo County,

California, (35°26'40" N, 120°56'42" W), on rocks at low tide.

Other Material Examined: Two specimens, 12.1—38.1

m, Carmel Submarine Canyon, north end San Jose Creek

Beach, Monterey County, California (36°31'59" N,

121°55'59" W), collected by J.H. McLean, 1960-1964

[LACM 1960-24.105]; two specimens, Monterey, Mon-

terey County, California [LACM 151093]; four specimens,

intertidal on shale ledges, west of Cayucos Creek, San Luis

Obispo County, Calitormia (35°26'48" N, 120°54'29" Ww).

collected by P.I. LaFollette, 11 December 1977 [LACM

1977—112.48]; four specimens, Cayucos, San Luis Obispo

County, California, collected by Bormann [LACM 72468];

one specimen, intertidal, Dinosaur Cave, Shell Beach, San

Luis Obispo County, California (35°9'11" N, 120°40'36"

W), collected by P.M. Oringer, 19 December 1968 [LACM

1968—48.23]; six specimens from Shell Beach, San Luis

Obispo County, California (35°9'24" N, 120°40'36" W),

intertidal on shale ledges, collected by J.H. McLean,

1961-1963 [LACM 1961- 11.50]; six specimens, near

Arroyo Grande, San Luis Obispo County, California

[LACM 59600]; three specimens, San Luis Obispo County,

California [LACM 59620]; one specimen, 3.2 km north of

THE NAUTILUS, Vol. 134, Nos. 3-4

Avalon, Santa Catalina Island, California Channel Ids.,

California (33°23'59" N, 118°22'0" W), collected by J.H.

McLean, 29 April 1962 [LACM 1962-5.31].

Distribution: Carmel Submarine Canyon, Monterey Co.,

California (36°) to Avalon, Santa Catalina Island, Cal-

ifornia Channel Ids., California (33°N); primarily on low

intertidal rocks, large rock overhangs and undersides of

giant boulders, to 38.1 m.

Remarks: The spiral morphology of the initial whorl are

identical (P1 and P2) to various other Paciocinebrina

species, with distinct s2 cord in the second whorl of P.

bormannae, third whorl with adis cord, a feature not seen

in comparable species, fourth whorl similar to Pacioci-

nebrina munda (Carpenter, 1864), with P3 cord absent in

that species, final whorl of P. bormannae with SP, abis, s4,

ads, ms, abs cords and s6 cord absent, all characters not

seen in P. munda (Figures 56-59). Superficially similar to

the more northern species, P. grandilurida new species,

but is lighter in coloration, has a less acute spire, one ad-

ditional axial rib on final whorl and a different spiral cord

morphology with P. bormannae having an additional t cord

after s2 and ABP, abs cords on final whorl (Figures 5, 10).

Etymology: Named in honor of the late Mary Bormann, a

collector of various Paciocinebrina species, who also had a

particular passion for this group of west coast muricids.

Paciocinebrina grandilurida new species

(Figures 1-7)

Description: Shell large for genus, holotype length

27.0 mm, width 15.0 mm. Shape acute rhomboid, outer

lip projecting, thick, spire acute, siphonal canal moder-

ately long, color dark brown to black, interspaces lighter in

appearance, canal face white, aperture interior often with

blue overtone. Initial protoconch whorl low laying, bul-

bous, tabulation faint, shoulder and median cords well

established, ribs becoming well developed, first tele-

oconch whorl with overriding spiral cords, further whorls

forming large, thick, tightly set cords, final whorl surface

heavily scabrous, especially between cords, obscurely

clathrate. First teleoconch whorl with P1, P2 spiral cords,

second with Pl, P2 cords, starting IP cord, third whorl

with IP, Pl, P2 cords, starting s1, s2 cords, fourth with IP,

Pl, sl, P2, s2 cords, starting adis, abis cords, final whorl]

With SP eadis TPoabis..P beh Pa: 62: Po s8 Pa. G4 PS sh:

P6, s6, ads, ADP, MP, ms spiral cords, eight very faint,

broadly spaced axial ribs on final whorl. Aperture elongate

ovate, with ID, D1 split and five additional denticles,

columellar lip wide, recessed into previous whorl, siphonal

canal rather short, pseudoumbilicus faint.

Type Material: Holotype LACM 3761, length 27.0 mm,

width 15.0 mm.

Type Locality: Shelter Cove, Humboldt County, Cal-

ifornia, (40°1'24" N, 124°4'24" W) on rocks at low tide.

Other Material Examined: 12 specimens, intertidal,

near Machi Brothers Resort, Shelter Cove, Humboldt

S.G. Wiedrick and R. Houart, 2020

County, California (40°01'29" N, 124°04'00" W), collected

by P. I. LaFollette, 19-20 July 1970 [LACM 70-70.40];:

four specimens, intertidal on rocks, Van Damne State

Park, Little River, Mendocino County, California

(39°17'42" N, 123°47'48" W), collected by J.H. McLean,

22 February 1964 [LACM 64-8.34]; two specimens, in-

tertidal, Albion, Mendocino County, California (39°14'30"

N, 123°46'30" W), collected by J.H. McLean, 11 No-

vember 1962 [LACM 62-15.30]; two specimens, inter-

tidal, 0.8 km south of Fort Ross, Sonoma County,

California (38°30'42" N, 123°13'59" W), collected by (Pak

McLean, 28 December 1963 [LACM 63-57.23].

Distribution: Shelter Cove, Humboldt Co., California

(40°N) to Fort Ross, Sonoma Co., California (38°N); low

to mid intertidal zone, on rock reefs, undersurfaces of

large boulders and on the girdle of the gumboot chiton,

Cyptochiton stelleri (Middendorff, 1846).

Remarks: The first three whorls are identical to Pacio-

cinebrina munda (P1, P2; IP, P1, P2; IP, Pl, sl, P2, s2),

with abis cord of fourth whorl absent in P. munda. See P.

bormannae section for comparisons to this species. Tal-

madge (1975: 414), Abbott and Haderlie (1980) and

McConnaughey and McConnaughey (1985: 358) re-

ported the feeding of Paciocinebrina lurida (Figures

60-63) on the gumboot chiton, Cryptochiton stelleri

(Middendorff, 1846), although field observations from

Shelter Cove have exclusively been by the larger P.

grandilurida, despite the mass presence of P. lurida at

that locality.

Etymology: In reference to the large size and similarities

to the smaller Paciocinebrina lurida (Middendorff, 1848),

grandi- meaning grandiose, lurida in reference to that

other species, gender feminine.

Paciocinebrina mininterfossa new species

(Figures 13-18)

CHRESONYMY

Ocenebra interfossa.—Rice, 1971: pl. 17, fig. 98.

Type Material: Holotype LACM 3762, length 13.1 mm,

width 6.1 mm (ex SGW 114).

Type Locality: China Rock, Seventeen Mile Drive,

Monterey County, California, (36°36'10" N, 121°57'42"

W), on rocks at low tide.

Other Material Examined: Five specimens, Crescent

City, Del Norte County, California, collected by E. P.

Chace [LACM 72475]; two specimens, 18 m, Isle of St.

James, North Farallon Islands, California (37°45'40" N,

123°5'57" W), collected by R.W. Schmieder, R/V CORDELL

EXPLORER, 14 September 1991 [LACM 1991-177.19]; two

specimens, intertidal on rocks, south side of Point Joe,

Seventeen Mile Drive, Monterey County, California

(36°36'29" N, 121°57'29" W), collected by P.I. LaFollette,

June 1962 [LACM 1962-43.4]; 10 specimens, intertidal,

Pacific Grove, Monterey County, California (36°37'22" N,

121°54'34" W), collected by J.H. McLean, 1959-1964

Page 121

[LACM 1959-12.63]; one specimen, intertidal on Phyl-

lospadix roots, Hopkins Marine Station, Pacific Grove,

Monterey County, California (36°37'30" N, 121°54'00"

W), collected by J.H. McLean, 1 December 1962 [LACM

1962-16.17]; three specimens, Pacific Grove, Monterey

County, California [LACM 151066].

Distribution: Crescent City, Del Norte Co., California

(41° N) to China Rock, Seventeen Mile Drive, Monterey,

Monterey Co., California (36°N); primarily at low to mid-

intertidal zones on rocks, near dense mats of purple algae,

as noted on one LACM record from near the surfgrass,

Phyllospadix, 1 subtidal lot at 18 m.

