THE NAUTILUS

Volume 131, Number 2

June 13, 2017

ISSN 0028-1344

A quarterly devoted

to malacology.

&. 1—

4o(

iAj MZ-

EDITOR-IN-CHIEF

Jose H. Leal

The Bailey-Matthews National

Shell Museum

3075 Sanibel-Captiva Road

Sanibel, FL 33957 USA

EDITOR EMERITUS

M. G. Harasewyeh

Department of Invertebrate Zoology

National Museum of

Natural History

Smithsonian Institution

Washington, DC 20560 USA

CONSULTING EDITORS

Riidiger Bieler

Department of Invertebrates

Field Museum of

Natural History

Chicago, IL 60605 USA

Arthur E. Bogan

North Carolina State Museum of

Natural Sciences

Raleigh, NC 27626 USA

Philippe Bouchet

Laboratoire de Biologie des

Invertebres Marins et Malacologie

Museum National d’Histoire Naturelle

55, rue Buffon

Paris, 75005 FRANCE

Robert H. Cowie

Center for Conservation Research

and Training

University of Hawaii

3050 Made Way, Gilmore 409

Honolulu, HI 96822 USA

Kenneth A. Hayes

Department of Biology

Howard University

Washington, DC 20001 USA

Steffen Kiel

Department of Paleobiology

Swedish Museum of Natural History

Box 50007

104 05 Stockholm, SWEDEN

Harry G. Lee

4132 Ortega Forest Drive

Jacksonville, FL 32210 USA

Charles Lydeard

Biodiversity and Systematics

Department of Biological Sciences

University of Alabama

Tuscaloosa, AL 35487 USA

Bruce A. Marshall

Museum of New Zealand

Te Papa Tongarewa

PO. Box 467

Wellington, NEW ZEALAND

Paula M. Mikkelsen

Paleontological Research

Institution

1259 Tmmansburg Road

Ithaca, NY 14850 USA

Diarmaid O Foighil

Museum of Zoology and Department

of Biology

University of Michigan

Ann Arbor, MI 48109-1079 USA

Gustav Pau lay

Florida Museum of Natural History-

University of Florida

Gainesville, FL 32611-2035 USA

Gary Rosenberg

Department of Mollusks

The Academy of Natural Sciences

1900 Benjamin Franklin Parkway

Philadelphia, PA 19103 USA

Elizabeth Shea

Mollusk Department

Delaware Museum of

Natural History

Wilmington, DE 19807 USA

Angel Valdes

Department of Malacology

Natural History Museum

of Los Angeles County

900 Exposition Boulevard

Los Angeles, CA 90007 USA

Geerat J. Vermeij

Department of Geology

University of California at Davis

Davis, CA 95616 USA

G. Thomas Watters

Aquatic Ecology Laboratory

1314 Kinnear Road

Columbus, OH 43212-1194 USA

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THE0NAUTILUS

Volume 131, Number 2

June 13, 2017

ISSN 0028-1344

CONTENTS

Edna Naranjo-Garcia First inventory of the introduced and invasive mollusks in Mexico . 107

Zoila G. Castillo-Rodriguez

John D. Taylor A new species of Lucinoma from 240-500 m on the continental shelf break

Emily A. Glover off Newfoundland (Bivalvia: Lucinidae) . 127

Patrick Anseeuw Bayerotrochus belauensis, a new species of pleurotomariid from the Palau

Lori J. Bell Islands, western Pacific (Gastropoda: Pleurotomariidae) . 138

M.G. Harasewych

Leonard G. Brown A redescription of Rissoina tnayori Dali, 1927, a junior subjective synonym

Bruce D. Neville of Opalia pumilio (Moreh, 1875) (Gastropoda: Epitoniidae) . 147

Erratum

150

THE NAUTILUS 1 31 (2): 1 07-126, 2017

Page 107

First inventory of the introduced and invasive mollusks in Mexico

Edna Naranjo-Garcfa

Instituto de Biologfa

Universidad Nacional Autonoma de Mexico

Apartado Postal 70-153

Mexico City, C.P. 04510, MEXICO

[email protected]

Zoila G. Castillo-Rodriguez

Instituto de Ciencias del Mar y Limnologia

Depto. de Biodiversidad y Ecologfa Acuatica

Universidad Nacional Autonoma de Mexico

Apartado Postal 70-153

Mexico City, C.P. 04510, MEXICO

zgcr@cm arl . u nam . mx

ABSTRACT

Early historical records are included in this first national in¬

ventory of species of mollusks introduced, whether intentionally

or not, into Mexico hv humans. Of the 56 exotic-invasive mol¬

lusks listed, 15 inhabit brackish and marine environments, 10

freshwater, and 31 are terrestrials. Thirty-six per cent of the

introduced species come from Europe and the Mediterranean,

18% from Asia- Australia-New Zealand, 46 % are cryptogenic,

coming from different regions of the planet, and the origin of

several others is uncertain. The best-represented families are

Mvtilidae, Teredinidae (brackish and marine), Ampullariidae,

Thiaridae, and Planorbidae (freshwater), and Helieidae, Agrioli-

maeidae, Limacidae, Subulinidae, and Vallonidae (terrestrial).

They involve Mytilus galloprovincialis Lamarck, 1819, Pomacea

canaliculata, Lamarck, 1819, and Dreissena polynwrpha (Pallas,

1771), species included among the world's worst invasive species.