Description: Shell small for the genus, holotype length

13.1 mm, width 6.1 mm. Shell elongate, profile narrow,

slenderly rhomboid, whorls ovate, tall, sutural indenta-

tions weak, size small, but robust, ribs subtle, scabrous

sculpture of even width throughout entire length of shell,

color dingy white, occasionally with light brown band on

P2 cord below shoulder, outer lip of aperture dull white,

columellar and parietal walls and interior flushed with

purplish brown. Spire acute, protoconch bulbous, worn in

nearly all specimens examined. Initial whorl somewhat

flattened, wide spanning, half set into proceeding whorl,

rotund, becoming shouldered by cord further into whorl

than most other Paciocinebrina species. Median cord

strong, slightly more projecting initially, becoming of

equal strength and pseudo-clathrate, spiral cords be-

coming projected nodes at intersection of axial ribs. Shape

of siphonal canal moderately small, tapering. First tele-

oconch whorl in holotype with P1, P2 cords, second with

Pl, P2 cords, starting IP cord, third with Pl, P2 cords,

starting, sl cord, fourth whorl with IP, P1, sl, P2 cords,

starting s2 cord, final whorl with SP, IP, Pl, sl, P2, s2, P3,

s3, P4, s4, P5, s5, P6, ADP, MP spiral cords and eleven

narrow, but widely spaced axial ribs. Aperture consistently

with weak D1, D2 (split), D3, D4, D5 denticles, strongest

centrally, split D2 denticles weak in small sized speci-

mens. Aperture moderately sized, ovate, posterior end

squarer in shape, lip heavily developed and thick at

subsutural ramp on final varices, slightly slimmer at

shoulder, thinning towards anterior end, parietal wall

extremely thin at top, wider anteriorly, slightly thicker at

columellar wall, terminated by a twist near anterior end.

Remarks: The initial three teleoconch whorls are iden-

tical to Paciocinebrina minor (Dall, 1919) (Figures

44-47), with P. mininterfossa having adis, abis, s1, s2,

s3, s4, s> and P6 cords on the fourth whorl, characters

absent in P. minor. This species is very small compared to

the similar appearing species, P. interfossa (Figures

64-67), of which many specimens have been haphaz-

ardly mixed into LACM lots labelled as such, a likely

assumption is that workers simply presumed a juvenile

state in this smaller species. Additionally, this taxon is a

dingy white, not brown, has thicker and much looser

sutural scabrocity and a different spiral morphology. A

specimen figured by Rice (1971), and identified as P.

interfossa, seems to match the holotype of this species and

enl22 THE NAUTILUS, Vol. 134, Nos. 3-4

Figures 13-18. Paciocinebrina mininterfossa new species. 13-14. China Rock, California, on intertidal rock, length 12.7 mm,

width 6.3 mm, (SGW 118). 15. Spiral cords morphology. 16-17. Holotype, China Rock, California, on intertidal rock, length 13.1 mm,

width 6.1 mm, (LACM 3762). 18. Apertural denticles morphology.

S.G. Wiedrick and R. Houart, 2020 Page 123

Figures 19-25. Paciocinebrina murphyorum new species. 19-20. Shell Beach, California, length 9.6 mm, width 5.6 mm (SGW

119). 21-22. Holotype, Jade Cove, California, on rock at 14.6 m, length 11.7 mm, width 7.2 mm, (LACM 3763). 23. Spiral cords

morphology. 24. Apertural denticles morphology. 25. Protoconch, scale bar: 1.0 mm (SGW 120).

Page 124

is here included, despite Rice (1971) neglecting to provide

a specific locality of the figured specimen. This species has

been observed in the field as feeding on Barleeia species.

Etymology: Gender feminine in the nominative case,

mini- meaning a miniature version of P. interfossa,

meaning having ditches among itself, in reference to the

sculpture of that species.

Paciocinebrina murphyorum new species

(Figures 19-25)

CHRESONYMY

Ocenebra interfossa clathrata.-Bormann, 1946: 39, pl. 4,

Beer Aon Oe

Description: Shell small for genus, holotype length

11.7 mm, width 7.2 mm. Shape rhomboid, severely

indented at whorl base, siphonal canal moderately long,

narrow, whorls tabulate, sculpture clathrate, of thick

intersecting cords and ribs, color chalky white with dark

brown bands at suture to mid-subsutural ramp, and

posteriorly before cords P3 through P4, aperture interior

usually brown or brownish purple, parietal wall lighter in

coloration, somewhat thin, twisted anteriorly. Initial

protoconch whorl upward projecting, rapidly downward

angled in shoulder cord, anteriorly tabulate, cords equal in

strength, rectangularly clathrate at rib intersections, deep

squarish pits on final whorl, early whorl growth lines faint,

crispate scales tightly spaced on final whorl, less evident in

pits. First two teleoconch whorls with P1, P2 cords, third

with P1, P2 cords, starting P3 cord, fourth whorl with adis,

TP. abis: Pl. sk P2282. PS. s8P4) 94’ Pb ADP: MPOABP

spiral cords, nine strong, widely spaced axial ribs, pro-

jecting at shoulder on final whorl. Aperture somewhat

small, ovate with D1—D4 denticles, outer lip projecting,

lip thickened, parietal wall closely attached, narrow,

siphonal canal long, spindle-like from basal indention.

Type Material: Holotype LACM 3763, length 11.7 mm,

width 7.2 mm.

Type Locality: Jade Cove, Big Sur, Monterey County,

Canforma.: (35°54'46""N.” 191°28'99"- WW) on .-roek at

14.6 m.

Other Material Examined: Two specimens, Roller Bay,

intertidal rocks, Hope Island, Vancouver Island, Rupert

District, Butish Columbia, Canada’ (50°55'36". N.

127°57'5" W), collected by J.H. McLean, 22 May 1963

[LACM 1963-31.22]; four specimens at 9-15 m, 100 m

inshore of buoy at Point Delgado, Humboldt County,

California (40°0'29" N, 124°04'00" W), collected by CC.

Swift, R/V SEARCHER, 29 July 1971 [LACM 1971-107.12];

one specimen at 0.3 m on rocks, northwest of Pico Creek,

San Simeon Village, San Luis Obispo County, California

(35°36'55" N, 121°9'4" W), collected by P.I. LaFollette, 8

August 2006 [LACM 2006-28.1]; three specimens,

Cayucos, San Luis Obispo County, California, collected by

R. and M. Bormann [LACM 72480]; six specimens, in-

tertidal on shale ledges, Shell Beach, San Luis Obispo

THE NAUTILUS, Vol. 134, Nos. 3-4

County, California (35°9'24" N, 120°40'36" W), collected

by J.H. McLean, 1961-1963 [LACM 1961-11.48]; 12

specimens, intertidal on shale ledges and boulder reef,

0.4 km northwest of South Point, Shell Beach, San Luis

Obispo County, California (35°9'18" N, 120°40'29" W),

collected by P.I. LaFollette, 10 December 1977 [LACM

77-111.44]; one specimen, intertidal, Avila Beach, San

Luis Obispo County, California (35°10'59" N, 120°43'59"

W), collected by P.M. Oringer, 28 July 1968 [LACM

1968—37.36]; two specimens, San Nicolas Island, Cal-

ifornia Channel Ids., California, MacGinitie collection,

17-19 July 1962 [LACM 59601].

Distribution: Hope Island, Vancouver Island, British

Columbia (50° N) to San Nicolas Island, California

Channel Ids., California (33° N); on and under intertidal

rocks, in gravel, rarely at shallow subtidal depths to 14.6 m.

Remarks: The initial three whorls are identical to

Paciocinebrina minor, with P. murphyorum having adis,

abis, sl, s2, s3 and s4 spiral cords on fourth whorl,

characters absent in P. minor. This species superficially

resembles P. atropurpurea (Carpenter, 1865) but has a

different morphology (Figures 52-55) than P. mur-

phyorum (Figures 23-24). Bormann (1946: pl. 4, fig. 1, 3)

figured a specimen of this species under the name cla-

thrata, now known as a synonym of P. atropurpurea

(Houart et al., 2019: 209).

Etymology: Named in the genitive case for the surname

Murphy, a name representing Brendan and William

Murphy, two individuals completely unknowingly of one

another, but coincidentally with same last name. Brendan,

a dear friend and high school colleague of the senior

author, facilitated the first visit to the type locality of this

species, a site where several specimens had subsequently

been collected and examined for this description. William

Murphy, a dear friend of the senior author’s father, was

gracious enough, along with wife Donna, to host Mr.

Wiedrick at their residence in Humboldt County, an

opportunity that afforded Mr. Wiedrick to conduct re-

search in northern California field sites. Specimens

sampled and identified as P. grandilurida trom Shelter

Cove, the city in which Mr. and Mrs. William Murphy

resides, is also the type locality of that new species, the

only location where this species was acquired and ana-

lyzed for spiral morphology.