Some have become naturalized: three brackish and marine

species, six freshwater, and twelve terrestrial. The increase in

exchange of goods, services, and transport litis assisted in the

transfer of species from distant places, as has intentional or

unintentional introduction of species of economic importance

(M. galloprovincialis, Crassostrea gigas (Thunberg, 1793),

C. sikanxea (Amemiya, 1928)), without consideration of the

epibionts, endobionts, and endoparasites that may also be

introduced. Effective strategies must be developed to minimize

the potential impact of biological invasions and raise public

awareness of the problem; this must include the rigorous ap¬

plication of more stringent regulations.

Additional Keywords: exotic, freshwater, terrestrial mollusks,

marine, mollusks, economic and biological risks

INTRODUCTION

Biological contamination in the world has increased

markedly since 2000 (Crocetta et ah, 2013), Pimentel

et al. (2001) estimated that some 480000 species have

been introduced around the world throughout the his¬

tory of humankind, and this is of great concern. The

effects that alien species may have in ecosystems

(Carlton, 1999) and in their interaction with native

organisms are poorly understood; however, we face loss

of diversity (Reyna et ah, 2013) of formerly diverse

ecosystems (Cowie, 1998, 2001; Cowie and Robinson,

2003; Lopez-Lopez et ah, 2009). It may be possible that

endemic species are the most vulnerable, although un¬

fortunately, knowledge of the diversity and abundance of

much of the world’s fauna remains unsatisfactory. In

M exieo, where it is estimated that 75 % of brackish and

marine species are known, it is suggested that 1 7 % of the

Pacific species, and 15% of those in the Gulf of Mexico

and the Caribbean Mexican coast are endemic (Castillo-

Rodriguez, 2014). On the other hand, fewer than 35 %

of the native non-marine mollusks are known and 85%

of the Mexican territory is in need of exploration

(Thompson, 2011).

In the case of the introduction pathways at global

level, marine vectors are well documented. Mexico has

a navigation infrastructure that facilitates the in¬

troduction of alien species; human activities on the

continental margins, and in the bays and estuaries of the

coastal zone, have evolved since the 16th century and are

now part of an impressive network of global marine

traffic. Currently there is the threat of climate change,

which will undoubtedly alter the structure and compo¬

sition of native communities. This will also alter the

functioning of ecosystems and become a stressor that will

further increase the risk of biological invasions in marine

and non-marine systems. The effects of climate change

on the environment will include substantial impact on

native species. Given the increase in threats to the native

fauna, inventories of exotic species become the foun¬

dation for future actions, including the control and

eradication of invasives (Mendoza et ah, 2014). This

article offers a review of the brackish-marine and non¬

marine introduced mollusks in Mexico, constituting the

first national inventory of this type elaborated in Mexico;

it also determines the naturalized species at the national

level. We examine potential vectors and recommend

measures that may help prevent the entry of additional

alien organisms and that could help control and serve as

essential protective measures for the fauna and the

environment at the national level in Mexico.

Page 108

THE NAUTILUS, Vol. 131, No. 2

Early Records of Introduced Mollusks in Mexico

In relation to introduced non-marine mollusks. Cornu

aspersum (Muller, 1774) (as Helix aspersa Muller, 1774)

was detected in Mexico by Alexander Humboldt between

1803 and 1804 (Martens 1890-1901); and it was recorded

by Pilsbry (1891) in Mexico City. It was also located in

Jaral, Guanajuato, and other unspecified sites (Martens

1890-1901). The species became a pest in gardens

throughout Mexico City (Ancona, 1947) and was later

recorded at Chapultepec, and in mountains between

Mexico City and the city of Cuernavaca (Jacobson, 1952).

Baker (1925) mentioned the presence of Phyllocaulis gatji

(Fischer, 1871) in Mazatlan, Sinaloa. Andrews and

Dundee (1987) stated for the first time the problems

caused by the slug Sarasinula plebeia (Fischer, 1868) in

Chiapas (1980) and Veracruz (1981), and Naranjo-Garcfa

et al. (2007) reviewed the distribution of the family

Veronicellidae nationwide, with particular reference

to Sarasinula plebeia (= Sarasinula dubia (Semper,

1885)). As for other introduced slugs, Cockerell (1923)

recorded Limacus flavus (Linnaeus, 1758) (as Limax

flavus Linnaeus, 1758) with numerous individuals or

populations in Mexico City. After that. Baker (1930)

Found various European slugs ( Limax nuiximus Linnaeus,

1758 in Desierto de Los Leones to Cuajimalpa; L. flavus in

Huachinango, Puebla; Deroceras laeve (Muller, 1774)

(apballic) in Desierto de Los Leones and Neeaxa; D. laeve

(phallic) (Muller, 1774) in Cuajimalpa and San Juan

Teotihuacan; and Milax gagates (Drapamaud, 1801) in

Desierto de Los Leones. The Cuban species Zachnjsia

auricomya havanensis Pilsbry, 1894 was recorded in

Yucatan (Bequaert and Clench, 1936) and Vallonia

excentrica Sterld, 1893 was found by Joshua L. Baily Jr. in

Cuernavaca (Pilsbry, 1948).

The freshwater clam Corbictda fluminea ( Nl idler, 1774)

(as Corbictda nianilensis (Philippi, 1844)) was recorded in

Baja California, northwestern Mexico (Fox, 1970) and

Hillis and Mayden (1985) summarized its distribution

along the coastal areas of Pacific and in the State of

Tamaulipas; it was later recorded in Lake Catemaco,

southern Veracruz (Torres-Orozco and Revueltas-Valle,

1996). The freshwater snail Melanoides tuberculata

(Muller, 1774) was found in the vicinity of Veracruz in

1973 (Abliott, 1973), and its presence in Mexico was

confirmed in 1975 by Pointier and McCullough (1989).

More details of its distribution in Mexico have subsequently

emerged (Contreras-Arquieta, 1998; Contreras- Arquieta

and Contreras-Balderas, 2000; Contreras-Arquieta, et al.

1995). The first record of Tarebia granifera (Lamarck,

1822) was at Lake Catemaco, Veracruz (Naranjo-Garcfa

et al., 2005), and other foci were later found in northern

Veracruz State (Dipez- Lopez et al., 2009), in southern

Oaxaca State, and in the Lacandona Forest, Chiapas

(Naranjo-Garcfa non-published data); it was later recorded

from 1 1 lakes of three municipalities in the State of Tabasco

(Rangel-Ruiz et al., 2011).

The terrestrial species Rumina decollate (Linnaeus, 1758)

was recorded in 1993 in the States of San Luis Potosf and

Tamaulipas (Correa-Sandoval, 1993, 1998; Correa-Sandoval

and Rodriguez, 2002), and soon afterward in Nuevo Leon

(Correa-Sandoval 1999b; Correa-Sandoval and Rodriguez,

2005; Correa-Sandoval et al., 2007). Huttonella bicolor

(Hutton, 1834) (as Gulella bicolor (Hutton, 1834)) was

collected at the archaeological site El Tajin, Veracruz

(Correa-Sandoval, 1999a, 2000).

In relation to marine mollusks introduced into Mexican

waters, Hendrickx (1980) and Salgado-Barragan and

Toledano (2006) provided data from specimens ob¬

served in situ and preserved in the Invertebrate Collec¬

tion (EMU) of Unidad Aeademica of Mazatlan, Instituto

de Ciencias del Mar y Limnologfa, Universidad Nacional

Autonoma de Mexico. There are also other published

records: for example, Okolodkov et al. (2007) and Ortiz-

Arellano and Salgado-Barragan (2012). According to

Carlton (1999), the diversity and abundance of exotic and

invasive marine species at worldwide level are still poorly

known; this is certainly true for Mexico, where lack

of awareness of the presence of exotic species is

compounded bv unfamiliarity of their effects on the

ecosystems.

MATERIALS AND METHODS

The present compilation was assembled based on records

of mollusks deposited in: Coleccion Nacional de Moluscos

(National Collection of Mollusks), Instituto de Biologfa,

Universidad Nacional Autonoma de Mexico); database of

the Invertebrate Zoology division of the Florida Museum

of Natural History, FLMNH; the Unidad Aeademica of

Mazatlan of the Instituto de Ciencias del Mar y

Limnologfa (ICMyL-MAZ, UNAM); the literature; field

observations by the authors and/or other specialists; a list

of organisms coming from Mexico and intercepted at

customs upon arrival in the USA; and additional in¬

formation provided by David Robinson (letter to E.

Naranjo-Garcfa, 4 September 2014). In certain cases,

where the literature is very extensive, the first and the

most recent articles or references were selected. Acro¬

nyms used are: CNMO, Coleccion Nacional de Moluscos,

Universidad Nacional Autonoma de Mexico; CM, Car¬

negie Museum of Natural History, Pittsburgh; FLMNH,

database of the Florida Museum of Natural History,

Gainesville; EMU, Coleccion de Invertebrados, Unidad

Aeademica de Mazatlan del Instituto de Ciencias del Mar

y Limnologfa.

RESULTS

In total, 56 species of exotic-invasive mollusks from the

classes Bivalvia and Gastropoda have been so far in¬

troduced into Mexico, whether intentionally or not: 15

brackish-marine, 10 freshwater, and 31 terrestrial mol¬

lusks (Tables 1, 2, and 3). Of these, 36 % came from

Europe and the Mediterranean (the majority are terres¬

trial mollusks), 18 % from Asia- Australia-New Caledonia

E. Naranjo-Garcfa and Z.G. Castillo-Rodriguez, 2017

Page 109

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Page 110

THE NAUTILUS, Vol. 131, No. 2

(several estuarine/brackish- marine and some freshwater

species), 46 % from different regions of the planet

(Table 4), with the place of origin of several being

uncertain.

Most of the brackish and marine species (Table 1)

belong to the families Mytilidae and Teredinidae

(Bivalvia). Among the freshwater mollusks (Table 2),

gastropods are the dominant group, in particularly

species in the families Ampullariidae, Thiaridae, and

Planorbidae. Most of the terrestrial mollusks (Gastro¬

poda) (Table 3) belong to the family Helieidae, followed

by the Agriolimacidae, the Limacidae, the Subulinidae,

and the Vallonidae. The species that have been

recorded only once are: Theora lubrica Gould, 1861

(marine), Helisoma duriji (Wetherby, 1879) (as Pla-

norbella duriji (Wetherby, 1879)) (freshwater), and

Phyllocaulis gayi, Cecilioides acicula ( M tiller, 1774),

Arion circumscriptum Johnston, 1828, Vallonia costata

(Miiller, 1774), Zachrysia auricomya havanensis, and

Cantareus apertus (Born, 1778) (as Helix aperta Born,

1778) (terrestrial).

DISCUSSION

Marine Mollusks

The ecosystems that now are home to exotic mollusks are

vulnerable to changes in their composition, regardless of

whether those mollusks were introduced inadvertently or

for commercial purposes. Especially threatening are those

species that have been restricted to environments dis¬

turbed either naturally or by human intervention, as is the

case of brackish and marine mollusks such as Mytilus

gallop rovindolis Lamarck, 1819 (Mytilidae). Mytilus

galloprovincialis, originally from the Mediterranean,

Black Sea and Adriatic Sea, has been categorized as one of

the 100 worst invasive alien species of the world. Despite

this, the species is cultivated for food in the states of Baja

California Norte and Sur, and this is without any

knowledge of its ecological impact on the native species.