Paciocinebrina pseudopusilla new species

(Figures 26-36)

CHRESONYMY

Ocenebra interfossa.-Radwin and D’Attilio, 1976: pl. 20,

Hee EY,

Description: Shell very small for genus, holotype worn,

length 7.5 mm, width 5.8 mm, paratype juvenile, length

5.4 mm, width 3.0 mm. Shell ovate rhomboid, siphonal

canal short, sculpture clathrate, axial ribs somewhat

narrow, cord thickness moderate, interspaces squarish,

large, color white to tan, pits slightly browner, occasionally

S.G. Wiedrick and R. Houart, 2020 Page 125

Figure 26-36. Paciocinebrina pseudopusilla new species. 26-27. Holotype, China Rock, California, length 7.5 mm, width

5.8 mm (LACM 3764). 28. Spiral cords morphology. 29. Apertural denticles morphology. 30-31. P. pseudopusilla, Neah Bay,

Washington, length 7.5 mm, width 4.3 mm (SGW 121). 32-33. Punta Popotla, Baja California, Mexico, length 7.6 mm, width 4.3 mm

(SGW 122). 34-35. Paratype, Jade Cove, California, on rock at 14.6 m, length 5.4 mm, width 3.0 mm, (LACM | 3766). 36. Protoconch,

scale bar: 1.0 mm.

Page 126

with sienna brown blotches on IP cord and/or Pl cord

white, rarely one brown thin band on base, interior of

aperture almost exclusively with dark brown band at

cords. Spire moderately blunt, apex relatively blunt,

tabulate, protoconch large, broad, whorl carination strong,

shoulder and median cord almost equal in strength, large,

distinct, slowly and subtly transitioning to clathrate

sculpture, cords more spaced than in P. pusilla, scabrocity

less evident than that species. First teleoconch whorl with

P1, P2 spiral cords, second with P1, P2 cords, starting IP

cord, third whorl with IP, P1, P2 cords, fourth with IP, P1,

P2, P3, P4, P5, ADP, MP spiral cords, final whorl with 15

narrow, moderately spaced axial ribs. Aperture elongate

ovate, moderately small, parietal and columellar wall

extremely thin, canal rarely fused. Apertural denticle

morphology somewhat weak, consisting of D1—D5.

Type Material: Holotype LACM 3764, length 7.5 mm,

width 5.8 mm; paratype LACM 3766, length 5.4 mm,

width 3.0 mm (Jade Cove, Monterey County, California

(35°54'46" IN, 121°28'29" W). on rocks at 14:6 -m.

Type Locality: China Rock, Seventeen Mile Drive,

Monterey County, California, (36°36'10" N, 121°57'42"

W) on intertidal rocks.

Other Material Examined: Three specimens, in beach

drift, Makah Bay, Clallam County, Washington, collected

by Tom Rice [LACM 182327]; one specimen, Anchor

Bay, Mendocino County, California, collected by D.

Brown, July 1958 [LACM 61635]; one specimen, inter-

tidal, Waddell Beach, Santa Cruz County, California

(37706'00" oN, .122°10'50" WV): collected zby. Elalda

McLean, 1967-1968 [LACM 1967-95.60]; eight speci-

mens at 18 m, Isle of St. James, North Farallon Islands,

California (37°45'40" N, 123°05'57" W), collected by R.W.

Schmieder, R/V CORDELL EXPLORER, 14 September 1991

[LACM 1991-177.23]; 23 specimens, intertidal on rocks,

Carmel Point, Monterey County, California (36°31'00" N,

121°57'0" W), collected by J.H. McLean, 13-15 October

1981 [LACM 1981-47.25]; three specimens at 12.1-38.1

m, Carmel Submarine Canyon, north end San Jose Creek

Beach, Monterey County, California (36°31'59" N,

121°55'59" W), collected by J.H. McLean, 1960-1964

[LACM 1960-24.106]; two specimens, intertidal on shale

ledges and boulder reef, 0.4 km northwest of South Point,

Shell Beach, San Luis Obispo County, California

(35°09'18" N, 120°40'29" W), collected by P. I. LaFol-

lette, 10 December 1977 [LACM 77-111.45]; one

specimen, intertidal to 7.6 m, 0.48 km southeast of Bay

Point, San Miguel Island, California Channel Ids., Cal-

ifornia (34°01'59" N, 120°17'59" W), collected by J.H.

McLean, 21-22 August 1967 [LACM 1967—38.57]; one

specimen, intertidal, Forney Cove, Santa Cruz Island,

California Channel Ids., California (34°03'29" N,

119°55'00" W), collected by J.H. McLean, 15 March 1969

[LACM 1969-11.37]; two specimens at 23-27 m, on

granite pinnacles, Wilson Rock, San Miguel Island, Cal-

ifornia Channel Ids., California (34°06'24" N, 120°23'41"

W), collected by J.H. McLean, 31 May 1982 [LACM

THE NAUTILUS, Vol. 134, Nos. 3-4

1982-59.33]; 20 specimens at 9.1—30.4 m, Isthmus Cove,

Santa Catalina Island, California Channel Ids., California

(33°26'30" N, 118°28'59" W), collected by J.H. McLean,

June-July 1971 [LACM 71-99.33]; six specimens at

9. 1—21.3 m, Santa Barbara Island, California Channel Ids.,

California (33°28'59" N, 119°1’30" W), collected by J.H.

McLean and J. Margetts, 8 July 1972 [LACM 72-97.38];

13 specimens, San Pedro, Los Angeles County, California,

collected by R.H. Tremper [LACM 59613]; five speci-

mens, Laguna Beach, Orange County, California [LACM

17420]; two specimens from Reef Point, Orange County,

California [LACM 17431]; three specimens, intertidal,

south side of Punta Banda, Baja California, Mexico

(31°43'36" N, 116°43'00" W), collected by ].H. McLean,

30 November 1963 [LACM 63-55.31]; 20 specimens at

15.2-30.4 m, Ben Rock, near Isla San Martin, Baja

California, Mexico (30°25'59" N, 116°07'00" W), collected

by J.H. McLean, 23 September 1972 [LACM 72-112.37];

five specimens at 13.7 m, northwest end of Isla Cedros,

Baja California, Mexico (28°21'00" N, 115°14'48" W),

collected by J.H. McLean, 24 September 1972 [LACM

72-114.36]; one specimen, intertidal, Punta Rompiente,

Baja California, Mexico (27°43'22" N, 115°00'06" W),

collected by J.H. McLean and P. I. LaFollette, R/V

SEARCHER, 21 October 1971 [LACM 71-162.43].

Distribution: Makah Bay, Clallam Co., Washington (48°

N) to Punta Rompiente, Baja California Sur, Mexico (27°

N): intertidal and subtidal zone on rocks to 125 m.

Remarks: The initial three teleoconch whorls are similar

to Paciocinebrina gracillima (Stearns, 1871) (Figures 48-51),

but P. pseudopusilla is absent of secondary cords. The

general morphology is most similar to P. pusilla, protoconch

features, spiral and denticle morphology (Figures 28-29,

39-40) are distinguishable characters in separating these

species, considering additional characteristics is problematic

and usually lend to improper identification. Protoconch of P.

pusilla small, bulbous, highly erect, keel indistinct, P. pseu-

dopusilla protoconch large, profile wide, tabulate, keel dis-

tinct, D6 cord present in P. pusilla, exclusively absent in P.

pseudopusilla. Superficially similar to P. pusilla, initial two

whorls identical, third whorl with P3 cord present in P.

pusilla, absent in P. pseudopusilla, fourth whorl of P. pusilla

with P6, s6 and ABP cords, D6 apertural denticle, and one

fewer axial rib, characters not seen in P. pseudopusilla.

Etymology: Gender feminine, pseudo- meaning spurious

and pusilla in reference to that morphologically similar

species in the nominative case.

Paciocinebrina pusilla new species

(Figures 37-43)

Description: Shell small for genus, holotype length

9.6 mm, width 5.2 mm. Shell elongate, rhomboid,

sculpture clathrate, ribs somewhat tight, spiral cords thick,

interspaces small, condensed, color white to dingy, gray

brown, rarely with sienna brown blotches on IP cord, P1

cord often white, occasionally P3 cord with faint brown

marks, interior of aperture dark purplish brown or white.

S.G. Wiedrick and R. Houart, 2020 Page 127

Figures 37-43. Paciocinebrina pusilla new species. 37-38. Holotype, Point Estero, California, on intertidal rocks, length 9.6 mm,

width 5.2 mm (LACM 3765). 39. Spiral cords morphology. 40. Apertural denticles morphology. 41-42. Paratype, Cayucos, California,

on intertidal rocks, length 8.8 mm, width 4.9 mm (LACM 3767). 43. Protoconch, scale bar: 1.0 mm (SGW 123).