Populations of the mussel Pema perna (Linnaeus, 1758)

are distributed from Texas to southern Veracruz State,

Mexico (Hicks and Tunnell, 1993, 1995; McGrath et al.,

1 998), and Hicks et al. (2001) consider that its occurrence

on Mexican coasts should be carefully monitored.

Mussels and shipworms can survive tough conditions in

variable estuarine and marine environments, as well as in

sheltered sites that may be favorable to their development

and dispersal: they are adapted to this survival by their

sessile habit, their filter feeding, and their modes of

reproduction, growth, and morphological protection

((McDonald and Koehn, 1988; Turner, 1966; Tuente

et al., 2002; Petes et al., 2007; Didziulis, 2007).

Tl re shipworms, such as Teredo navalis Linnaeus, 1758,

bore into submerged wood substrata all over the world,

and there are few records of the species in Mexico. Lopez-

Garrido (2008) recorded the species from sunken boats in

the state of Campeche, in the southern Gulf of Mexico.

There are also references from the early 1900s (Dublan

and Lozano, 1901; Mariscal, 1902) regarding re¬

quirements of the asphalt composition in Submarine

Telegraph cables between the port of Veracruz and

Campeche in order to avoid damage caused by the

“marine worm”, T. navalis. More recently, there are

checklists, reports and theses records of its presence in

estuaries on the Mexican coast of the Gulf of Mexico, such

as the Tampamachoco Lagoon, Veracruz (without ref¬

erence code), where it has been an element of the epi-

biosis on the mangrove Rhizophora mangle Linnaeus,

1753 since 1980.

Oysters of Asian origin, Crassostrea gigas (Thunberg,

1793) and, more recently, C. sikamea (Amemiya, 1928),

are cultivated in Baja California Norte and Sur in Mexico.

Sessile oysters adhering by cementing to any hard sub¬

stratum, together with the large accompanying fauna on

the surface of their shells, represent a potential risk to

wildlife; hence, a study of their ecological impact would be

most important. Crassostrea gigas exists as an exotic

species on the southern Pacific coast and, since un¬

controlled introduction is possible through ballast water

and aquaculture practices, it might also be expected to

eventually be introduced along the southern Gulf of

Mexico, in locations as Veracruz State.

Anadara transversa (Say, 1822), a clam from the

northwestern Atlantic, is considered a non-invasive exotic

species. It was recorded in Tamiahua lagoon, Veracruz by

Garcia-Cubas (1969) and Abbott (1974). Although there

are no recent records of substantial living populations,

abundant disjointed valves have been reported.

Theora lubrica Gould, 186L originally from Asia, is

considered as one of the most important invasive species

in Europe (Balena et ill., 2002). According to Steneck and

Carlton (2001), it is one of the 15000 species that have

been transported across the world in ballast water. It is

recorded from Baja California Norte, where it possibly

arrived secondarily introduced from San Francisco Bay,

U SA; it can be considered a potentially invasive species for

the Pacific States of Mexico.

There are fewer records of exotic marine gastropods

species in Mexico than of exotic bivalve species. Among

these gastropods is the pyramidellid Boonea bisuturalis

(Say, 1822), a native of the northern coast of the Atlantic

(Canada and USA) that feeds on the body fluids of in¬

vertebrates (Fretter and Graham, 1949; 1962; Fretter,

1951), including polychaetes, gastropods, and bivalves,

and minor groups such as polyplacophorans and some

eehinoderms (Robertson and Orr, 1961). The Mexican

coast of the Gulf of Mexico houses marine resources that

include the Eastern Oyster C. virginica (Gmelin, 1791),

which represents 90% of the catch produced along that

coast (Caeeres-Martinez and Vasquez-Yeomans, 2013).

Despite studies on diseases and ectoparasites of C.

virginica in Mexico (Aguirre-Macedo et al., 2007;

Caeeres-Martinez and Vasquez-Yeomans, 2013), a study

of non-native endo- and ecto-parasites of mollusks in the

coastal lagoons and coral reefs of the southern gulf is still

lacking. However, B. bisuturalis (Say, 1822) has been

registered by De la Cruz and Gcnzalez-Gandara (2006)

Table 2. Exotic freshwater mollusks recorded in Mexico, where are shown: place of origin, references or source, means of introduction (vector), habitat and impact with regard

to the damage they may cause;* = Naturalized,** = data provided by David G. Robinson.

Fami]y Species Origin Source/ record CNMO Vectors Habitat Impact

E. Naranjo-Garefa and Z.G. Castillo- Rodriguez, 2017

Page 111

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Page 112

THE NAUTILUS, Vol. 131, No. 2

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Page 120

THE NAUTILUS, Vol. 131, No. 2

Table 4. Species introduced in Mexico per region of origin

and percent which they represent.

on Lobos Reef, Veracruz. Since this species can survive

as an ectoparasite on various invertebrates, it could have

been introduced with species such as C. virginica as far

back as the last century. Its survival would have been

favored by the diversity of species that exist on the reef

plain of Veracruz; hence, B. bisuturalis is likely to be

more common in the region than suggested by this single

record. Its planktotrophic larvae (Robertson and Mau-

Lastovicka, 1979) would enable B. bisuturalis to be

transferred in ballast water and install itself on various

maerobenthic species. Unfortunately, it has not been

recorded as introduced to Mexico, perhaps as it is

considered by some to be native/naturalized, or because

the research has been limited to compilation of a simple

checklist. Boonea species can seriously affect oyster

fisheries and aquaculture (Wilson et ah, 1988; Gumming

and Alford, 1994), so that establishment large pop¬

ulations of B. bisuturalis should be considered as a po¬

tential threat to populations of the oysters in Mexico.