Page 128 THE NAUTILUS, Vol. 134, Nos. 3-4

Figures 44-55. Paciocinebrina species. 44-47. Paciocinebrina minor (Dall, 1919). 44-45. Holotype, Santa Catalina Island, Cal-

ifornia, length 7.3 mm, width 3.7 mm (USNM 56912). 46. Apertural denticles morphology. 47. Spiral cords morphology. 48-51.

Paciocinebrina gracillima (Stearns, 1871). 48-49. Syntype, at 18.2 m, San Diego, California, length 13.4 mm, width 7.4 mm (USNM

46920). 50. Apertural denticles morphology. 51. Spiral cords morphology. 52-55. Paciocinebrina atropurpurea (Carpenter, 1865).

52-53. Lectotype, Neah Bay, Washington, length 14.0 mm, width 7.6 mm (USNM 15528b). 54. Apertural denticles morphology. 55.

Spiral cords morphology.

S.G. Wiedrick and R. Houart, 2020 Page 129

| ww

ADP MP ABP

Figures 56-67. Paciocinebrina species. 56-59. Paciocinebrina munda (Carpenter, 1864). 56-57. Holotype, Santa Catalina Island,

California, length 16.4 mm, width 8.3 mm (USNM 46708). 58. Apertural denticles morphology. 59. Spiral cords morphology. 60-63.

Paciocinebrina lurida (Middendorff, 1848). 60-61. Syntype, Sitka, Alaska, length 18.0 mm, width 10.5 mm (ZISP 62131). 62. Apertural

denticles morphology. 63. Spiral cords morphology. 64-67. Paciocinebrina interfossa (Carpenter, 1864). 64—65. Holotype, Monterey,

California, length 18.8 mm, width 9.3 mm (USNM 4636). 66. Apertural denticles morphology. 67. Spiral cords morphology.

Page 130

Spire acute, protoconch small, relatively erect, initially

projecting upward, bulbous, keel weak, shoulder and

median cord almost equal in strength, intersected im-

mediately by clathrate sculpture transitioning to pustu-

lated intersections with evenly-spaced ribs, scabrocity

highly prominent on subsutural ramp, between cords and

on siphonal canal. First two teleoconch whorls with IP, P1,

P2 spiral cords, third with IP, P1, P2 cords, starting SP

cere. fourth whorl wit [PPPs P23) P4- Po PG. 66:

ADP, MP, ABP cords, 16 narrow, but tightly spaced axial

ribs on final whorl. Aperture elongate ovate, moderately

large, somewhat restricted, parietal and columellar wall

extremely thin, canal almost exclusively fused. Denticle

morphology absent or very weak, only detectable in fully

matured specimens, D1—D6 denticles thereafter.

Type Material: Holotype LACM 3765, length 9.6 mm,

width 5.2 mm; one paratype LACM 3767 (Cayucos, San

Luis Obispo County, California (35°26'50" N, 120°55'14"

W), on intertidal rocks).

Type Locality: Point Estero, San Luis Obispo County,

Califomia (35°27'34" N, 120°58'15" N), on intertidal rocks.

Other Material Examined: One specimen, intertidal on

rocks, Carmel Point, Monterey County, California

(36°31'0" N, 121°57'0" W), collected by J.H. McLean,

13-15 October 1981 [LACM 81-47.26]; four specimens,

Pacific Grove, Monterey County, California, collected by

A. G. Smith, 1910 [LACM 182324]; two specimens,

Monterey, Monterey County, California, Falkenthal

collection [LACM 182323]; three specimens, Monterey,

Monterey County, California [LACM 182325]; five

specimens, Monterey, Monterey County, California,

collected by Howard Hill [LACM 182326]; one specimen,

siftings from Phyllospadix roots, Hopkins Marine Station,

Pacific Grove, Monterey County, California (36°37'30" N,

121°54'00" W), collected by J.H. McLean, 1 December

1962 [LACM 62-16.20]; one specimen, intertidal, Di-

nosaur Cave, Shell Beach, San Luis Obispo County,

California (35°09'01" N, 120°40'36" W), collected by P.M.

Oringer, 19 December 1968 [LACM 68-48.24]; two

specimens, intertidal, near E] Cortez Hotel, north of

Ensenada, Baja California, Mexico (31°52'30" N,

116°40'48" W), collected by J.H. McLean and P.M.

Oringer, 19 December 1964 [LACM 64-32.40]; one

specimen, intertidal on rock ledges and boulders, 1.6 km

south of Puerto Santo Tomas, Baja California, Mexico

(31°34'59" N, 116°40'00" W), collected by J.H. McLean

and P.M. Oringer, 4 January 1966 [LACM 66-1.47]; six

specimens, intertidal, cement plant halfway between

village and beach at Puerto Santo Tomas, Baja California,

Mexico (31°33'00" N, 116°40'00" W), collected by J.H.

McLean, 8-10 January 1967 [LACM 67-2.58].

Distribution: Carmel Point, Monterey Co. California

(36° N) to south of Puerto Santo Tomas, Baja California,

Mexico (31° N); chiefly intertidal on rocks. This species

seems to inhabit cold waters to the north and reappears in

the cold upwelling regions of northern Baja California.

THE NAUTILUS, Vol. 134, Nos. 34

Remarks: The first whorl with visible IP, Pl and P2 cords

which is identical to other Paciocinebrina species, second

whorl identical to P. lurida and P. interfossa, fourth and

final whorl of P. pusilla lacking secondary cords not

present in the other two species. See P. pseudopusilla

section for comparisons of this similar species. This

species is commonly confused as juvenile specimens of

Paciocinebrina atropurpurea and P. interfossa.

Etymology: Gender feminine, pusilla meaning little in

the nominative case.

ACKNOWLEDGMENTS

We graciously thank Lindsey T. Groves (Collections

Manager, Malacology Department, LACM) for access to

the museum’s collection, assistance with LACM voucher

numbers, permission to analyze LACM material and

essential literature at the LACM library, and for his

suggestions that greatly improved the initial manuscript.

Bernard Garrigues was additionally instrumental in pro-

viding constructive remarks which further refined the

development of this work. We are also especially grateful

to Elizabeth Kools (CASIZ), Erica Clites and Dave Strauss

(UCMP), Andreia Salvador (NHMUK), Boris Sirenko

(ZIN), and Ellen Strong (USNM) for either allowing

access to respective collections or for sending images of

specimens essential to the completion of this paper.

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THE NAUTILUS 134(3-4):132-137, 2020

Page 132

A new species of Arca (Bivalvia: Arcidae) from the lower Miocene

Asahi Formation on the Japan Sea side of central Honshu, with

remarks on the westward faunal migration from the eastern Pacific

Hiroshi Kurita

Department of Geology

Faculty of Science

Niigata University

Niigata 950-2181, JAPAN

Kazutaka Amano

Department of Geoscience

Joetsu University of Education

1 Yamayashiki

Joetsu 943-8512, JAPAN

ABSTRACT

The arcid bivalve, Arca (Arca) budoensis new species, is de-

scribed from the Budo Mudstone Member of the Asahi For-

mation in northern Niigata Prefecture, central Honshu. The age

of the member has been assigned to 16.6-15.9 Ma (late early

Miocene) on the basis of dinoflagellate cysts. Judging from the

arcid, dinoflagellate cysts and sedimentary facies, the paleo-

environment of the member was a shallow embayment influenced

by a warm-water current. The reason that the molluscan fauna

of the Budo Member contains no characteristic species of the

subtropical to tropical Arcid-Potamid fauna (17.0-16.7 Ma) is that

the Budo fauna is slightly later than that Arcid-Potamid fauna.

Because the new species resembles fossil and Recent species of

the eastern Pacific, A. budoensis is accepted as derived from an

ancestor that migrated westward to Asia. The senior author,

Kazutaka Amano, is the single author of the new species.

Additional Keywords: Marine, fossil, shallow sea, warm-current

INTRODUCTION

The Japan Sea was formed in the latest Oligocene (ca. 25

Ma) by separation of the Japanese Islands from the

Eurasian continent (Yanai et al., 2010). The oldest “ma-

rine” trace fossils from the Japan Sea side were recovered

from the Shiose-no-Misaki sediments which were in-

truded by a dolerite dated at 20 Ma (Ohguchi et al., 2005).

However, Sato et al. (2009) argued that these fossils were

non-marine. According to Sato et al. (1991, 2009), the

oldest marine fossils including the nannofossil, Spheno-

lithus heteromorphus were from the lower part of the

NN4 zone (Martini, 1971), later than 17.75 Ma according

to Backman et al. (2012).