Cenchritis muricatus (Linnaeus, 1758) is common in

the Caribbean, southern Florida, and the Bahamas

(Clench and Abbott, 1942; Abbott, 1954; Trussell. 1997),

where it is distributed from the shoreline to a depth of

about 3.6 in (Lang et ah, 1998; Emson et ah, 2002). It

withstands desiccation and extreme heat at low tide. It was

first recorded in 1992 in the northern part of the Gulf of

California.

Diala albugo (Watson, 1886) is a small gastropod of the

Indo-Paeific that, as evidenced by its protoconch, pos¬

sesses a planktotrophic larva (Ponder and De Keyzer,

1992). It was included by Aguilar-Estrada et al. (2014) in

a checklist of a reef community in Veracruz that con¬

sisted mainly of dead specimens. It is not officially

registered as introduced in Mexico and, because its

identification is difficult, it is not included here nor in

Table 1.

Nudibraneh gastropods (Heterobranehia) of the genus

Anteaeolidiella Miller, 2001, A. foulisi (Angas, 1864), A.

cacaotica (Stimpson, 1855), and A. indica (Bergh, 1888),

recorded in Mexico (Hermosillo et al., 2006; Hermosillo

and Gosliner, 2008; Hermosillo, 2009), are not considered

here because the systematic^ of the Aeolidiidae has been

undergoing review on the basis of morphological and

molecular data (Carmona et id., 2013); those studies will

affect the distribution records of previously unrecognized,

potentially cireumtropical species (Angel Valdes, personal

communication).

Freshwater Mollusks

The gastropod Tarebia granifera (Lamarck, 1816) from

Madagascar, India, and Asia, is ovoviviparous, repro¬

ducing by parthenogenesis, and matures to a short length

(5.5 to 8.0 mm) (Appleton et al., 2009), features that are

advantageous in competing with native species following

invasion events. In other countries, it can displace native

species that display similar habitat requirements, such as

those in the genus Pachychilus Lea, 1850. In the Ca¬

ribbean and South America, it displaces and/or regulates

populations of species in the genus Biomphalaria Preston,

1910 (Pointier and Augustin, 1999; Pointier et al., 1998).

Its high reproductive potential allows it to quickly invade

bodies of water where it is introduced, and to literally

modify its physical conditions; in Mexico, it is considered

an invasive species (CONABIO, 2015).

Melanoides tuberculata (O.F. Muller, 1774) (gastropod)

and Corbicida fluminea (O.F. Muller, 1774) (bivalve), both

originating from Asia, are widely distributed in Mexico

(Contreras-Arquieta et al., 1995; Contreras-Arquieta, 1998;

Contreras-Arquieta and Contreras-Balderas, 1999) and are

recognized as invasive species (CONABIO 2015). Mela¬

noides tuberculata is ovoviviparous and reaches maturity at

about 3.5 mm or in about six months (Guticrrez-Amador

et al., 1995; Appleton et al., 2009), which has allowed it

to produce large populations very quickly. Corbicula

fluminea tolerates changes in environmental conditions

(Avelar et al., 2014), feeds on suspended material by

filtering and through pedal feeding, and influences the

abundance of surrounding benthic and pelagic fauna, as

well as the organic-matter cycle (Hakenkamp et al., 2001).

It may be possible that C. fluminea competes for space

with native clams of the family Unionidae (Britton and

Fuller, 1979).

Pomacea canaliculata (Lamarck, 1822) originally from

Argentina, South America (Cowie and Thiengo, 2003),

was registered for the first time in the wild in Mexico in

2013. It is presumed that the population in Mexico came

from the Colorado Riser, since it was located in 2005 in

that river in Yuma, Arizona, and the Colorado River

continues its course in Mexico (Campos et al., 2013). In

addition, specimens of apple snails found in California and

Arizona have been confirmed to be P. canaliculata based

on 46 unique mtDNA haplotypes (Rawlings et al., 2007).

The presence of non-native apple snails is of great concern

due to their ability to spread fast and because they are

E. Naranjo-Garcfa and Z.G. Castillo- Rodriguez, 2017

Page 121

recognized as agricultural pests (e.g., in rice fields in Asia).

Invasion of exotic apple snails poses a treat to marshland

habitats, with the possibility of changes in their diversity

and ecological processes, as happened in Laos, South East

Asia (Carlsson and Lacoursiere, 2005; Rawlings et ah,

2007). Apple snails are listed among the world’s 100 worst

invasive species (Lowe et ah, 2000). Its amphibian status

and herbivore habits are conducive to its establishment

and possibly to its success in environments such as rice

fields in Japan, Philipines, China, and other Asiatic

countries (Thiengo et ah, 1993; Cowie et ill., 2006;

Rawlings et ah, 2007; Ziyuan and Yuansheng, 2012). From

the human health point of view, P. carudiculata, along

with various freshwater or terrestrial mollusean species, is

an intermediate host of the low specific host nematode

Angiostrongylus cantonensis (lung worm); in nature its

definite hosts are several species of rodents. Pomacea

flagellata (Say, 1829), a native of the Gulf of Mexico states,

was introduced to the Pacific coast. Its present distribu¬

tion has facilitated the expansion of the distribution of the

snail-eating kite Rostrhamus sociabilis major Nelson and

Goldman, 1833 by about 900 km into the Pacific region

(Hernandez-Vazquez et ah, 2013). In addition, the “ear-

rao” Animus guarauna clolosus Peters, 1925 (naturally

distributed in the States of Veracruz, Chiapas, and

Yucatan) is now found in Laguna del Tule, Barra de

Navidad, Jalisco, on the Pacific coast (Hemandez-Vazquez

et ah, 1999; Palomera-Garcia et ah, 2006). Dispersals of

introduced of species may affect native species of birds in

the longer term.