In the northern part of Niigata Prefecture, the marine

sediments occur in the Miocene Asahi Formation

(Nishida, 1958: Takahama et al., 1976), which uncon-

formably overlies the Tenjosan Formation and is overlain

by the Osudo Shale containing the Osudo flora (Kamoi

et al., 1978). This flora corresponds with the late early to

early Middle Miocene Daijima-type flora (Takahama

et al., 1976; Kamoi et al., 1978). The Asahi Formation

consists of the Budo Mudstone, Nagasakatoge Rhyolite

and Arasawa Sandstone and Conglomerate Members in

ascending order (Takahama et al., 1976). The large for-

aminifer Operculina complanata japonica Hanzawa, from

the Arasawa Member indicates a late early to early late

Miocene age (Kamoi et al., 1978).

The following mollusks and brachiopods (including

Terebratulina spp.) were recorded from the Budo

Member by Nishida (1958), Tsuda (1965) and Taka-

hama et al. (1976): Chlamys iwasakiensis, Pecten sp.,

Cardium sp., Geloina yamanei, Panomya simotomensis,

and Littolinopsis miodelicatula. Of these, Geloina

yamanei and Littolinopsis miodelicatula are charac-

teristic species of late early Miocene mangrove swamp

fauna (Oyama, 1950). However, these species names

were only listed up from unknown localities and have

never been described nor illustrated. Moreover, the

exact age of this member has not been determined from

microfossils.

Many marine fossils have been recovered from a

previously unknown locality in the Budo Member. The

species composition including a new arcid species 1s very

different from the above listed species. In this paper, we

determine the age by dinoflagellate cysts, describe the

new bivalve species of Arca and discuss the paleobio-

logical significance of the fauna.

MATERIALS AND METHODS

The fossils were collected from a small outcrop along a

rice field at 250m east from Budo (Figure 1; 38°23'23" N,

139°33'27” E). At the fossil locality, hard gray mudstones

yielding many shell-dissolved fossils are exposed. From

this locality, the following molluscan species are recorded:

Arca budoensis new species, Arcuatula? sp. and Cav-

ilucina? sp. Moreover, the fossils also include three

species of brachiopods: Discinisca sp., Coptothyris grayi

(Davidson), Terebratalia sp. and one fragment of Cirri-

pedia, Capitulum? sp.

kK. Amano and H. Kurita, 2020

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Page 133

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Figure 1. Locality of fossils. Base map from “Budo”, original scale 1:25,000; topographical map published by the Geospacial In-

formation Authority of Japan.

Dinoflagellate cysts were picked for age determination

from the rock subsampled in the mollusk-bearing mud-

stone. Taxonomic identification of dinoflagellate cysts

follows Fensome et al. (2008), where complete biblio-

graphic references were provided. The sample was treated

successively with HCl and HF to eliminate carbonate and

silicate minerals. Then heavy liquid zinc bromide (specific

gravity 2.0) was used to concentrate organic particles from

the residues after the acid treatment.

We used digital calipers (Mitsutoyo Company, model

CD-20) to measure specimens of Arca to the first decimal

place. The terminology on Arca is follows Noda (1966). All

specimens of Arca are deposited at the National Museum

of Nature and Science, Tsukuba (NMNS).

DINOFLAGELLATE AGE

The sample yielded a number of dinoflagellate cysts

whose preservation was sufficiently good for identifica-

tion. Based on 393 counted specimens, the dinoflagellate

cyst assemblage is characterized by abundant to common

Page 134

occurrences of Achomosphaera ramulifera, A. spongiosa,

Cribroperidinium giuseppei, C. granomembraneceum,

Diphyes latiusculum, Heteraulacacysta campanula,

Hystrichokolpoma_ rigaudiae, Lingulodinium machaer-

ophorum, Operculodinium centrocarpum, Spiniferites

pseudofurcatus and Systematophora placacantha with

fewer Nematosphaeropsis lemniscata, Reticulatosphaera

actinocoronata, and Tuberculodinium vancampoae. In the

assemblage, protoperidinioid species are very minor in

abundance and include Brigantedinium sp., Lejeunecysta

spp., and Selenopemphix nephroides (Table 1). In addition,

a few specimens of an acritarch species Paralecaniella

indentata and a freshwater green alga Pediastrum sp. were

also recorded.

The abundant to common occurrences of Cribroper-

idinium giuseppei, C. granomembraneceum, Diphyes

latiusculum, Spiniferites pseudofurcatus and System-

atophora placacantha indicate evident correlation with

the basal part of the Subzone b of Diphyes latiusculum

Zone originally proposed by Matsuoka et al. (1987) and

subsequently modified by Obuse and Kurita (1999).

According to Obuse and Kurita (1999), this part is coeval

Table 1. List of dinoflagellate cysts and acritarchs associated

with the molluscan fossils. Relative abundance of each dino-

flagellate cyst taxon is expressed as VA (very abundant, 20 % and

more of the total specimen count), A (abundant, 20-10 %), C

(common, 10-3 %), R (rare, 3-1 %) and VR (very rare, less than 1

%).

Species Abundance

DINOFLAGELLATA

Achomosphaera ranulifera C

Achomosphaera spongiosa C

cf. Achomosphaera spongiosa R

Batiacasphaera ? spp. VR

Brigantedinium sp. VR

Cleistosphaeridium ancyrea VR

Cribroperidinium giuseppei Cc

Cribroperidinium granomembraneceum C

Diphyes latiusculum ®

Heteraulacacysta campanula R

Hystrichokolpoma rigaudiae R

Impagidinium sp. VR

Lejeunecysta spp. VR

Lingulodinium machaerophorum c

Lingulodinium sp. VR

Nematosphaeropsis lemniscata VR

Operculodinium centrocarpum CG

Reticulatosphaera actinocoronata VR

Selenopemphix nephroides VR

Spiniferites membraneceus R

Spiniferites pseudofurcatus C

Spiniferites ramosus R

Spiniferites sp. A

Systematophora placacantha V

Tuberculodinium vancampoae R

Other organic algal microfossils

Paralecaniella indentata

Pediastrum sp.

present

present

THE NAUTILUS, Vol. 134, Nos. 3-4

with the diatom Denticulopsis praelauta Zone (NPD3B)

that is calibrated to the age interval of 16.6—15.9 Ma (latest

early Miocene) by Yanagisawa and Akiba (1998) and

Watanabe and Yanagisawa (2005).

SYSTEMATICS

Family Arcidae Lamarck, 1809

Subfamily Arcinae Lamarck, 1809

Genus Arca Linnaeus, 1758

Subgenus Arca Linnaeus, 1758

Type Species: Arca noae Linnaeus, 1758 by subsequent

designation.

Remarks: The senior author, Kazutaka Amano, is

the single author of the new species. Reinhart (1935)

recognized four subgenera: Arca, Litharca Gray, 1842,

Arcoptera Heilprin, 1887, and Eonavicula Arkell, 1929.

According to Lutaenko and Maestrati (2007), Litharca is

an independent genus and Arcoptera is independent from

Arca (s.s.). Oliver and Chesney (1994) recognized Tet-

rarca Nordsieck, 1969 (type species: Arca tetragona Poli,

1795). When Oliver and Holmes (2006) subdivided Arca

into A. noae, A. avellana, and A. tetragona groups, they

did not treat the last group as a separate subgenus.

Consequently, Lutaenko and Maestrati (2007) separated

the extinct Arcoptera as a subgenus and Arca (s. s.) into

three types: Type 1 ( A. avellana, A. boucardi, A. tetra-

gona), Type 2 (A. navicularis, A. pacifica, A. zebra) and

Type 3 (A. koumaci), based on the presence of postero-

dorsal wing and posterior sulcus. Huber (2010) sub-

divided the modern Arca into four subgenera as Arca

(s. s.), Tetrarca, unnamed I (A. avellana, A. imbricata, A.

mutabilis, A. ventricosa) and unnamed II (A. boucardi).

From the view point of valve margins, Vermeij (2013)

subdivided Arca into three groups as A. imbricata group

with smooth edge (A. avellana, A. imbricata, A. mutabilis),

A. zebra group with obsolete ventral crenulations and

well-developed anterior and posterior ones (A. zebra, A.

pacifica, A. navicularis, A. ventricosa, A. noae). Vermeij

(2013) separated A. boucardi from these two groups by its

continuously crenulated ventral margin and a much

narrower hinge plate. Molecular data by Feng et al. (2015)

and Kong et al. (2020) support the subdivision of Arca (s.

s.) by Oliver and Holmes (2006) and Lutaenko and

Maestrati (2007) except for the separation of A. boucardi

as a subgenus level supporting Huber’s (2010) and Vermeij’s

(2013) opinions. In this paper, the new species is treated as a

A. noae group (Subgenus Arca s. s.) from its shell shape.

Arca (Arca) budoensis Amano new species

(Figures 2-6)

Diagnosis: Small Arca having low elongate shell, pointed

anterior end, truncated posterior margin, strong radial

ridge from umbo to postero-ventral corner and posterior

sulcus. Surface ornamented with 64 radial ribs and granu-

lated with growth lines on posterior part.

kK. Amano and H. Kurita, 2020

Page 135

Figures 2—6. Arca (Arca) budoensis Amano new species. 2. Para

Description: Shell small (to 25.8 mm long), elongate

quadrate, much lower than high especially in adults (H/

L= 0.33 to 0.56), inequilateral (AL/L= 0.27 to 0.41),

strong radial ridge extending from umbo to postero-

ventral corner. Antero-dorsal margin straight and

horizontal; anterior end of dorsal margin pointed; antero-

ventral margin posteriorly oblique; middle to posterior

ventral margin broadly concave as byssal notch in adult

but straight in younger specimens (length <7.3 mm);

postero-ventral corner acutely rounded; posterior margin

subtruncated and concave; postero-dorsal margin straight

and horizontal making right angle with posterior margin.

Umbo produced above dorsal margin; beak located at

anterior about one-third to two-fifths. Surface of anterior

part of shell in front of radial ridge ornamented with 51

flat and fine radial ribs separated by nearly equal inter-

spaces; anterior eight stronger than other ribs; posterior

area of shell behind radial ridge sculptured by 13 radial

ribs, lamellated growth ribs making granules at their

crossing points; granulation sometimes seen in front of

radial ridge in younger shells. Small taxodont teeth ob-

servable in terminal of hinge. Inner structure unknown.

Type Material: Holotype: Left valve (NMNS PM no.

65046), length 25.8 mm, height 8.6 mm, anterior length

7.6 mm. Paratypes: Left valve (NMNS PM no. 65047), length

19.6 mm, height 7.7 mm, anterior length 8.0 mm; left valve

(NMNS PM nos. 65048), length 13.8 mm, height, 5.0 mm-+;

left valve (NMNS PM no. 65049), length 7.3 mm, height

4.1 mm, anterior length 2.9 mm; right valve (NMNS PM no.

65050), length 3.7 mm, height 1.9 mm, anterior length 1.0 mm.

Type Locality: 250m east from Budo, Murakami City; upper

lower Miocene Budo Mudstone Member of Asahi Formation.

Material Examined: Twenty-eight specimens from the

type locality.

e, left valve, NMNS PM 65048. 3. Paratype, right valve,

NMNS PM 65050. 4. Paratype, left valve, NMNS PM 65049. 5. Holotype, left valve, NMNS PM 65046. 6. Paratype, left valve,

NMNS PM 65047. Scale bars = 5 mm.

Remarks: From the point of view of shell shape, there is

no similar modern or fossil species in Japan. Arca sp. from

the uppermost lower Miocene Kubohara Formation in

Gifu Prefecture by Itoigawa et al. (1981, 1982) has a

similar shell size and an elongate shell. However, Arca

(Arca) budoensis new species has a pointed anterior

end and lower shell than Arca sp.

Arca (Arca) budoensis new species can be included

in the A. noae group of Oliver and Holmes (2006) and

Type 2 by Lutaenko and Maestrati (2007) because of its

wing shape and posterior sulcus. In the western Pacific,

there is no fossil record other than the modern species,

Arca (Arca) navicularis Bruguiére, 1789 belonging to the

same group and type. However, the oldest record of A.

(A.) navicularis is from the Pliocene in Indonesia (Kase

et al., 2008). Arca (Arca) budoensis new species differs

from A. (A.) navicularis by having a lower shell, no

pointed postero-dorsal end and finer radial ribs. In con-

trast, some similar species have been described from the

eastern Pacific. Arca (Arca) cf. hawleyi Reinhart, 1943

from the Eocene Tejon Formation in California, is sim-

ilar to the new species by having a similar size elongated

shell with a pointed anterior end and a strong ridge from

beak to postero-ventral corner. However, Arca (Arca)

budoensis new species has a more posteriorly situated

beak and more distinct radial ribs. A Recent species, Arca

(Arca) pacifica (Sowerby, 1833) from Baja California

to the Galapagos Islands (Coan and Valentich-Scott, 2012)

is similar to the new species in having pointed anterior

end and a concave area behind the strong ridge from

beak to postero-ventral margin. However, Arca (Arca)

budoensis new species has a lower elongate-quadrate

shell, no pointed posterior end, shallower byssal notch and

more numerous fine radial ribs and is much smaller than

A. (Arca) pacifica. Another similar species is Arca (Arca)

truncata (Sowerby, 1833) from San Lucas, Baja California

Page 136

to the Galapagos Islands (Coan and Valentich-Scott,

2012). It resembles Arca (Arca) budoensis new spe-

cies in its low elongate quadrate outline. Arca (Arca)

budoensis new species has a pointed antero-dorsal end,

a posterior sulcus and an oblique posterior margin which

are never observed in the much larger A. (A.) truncata.

Distribution: Only from the type locality.

Etymology: Named for locality from where this species

was collected.

DISCUSSION

The present paper is the first to describe and illustrate

fossils from the Budo Member. Arca (Arca) budoensis

new species and its associated species indicate a

shallow-marine environment. The occurrences of the

dinoflagellate cysts Lingulodinium machaerophorum (cold-

intolerant/thermophilic) and Tuberculodinium vancam-

poae (tropical to subtropical) suggest the influence of a

warm current (Head, 1997: de Vernal and Marret, 2007).

Moreover, the occurrence of a freshwater alga Pediastrum

sp. suggests proximity to a river that fed the embayment.

This inference was also supported by the sedimentary

facies analysis (Igarashi and Kurita, 2007). The shallow-

marine molluscan fauna herein described is different from

the tropical to subtropical Arcid-Potamid fauna (Tsuda,

1965), despite them sharing a similar paleoenvironment.

Recently Yanagisawa and Watanabe (2017) postulated that the

occurrence of the Arcid-Potamid fauna in Japan was confined

to the interval of 17.0 to 16.7 Ma. As the Budo Member was

deposited slightly later (16.6 to 15.9 Ma) than the Arcid-

Potamid fauna, it does not include any characteristic species of

the fauna. However, most species of Type 2 and the Arca noae

group to which Arca (Arca) budoensis new species belongs

live in warm water which is concordant with the paleo-

environment inferred by the dinoflagellate cysts.

As described here, Arca (Arca) budoensis new

species resembles both fossil and modern species from

the eastern Pacific, not from the western Pacific. In the

early Miocene, six bivalve genera and one subgenus have

been recognized as immigrants from the eastern Pacific to

the west during the early to early middle Miocene. The

taxa include the venerids Securella, Kaneharaia, Comp-

somyax, the hiattelid Panomya, the rock-boring myid

Platyodon, the Pholadid Penitella, and the tellinid sub-

genus Rexithaerus (Amano, 2005). One modern species of

the Type 2 (Lutaenko and Maestrati, 2007) in the western

Pacific, Arca (Arca) navicularis first appeared only in the

Pliocene of Java, Indonesia (Kase et al., 2008). Thus, Arca

(Arca) budoensis new species was derived from the

ancestor which migrated from the eastern Pacific to Japan

at least by the early Miocene.

ACKNOWLEDGMENTS

We are grateful to Geerat J. Vermeij (UC Davis) for his

critical reading of the manuscript and useful suggestions.

THE NAUTILUS, Vol. 134, Nos. 3-4

We thank Sven N. Nielsen (Universidad Austral de Chile)

and an anonymous reviewer for their useful comments.

We also thank Kazuo Kawauchi (Niigata University of

Pharmacy and Applied Life Sciences) and Yukihiko Kamoi

(Niigata City) for kindly helping one of the authors, KA to

collect the molluscan species.