Pomacea diffusa Blume, 1957, a species originally from

the region of Santa Cruz, Bolivia (Cowie and Thiengo,

2003) has been intercepted in shipments arriving in

the USA from an unspecified part of Mexico (David

Robinson, personal communication). As Howells et ah

(2006) pointed out, “introduced species pose a serious

threat to native biodiversity, second only to habitat loss”.

The Zebra Mussel Dreissena polymorpha (Pallas, 1771)

is a great ecological threat (Schloesser and Schmuekal,

2012) and is among the 100 worst invasive species (Lowe

et ah, 2000). Young specimens have been found at two

sites in the State of Veracruz: Rio Tonala, San Jose, and

Rio Coatzacoalcos, Napa Creek. (CNMO 3257: Las

Choapas. Rio Tonala, San Jose. Veracruz; and CNMO

6060: Rio Coatzacoalcos, Arroyo Napa, Veracruz. So far,

these are the first record of the species in Mexico.)

Freshwater species already naturalized in Mexico are:

Pomacea canaliculata, Pomacea flagellata (in the western

side of the country where it was not native), Melanoides

tuberculata, Corbicula fluminea, Tarebia granfera and

Radix auricularia (Linnaeus, 1758).

Terrestrial Mollusks

The veronicellid slug Sarasinula plebeia (P. Fischer,

1868), originally from New Caledonia (Gomes and

Thome, 2004), is gregarious, able to self-fertilize, and

oviparous, attributes that favor its potential as invasive

species. The species is thought to have displaced native

counterparts in Central America (Caballero et ah, 1991).

It is a serious pest of assorted agricultural crops in the

southern Catemaco Region, Veracruz (Naranjo-Garcfa

et ah, 2007), and of vanilla in the northern Veracruz

State (Velazquez- Montes de Oca et ah, 2014).

Phyllocaulis gayi (P. Fischer, 1871) is known to occur

in Valdivia, Chile (Thome, 1971, 1976). However, the

species was recorded in the city of Mazatlan, Sinaloa,

Mexico in by Baker (1925). Baker suggested that the

introduction of the species was probable due to

Mazatlan’s status as one of the main Pacific ports in

Mexico. In 2006, Naranjo-Garcfa visited the city of

Mazatlan looking for veronicellid slugs, but was unable to

confirm the occurrence of P. gayi (Naranjo-Garcfa et ah,

2007). The record of P. gayi in Mazatlan was either

a misidentification or, if it was present at that time, it did

not succeed there.

Cornu aspersion (O.F. Muller, 1774), originally from

Europe, tends to be gregarious and to produce large

numbers of offspring, attributes that have made it an

invasive pest in gardens in Mexico City (Ancona, 1947).

They have also destroyed orchards and gardens elsewhere

in Mexico (Maria Villaroel, personal communication), as

they have done in other countries (Apablaza, 1984; Cowie,

2000). Introduction may be at the egg stage or as juveniles

on imported plants; it was re-introduced into Mexico

(1991, CNMO 153) in imported strawberry seedlings, and

lias become a pest on cabbage crops in the State of

Michoacan (Naranjo-Garcfa, unpublished data).

With regard to slugs, Boetgerilla pattern Simroth, 1912

(Table 3) is believed to be a pest in gardens and green¬

houses in Europe (Welter-Schultes, 2012). Judging by

their background record in other places of the world,

Deroceras reticulation (O.F. Muller, 1774), Lehmannia

valentiana (Ferussac, 1821), Linuicus jlavus (Linnaeus,

1758), L. maximus Linnaeus, 1758, Rumina clecollata

(Linnaeus, 1758) are potential pests on crops and gardens.

Deroceras reticulation is a very destructive slug that feeds

on various cultivated plants (particularly at the seedling

stage) such as cauliflower, cabbage, potato (Pilsbry, 1948;

Castillejo, 1998). Hausdorf (2002) believes that Deroceras

invadens Reise, Hutchinson, Schunack and Schlitt, 201 1 is

a serious pest.

Some authors consider Deroceras laeve (O.F. Muller,

1774) to as introduced in Mexico. However, there are

fossil shells of what some believe to be this species of slug

in Canada, the USA, and Mexico (El Cedral, San Luis

Potosf; Olivera-Carrasco, 2007). If that is the case, the

species has then been in North America since the

Pleistocene. In Mexico, its two morphs are present, phallic

( = euphallic) and aphallic. The species is well-suited with

diverse life history traits that ensure it leasing de¬

scendants: it has a short life cycle, presents the two

morphs, auto-fertilize, and, in rare occasions, present

outcrossing. It is also tolerant of diverse ecological regi¬

mens and, under appropriate conditions, can reproduce

all year long (Gomez, 2001; Jordaens et ah, 2006). Der¬

oceras laeve has been observed that become a pest in

green houses (Wiktor, 2000).

Page 122

THE NAUTILUS, Vol. 131, No. 2

The snail Rumina decollata in the area of Santiago,

Nuevo Leon has been associated with crops of squash,

onion, and cucumber (Correa-Sandoval, 1993).

Terrestrial species are Saras inula plebeia. Cornu

aspersion, Paralaoma servilis (Shuttleworth, 1852), Arion

circumscriptns , Deroceras laeve, Deroceras reticulatum,

Lehmannia valentiana, Limacus flavus, Umax maximus,

Rumina decollata, Oxychilus draparnaudi (Beck, 1837)

and Zonitoides arboreus (Say, 1816).