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THE NAUTILUS 134(3-4):138-142, 2020

RESEARCH NOTE

Page 138

Imposex in the city: First evidence of

female masculinization in Queen Conch

Aliger gigas (Linnaeus, 1758)

(Gastropoda: Strombidae) in Florida

Justin N. Voss

Gabriel A. Delgado

Fish and Wildlife Research Institute

Florida Fish and Wildlife Conservation Commission

Marathon, Florida 33050, USA

Nancy J. Brown-Peterson

Center for Fisheries Research and Development

School of Ocean Science and Technology

University of Southern Mississippi

Ocean Springs, Mississippi 39566, USA

Einat Sandbank

Robert A. Glazer

Fish and Wildlife Research Institute

Florida Fish and Wildlife Conservation Commission

Marathon, Florida 33050, USA

Organotins, such as tributyltin (TBT), are chemical

compounds added to antifouling paint that were used

worldwide on oceangoing vessels from the 1960s until 2008,

when the International Maritime Organization banned

them globally through the International Convention on

the Control of Harmful Antifouling Systems on Ships

(Horiguchi, 2017; Laranjeiro et al., 2018). The use of

organotins adversely impacted growth, development, and

reproduction in numerous marine organisms (Horiguchi,

2017; Vogt et al., 2018). Despite the global ban, organotins

can persist in the environment for decades (Filipkowska

et al., 2014; Horiguchi, 2017), thus still posing a threat to

susceptible species. In gastropods, organotin exposure can

lead to the development of a condition known as imposex

(Bryan et al., 1988; Horiguchi, 2017). Imposex is the

superimposition of male reproductive tissues (ie., penis

and vas deferens) onto female gastropods and is caused

by disruption of the neuroendocrine system (Oberdorster

et al., 2005; Horiguchi, 2017). Continued exposure to

organotins increases masculinization in females and can

reduce fecundity (Averbuj and Penchaszadeh, 2010). In

severe cases, the vaginal opening is blocked, resulting

in sterility (Bryan et al., 1988; Horiguchi et al., 1995),

ultimately leading to large population declines (Cadee

et al., 1995; Horiguchi et al., 2006). Imposex in the natural

environment is almost exclusively associated with orga-

notin exposure (Oehlmann et al., 2007; Sternberg et al.,

2010; Horiguchi, 2017), to the point that imposex in

gastropods is used as a biomarker for TBT contamination

(Oehlmann et al., 2007; Horiguchi, 2017).

The incidence of gastropod imposex is higher near

shipping channels, industrial seaports, and marinas (Li

and Collin 2009; Averbuj and Penchaszadeh, 2010;

Laranjeiro et al., 2018). The Queen Conch, Aliger gigas

(Linnaeus, 1758), is a large, gonochoristic marine gas-

tropod that has been protected in Florida since 1986

because of overfishing. A well-documented breeding

ageregation of Queen Conch (Berry et al., 2016) sits

adjacent to Port Everglades (Figure 1), one of the busiest

industrial seaports in the United States (http:/Awww.

porteverglades.net). And while TBT-induced imposex

in areas of elevated boating activity occur in the Caribbean

(Titley-O’Neal et al., 2011), imposex has never been re-

ported in Queen Conch in Florida.

We conducted a demographic survey of the Port Ev-

erglades Queen Conch aggregation on August 30, 2007,

and another on June 27, 2018. A 100-m tape was deployed

along one margin of the aggregation; five secondary tapes

were then placed perpendicular to the primary tape at

random intervals. A secondary tape extended either 100 m

into the aggregation or, if positioned at a point at which

the aggregation was less than 100 m wide, to its distant

margin. Scuba divers counted all adult Queen Conch

(defined as having a fully flared lip approximately 10 mm

thick or more) within 1 m either side of the secondary

tapes to estimate the density of the aggregation. Identi-

fying the sex of a conch in situ requires turning it over and

patiently waiting for its body to emerge. Because of the

time constraints of scuba surveys, individuals were not

sexed, but reproductive behaviors (i.e., mating and egg

laying) were noted.

Our two aggregation surveys yielded comparable re-

sults in terms of density and the percentage of Queen

Conch engaged in reproductive behaviors. In 2007, we

surveyed 1000 m* and counted 73 adult Queen Conch, for

a density of 730 ha +. Of these, 5.5% were mating and

9.6% were laying egg masses. During the 2018 survey, we

surveyed 850 m~ and counted 42 adult Queen Conch, for

a density of 494 ha *. Of these, 4.8% were mating and

11.9% were laying egg masses.

During the 2018 survey, nine female conchs (so

identified because they were mating or egg laying) were

collected for assessment of ovarian condition. After re-

moving the bodies from their shells, we observed and

recorded that some of these females exhibited imposex

(Figure 2). We moved on to collect a sample of ovary

tissue from each female, excising a piece of tissue ap-

proximately 1 cm® from the middle of each ovary. The

samples were placed in labeled plastic cassettes and fixed

for seven days in 10% neutral buffered formalin. Fol-

lowing fixation, we prepared the ovarian tissues for

histological assessment of ovarian maturity and the per-

centage of developed ovarian tissue using the methods of

Delgado et al. (2004). For each female, we also recorded

J.N. Voss et al., 2020

“South Florida

Figure 1. Map of South Florida indicating the location of Port

Everglades. Inset shows the site of the Queen Conch aggre-

gation found at the entrance to Port Everglades.

four morphological measurements, as described by

Delgado et al. (2019): shell length, lip thickness, shell

weight, and body weight. Unpaired t-tests, performed in

Sigma Plot 13 (Systat Software Inc., San Jose, CA, USA),

were used to test for differences in histology and mor-

phology between females exhibiting imposex and normal

females. Results were considered statistically significant if

Pi 0205,

For each female, we ranked the severity of imposex

using the scale developed by Cob et al. (2011), which

ranges from 0 (i.e., normal; no imposex) to 3 (i.e., having a

complete penis). Imposex was observed in four of the nine

females (44.4%). One female was characterized as having

stage-1 imposex (Figure 2). The other three females were

characterized as having stage-2 imposex (Figure 2). From

histology, all four females with imposex were classified as

ripe and spawning-capable (i.e., they had tertiary vitel-

logenic oocytes, with oocytes in the oviduct; Figure 3).

Imposex-affected and normal females did not differ sig-

nificantly in shell length (¢ = —1.514; df = 7; P = 0.174),

Page 139

lip thickness (¢ = 0.037; df = 7; P = 0.972), shell weight

(t = 1.275; df = 7; P = 0.243), body weight (t = —1.976;

df = 7; P = 0.089), or the percentage of developed ovarian

tissue. (f= 1 O3ldt ="7.-P = 0.225) able 1);

This is the first documented case of imposex in Queen

Conch in Florida. While we did not test for the presence

of organotins at the aggregation site, the proximity to a

major shipping port suggests that organotin exposure was

the likely cause of the malady (Oehlmann et al., 2007;

Sternberg et al., 2010; Horiguchi, 2017). The marine

habitats surrounding Port Everglades are almost certainly

contaminated with organotins that predate the antifoulant

ban in 2008 as evident by the fact that the nearby disposal

site for dredged material from Port Everglades is heavily

contaminated by organotins (U.S. EPA and USACE

2020). Although organotin is banned from being sold in 67

countries, it is still manufactured and sold in many Ca-

ribbean and South American countries (Turner and

Glegg, 2014). Ships originating from these countries may

be a more recent source of contamination.

There may be other endocrine disruptors that might be

causing imposex at Port Everglades, but the over-

whelming evidence from the literature suggests organotin

exposure (Oehlmann et al., 2007; Sternberg et al., 2010;

Horiguchi, 2017). That said, the extent of organotin

contamination around South Florida is unknown. Future

research should include water, sediment, and tissue assays

to determine the extent of organotin contamination

around South Florida and could also include a local

member of the widespread Stramonita haemastoma

(Linnaeus, 1767) species complex such as Stramonita

floridana (Conrad, 1837) to serve as a bioindicator (sensu

Stickle and Zhang, 2003; Limaverde et al., 2007)

In contrast to studies finding that imposex negatively

affects reproduction in gastropods (Horiguchi et al., 2006;

Lahbib et al., 2009; Averbuj and Penchaszadeh, 2010),

Figures 2. Two imposex-affected female Queen Conch from Port Everglades, Florida (removed from shells).

Page 140

THE NAUTILUS, Vol. 134, Nos. 3-4

Figure 3. Histological section of ovarian tissue from an imposex-affected Queen Conch from Port Everglades, Florida. This female

was classified as ripe and spawning-capable. Note the abundant tertiary vitellogenic oocytes (Vtg3) throughout the ovarian tissue; minor

atresia (i.e., resorption of some vitellogenic oocytes) can also be seen.

the >40% incidence rate of imposex in our samples did

not seem to affect reproductive activities. The percentage

of Queen Conch mating and laying eggs in the Port

Everglades aggregation was similar to that reported

elsewhere in Florida (Delgado and Glazer, 2020), where

no imposex has been reported (Delgado et al., 2004,

2019). Based on histology, the ovaries of the females

afflicted with imposex were indistinguishable from those

of normal, ripe females. This may be because, while the

incidence rate of imposex was relatively high, the severity

of the affliction was not in the advanced stages. The

Table 1. Morphological and histological measurements

(mean + | standard error) of female Queen Conch from Port

Everglades, Florida.

Imposex-affected Normal

Shell length (cm) DOB. a O62 23.6: 0.58

Lip thickness (mm) 28.3. 2 D301 280 ae, 1,32

Shell weight (g) a GY) re emus 5) 9 238 f IAG

Body weight (g) 355 = 29:6 ATA ee AGS

Ovarian tissue (%) 97.0 + 1.92 09.8: 2 35

females we examined were engaged in reproductive ac-

tivities, which may have biased our results; there may be

females within the aggregation with more severe cases of

imposex that are not reproductively active. Future re-

search might explore the relationship between age and

imposex severity and incidence rate, because older fe-

males will have been exposed longer.

Studies of imposex in other strombids have reported

morphological differences between imposex-affected and

normal females. In Strombus pugilis Linnaeus, 1758 and

Conomurex luhuanus (Linnaeus, 1758), imposex-affected

females were larger and heavier than normal females

(Reed, 1993, 1995). In Laevistrombus canarium (Lin-

naeus, 1758), imposex-affected females had larger and

heavier shells but lighter bodies than did normal females

(Cob et al., 2008). However, we did not observe statis-

tically significant differences in gross morphology be-

tween imposex-affected and normal females. This may

have been because of the comparatively mild severity of

imposex in the females we examined.

Imposex can cause population declines in gastropods

(Cadee et al., 1995; Horiguchi et al., 2006), but this was

not observed in Queen Conch at Port Everglades. The

J.N. Voss et al., 2020

demographics of the aggregation remained steady over

our two surveys and were consistent with those reported

by Berry et al. (2016). Berry et al. (2016), however, noted

abnormal egg masses at the Port Everglades aggregation.

Normal egg masses are crescent shaped and covered with

sand; the abnormal ones were stringy and not covered

with sand (Berry et al., 2016: fig 7). We also observed

females laying abnormal egg masses. In other gastropods,

egg laying becomes disrupted in advanced stages of

imposex (Horiguchi, 2017), which may explain why some

Queen Conch females at Port Everglades lay abnormal

egg masses, but further study is needed.

It is unknown whether the offspring of imposex-

affected females are as viable as those from normal

Queen Conch. Other pollutants can cause transgenera-

tional effects in mollusks that reduce larval growth and

survival (Nice et al., 2003) and juvenile fitness (Osborne

et al., 2020). If the veligers and juveniles from imposex-

affected Queen Conch are compromised, this may have

negative consequences for larval connectivity to down-

stream populations and metapopulation persistence.

Beyond the effects on Queen Conch, the results from

this study highlight the deleterious impacts that society may

have on wildlife resources that are juxtaposed closely to

urban centers. Organotins represent only one in a much

wider suite of chemicals that enter the South Florida

nearshore marine environment from sources as diverse as

agriculture (e.g., fertilizers, herbicides, pesticides), indus-

try, recreation (e.g., golf course fertilization and pesticide

application), and residential activities (e.g., sewage). How

society values these resources and prioritizes the mitigation

of the pernicious threats associated with point and non-

point sources of pollution will ultimately determine the

overall health of nearshore ecosystems and the sustain-

ability of the resources they support.

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Meu ft bie Ss

Volume 134

2020

AUTHOR INDEX

PWBINOS Res O oak 2) essa hl, Nie cheat ct a a rch aS li de L Dll dl 32, J OVE Sia ta SMe Pew ie tte ne ae Ta Oe RI na A 36

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BERPSCH, TAY 2 cy Piece See ae ee eee 71 PEL Hey ee NPM LS OR eMC Sirs See AOD aE RL eo Soe ae Swe

PTET ERS EN, aie drcate acral MOREOCT dara nae ee 61 (Ere Stadha oe HEN ON et net eee ew, ee Oe eee a eee 107

BROWN-PETERSON, NANCY J. sseceesescesstestesees te teeeeecs nee teeseeees 138 SUSIE VAGNIR, We et Sins hoe tt tang eae, Be Mee aaa ote la RS Ae face 89

COSTA, P.MAS. seis neeesneeseneecsnecennncesscteneteeesccennnaten 95 eISIGIE: Weiter | Mebane any Sere PG Nae Seco nethe Hater 1 eer 61

CUCEINE TRO) G2 ltin 2.5 0a dee a cine eit eae 92 SEHR TUE ria cs fe tar aN ge CES GEE wy ah a 89

DELGADO, GoAn voices eeeieessneeeenetenieennetcneicen 89, 138 SSA HISD oe WAI cg hs SM AG of Lien ie i elie et oPh ali oie Mele fly i!

TASH ON. PPE s en sencas cee Binh te nace tere Nestle te Aine, eetechet ee atest 61

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FRCS Ae AE on cue nels ees re de SE ene Ue Ore ade 117 HSER IER) Led ea RON ONE erie nein We Rotel, MECN A we RE ae 36

KuRITA, | RAB AIL By de at ae Earl vr gene 8 Del ne a oah 132 Voss, J.N. MESSER HORT ah ee AD Rn aint soot Ravicgh-tioie RATE ITE! tairtibn Weel nat beak 138

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NITMICE BROOKS Nia cckcbnuee une rans okt ei cee eee 92 PASC ice cdhs | (PRC ee ABET A) Reha GREE TEA Chaar iE mn ARMA Tad rn, 45

NEW TAXA PROPOSED IN VOLUME 134

GASTROPODA

Bathyacmaea austrina S.-Q. Zhang and-S.-P. Zang, 2020, new species (Peetimodontidae) cc...) \lcclesct-sctesteconesseteeccbetseceoteecterens 46

Bathyacmaea brevidentata S.-Q. Zhang and S.-P. Zhang, 2020, new species (Pectinodontidae) ..........cccccseeseeeteteesereneeeeneneeneneees 48

Paciocinebrina bormannae Wiedrick and Houart, 2020, mew species (Miuricidae 20. .521c...10.0.tcke be shea velo eves eee nateth oreo en sete 118

Paciocinebrina grandilurida Wiedrick and Houart,2020) new species (Mbumeig@ ae) \..6.2252.5 io sisctend erie kya xe ete attest 120

Paciocinebrina mininterfossa Wiedrick and Houart,' 2020 new species ( Mumicidae) sqjscaciczsvested one iegseu jopsethen dsesteay- ca tae akin eal

Paciocinebrina murphyorum Wiedrick and Mota. 2020 mew species (I Viineldlae ) 05.0.0 sae dad caade nie ds, 0 es eaten 124

Paciocinebrina pseudopusilla Wiedrick and: Mouart, 2020, new species (Miiricicdar) <..h.sc.in<-cnsncsoessisotasysesesnaveras gehnvaueavenensatt tetaeetee 124

Paciocinebrina pusilla Widdrielk-and Hewert, 2080,.mew ameciess(M mic aeis 6h. sen dade livocssusotuneore- tierra yan bona 126

Rolleia simonaikem Watters, Smith ,and-Snedden, 2020 new species AMA NiGae) occ. ccccetacycnsactac causa esos caso eGbavosecpadh rena toa tnaasee 30

Tritonoharpa curvapex Souza, Gomes, and ‘Cosia,,2020, new species {Camoellamidae) o.-<.c5..ce..20+<4400 vss nennyouvasEeonsbason oosnnsesiesaseynstbonsna 96

BIVALVIA

Arca (Arca) -budoensis Amano, 2020, mewesbecies.( Avoidance. fagsil)-< cnr rene ten ee hepa mete sree tea teeta ne 124

Cuspidaria inowei Amano and Kurita, 2020; new species (Cuspidatiaaes tonsil 4 it in.dic,.ctr eaten sah atts meen eae 53

REVIEWERS FOR VOLUME 134

Beck, Lothar

Behrens, David

Bieler, Riidiger

Bogan, Arthur E.

Breure, Abraham

Coan, Eugene V.

Davis, Megan

Fallon, Philip

Flint, Mark

Frost, Thomas P.

Garrigues, Bernard

Groves, Lindsey

Harasewych, M.G.

Jenkins, Robert G.

Kantor, Yuri

Kiel, Steffen

Lee, Harry G.

McGann, Mary L.

Nielsen, Sven

Oberdoster, Eva

Oliver, Graham

Orstan, Aydin

Lutaenko, Konstantin A.

Pearce, Timothy A.

Portell, Roger W.

Rosenberg, Gary

Ter Poorten, Jan Johan

Slapcinsky, John

Stanley, Richard G.

Stout, Carla

Valentich-Scott, Paul

Verhecken, André

Walker, Brian

Zelaya, Diego

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