CONCLUSIONS

There are 56 species of mollusks introduced in Mexico.

These records are confirmed by live material deposited in

collections and from the literature. This number may

increase as searches intensify. Until now, such information

is contained in works consisting mostly of lists that do not

indicate the status of the species treated and whether they

are exotic (non-native) or invasive (established and nat¬

uralized). Species intentionally brought for aquaculture

may contribute to the introduction of associated, poten¬

tially invasive species. Examples of this potential are

mollusks with a byssus (mussels) and with live epifauna on

the upper surface of an oyster.

Between 1980 and 2009, fewer than five authors have

recorded living exotic species on the Mexican Pacific

coast. Existing legislation should be applied rigorously or

improved ( Ortiz- Monasterio, 2014). Administration and

management of ports and customs must protect the na¬

tional territory and conserve biodiversity. Introduction of

species can harm life cycles of other taxa, with direct

damage to human health and the national economy.

Some of the mollusks here mentioned may succumb

under the effects of climate change, but others may

survive and colonize areas where they currently cannot

survive due to the constraints imposed by the climate.

Hence, it is important to know the status of each species,

and to monitor their presence and effects over habitats

and native wildlife at national level. Molecular studies will

further add to the knowledge of the systematic^ and

population structure of these alien species.

Mollusks are among the most biodiverse groups of

invertebrates, and the creation of a Mexican monitoring

network devoted to exotic species and their effects would

help to protect native endemic species and could examine

with scientific basis the effects of introduced organisms on

human health and environment.

ACKNOWLEDGMENTS

Thanks to David Robinson (USDA APHIS) for sending us

a list of mollusks intercepted at the border of the United

States and Mexico. Maria Teresa Olivera-Carrasco con¬

tributed to this study. Timothy A. Pearce gave us data of

Vallonia cost at a deposited at the Carnegie Museum.

Georgina Leite and Miguel Angel Martinez, Institute of

Biology provided us with literature of difficult location.

Angel Valdez (California State Polytechnic University,

Pomona, USA) advised us on the presence or absence of

introduced marine slugs. Ann Grant revise our English.

David Robinson and another anonymous reviewer made

comments that greatly improve our manuscript.

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Abbott, R.T. 1974. American Seashells, 2nd edition. Van

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Aguilar-Estrada, L., D. Ortigosa, B. Urbano, and M. Reguero.

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arrecifal de Isla Verde, Veracruz, Mexico. Revista Mexicana

de Biodiversidad 85 (2): 502-512.

Aguiar, P.H.. P. Morera and J. Pascual. 1981. First record of

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THE NAUTILUS 1 3 1 (2): 127- 1 37, 2017

Page 127

A new species of Lucinoma from 240-500 m on the continental

shelf break off Newfoundland (Bivalvia: Lucinidae)

John D. Taylor

Emily A. Glover

Department of Life Sciences

The Natural History Museum London SW7 5BD,

UNITED KINGDOM

[email protected] [email protected]

ABSTRACT

A new species of the lucinid bivalve genus Lucinoma is described

from shells dredged at depths of 240-500 in from the edge of the

continental shelf off southern Newfoundland. It differs from the

other northern species, Lucinoma filos a, in shape, ligament, and

characters of the anterior adductor muscle scar. It also differs

from the poorly known Lucinoma atlantis from the outer shelf

off Maryland that is longer than high and Inis both anterior and

posterior sulci, and from L. blakeana from deep water off North

Carolina, a smaller species with a truncate posterior margin.

Other Lucinoma species are recorded further south in the

northern Gulf of Mexico, particularly from hydrocarbon seeps,

although the taxonomy is confused for those taxa.

INTRODUCTION

The chemosym biotic Lucinidae is now recognized as one

of the most speciose of marine bivalve families (Huber,

2015; WoRMS) and further new species and genera

continue to be described (Glover and Taylor, 2016).

Lucinoma, with at least 30 living species, is the best known

and widespread of the deeper-water lucinid genera. The

genus has a latitudinal range from 70° N to 55° S and from

the intertidal zone to deeper than 2500 m (summary

figure in Taylor and Glover, 2010, fig 5.9). The majority of

Lucinoma species are found from >200 m to mid-bathyal

depths and are often abundant at hydrocarbon seeps, mud

volcanoes and oxygen minimum zones (Cary et ah, 1989;

Okutani and Hashimoto, 1997; Callender and Powell,

1997, 2000; Salas and Woodside, 2002; Olu-Le Roy

et ah, 2004; Holmes et ah, 2005; Cosel, 2006; Cosel and

Bouehet, 2008; Oliver and Holmes, 2006; Dupe iron et ah,

2007; Oliver et ah, 2012; Zamorano and Hendrickx, 2012).

Around 40% of Lucinoma species have been described

within the last 1 5 years suggesting that diversity has not yet

been fully sampled. Nonetheless, it should be noted that

some Lucinoma species reported from Japan (Okutani

and Hashimoto, 1997), L. aclamsiana Habe, 1958 and L.

japonica Habe, 1958, are now classified in other genera

and subfamilies (Glover and Taylor, 2016).

The phylogenetic placement of Lucinoma has been

problematic. An initially surprising result from molecular

analyses of Lucinidae was that Lucinoma species grouped

within a major clade (subfamily Codakiinae) of otherwise

largely shallow water species of Ctena and Codakia

(Taylor et ah, 2011; 2014; 2016). Previously, using shell

characters, Dali (1901) had classified Lucinoma as

a subgenus of Phacoides, a placement also followed by

Britton (1970). By contrast, Chavan (1938; 1969) con¬

sidered the genus to be related to Myrtea and placed it in

the Myrteinae (also Abbott, 1974), while Bretsky (1976)

classified it as a subgenus of Miltha. At species level

Lucinoma are difficult to discriminate, but useful shell

characters are: overall shape, form of commarginal

sculpture, presence of posterior and anterior sulci,

thickness of hinge line and size of cardinal teeth, presence/

absence of anterior lateral teeth, and in particular,

characters of the anterior adductor muscle scar, notably

the length, width and angle of ventral detachment from

the pallial line.

On the western Atlantic continental margin three

species of Lucinoma have been described: Lucinoma

filosa (Stimpson, 1851), L. blakeana (Bush, 1893), and L.

atlantis (McLean, 1936). Unfortunately, only dead shells

are available for study and no Lucinoma species from the

Western Atlantic have been included in molecular ana¬

lyses. Recently, we located in the collections of the

Museum national d’Histoire naturelle, Paris, several

samples of a Lucinoma species collected in 1985 from

dredging on the edge of the continental shelf off southern

Newfoundland. These shells had been labelled as Luci¬

noma filosa, but differed from that species and also from

the two other Lucinoma species described from the

northwestern Atlantic. The samples were obtained during

a survey of fish stocks (ERHAPS 851) in the territorial

waters of Saint-Pierre and Miquelon, a small archipelago

of French territory ( Collectivite d’Outre-mer de Saint-

Pierre-et-Miquelon ) off the southern coast of New¬

foundland. A narrow strip (10.5 nautical miles wide) of

territorial waters (Exclusive Economic Zone) extends

southwards across the continental shelf. Cosel (1986)

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THE NAUTILUS, Vol. 131, No. 2

published a semi-popular account of the ERHAPS 851

cruise and a list of the mollusks recovered. In this paper

we describe this new species of Lucinoma and compare it

with congeners from the region.

MATERIALS AND METHODS

The length of anterior adductor muscle scars in relation to

shell height and length were measured on shells using

calipers, while measurements from L. filosa were mainly

taken from scaled images of shell interiors from speci¬

mens in ANSP, USNM, and MCZ (latter images courtesy

of G. Giribet and A. Raldinger). Similar measures were

made from images of the holotypes of L. blakeana and L.

atlantis. Outline drawings of shell interiors were made

from digital images using Photoshop.

Institutional acronyms used: ANSP, Academy of Nat¬

ural Sciences Philadelphia at Drexel University, USA;

MCZ, Museum of Comparative Zoology, Harvard Uni¬

versity, Cambridge, USA; MNIIN, Museum national

d'Histoire naturelle, Paris, France; NHMUK, The Nat¬

ural History Museum, London, U.K.; USNM, United

States Museum of Natural History, Washington, USA.

Other abbreviations: aas, anterior adductor muscle scar;

fms, fathoms; H, shell height; L, shell length; LV, left

valve; pv, pain'd valves; RV, right valve; v, single left or

right valve.

SYSTEMATICS

Bivalvia

Family Lucinidae Fleming, 1828

Subfamily Codakiinae Iredale, 1937

Genus Lucinoma Dali, 1901

Type Species: Lucina filosa Stimpson, 1851 Original

designation

Description: Medium to large shells. Subcircular to

anteriorly tapering. Posterior sulcus with marginal sinus

present in some species, anterior sulcus rarely present.

Sculpture of narrow, regularly spaced, commarginal la¬

mellae with finer lamellae between, radial sculpture ab¬

sent. Lunule lanceolate, slightly sunken. Ligament long,

external, protruding or in groove. Hinge with two cardinal

teeth in each valve, larger teeth usually bifid, small to

obscure anterior lateral teeth. Anterior adductor muscle

scar usually long and detached from pallial line for about

% of length. Inner shell margin smooth.

Lucinoma thula new species

(Figures 1-13, 14-15)

Description: L to 70 mm, H to 63 mm, slightly longer

than high (H/L=0.94±0.04, SD, n = 28). Robust, sub-

circular, shallow posterior sulcus, posterior margin truncate

with shallow sinus, anterior margin slightly projecting,

ventral margin broadly rounded. Umbones broad, low,

prosogyrate. Periostracum: thick, dark olive-brown.

Sculpture: widely spaced, low, sharp, commarginal lamellae

with .5-7 finer commarginal “cords” in interspaces. Major

lamellae are more closely spaced ventrally in larger

shells. Lunule: long, lanceolate. Ligament: external,

long, set on nymph. Hinge: RV with two cardinal teeth,

anterior thin, posterior huger, slightly bifid, lateral teeth

absent; LV with two cardinal teeth, anterior larger,

slightly bifid. Anterior adductor muscle scar long,

ventrally detached from pallial line at an angle of

about 20°; posterior adductor scar ovoid, anterior pedal

retractor muscle scar separate and dorsal to anterior

adductor scar. Pallial line entire, shell within line with

subcircular scars of mantle attachment, pallial blood

vessel trace visible, inner shell margin smooth.

Type Material: All type material from ERHAPS 851

cruise N.O. CRYOS; Holotype: One whole shell. Station

L219, 345-512 m, 45°0L N, 54°57’ W, 09 March 1985,

L=61 .3 mm, H=58.1 mm, tumidity single valve = ! 5.2 mm,

MNIIN I M -2000-33 102; Paratypes: MNHN IM-2000-

33103, station L181, 310-308 m, 46°32’ N, 57°3L W, 01

March 1985, 6 RV 9 LV (L=69.8, 57.2, 50.8, 54.8, 47.0,

45.8, 44.5, 40.3, 33.5 mm); MNHN I M -2000-33 104, station

LI 84, 243-244 m, 46°2T N, 57°2T W, 01 March 1985, 1 LV

(L=55.6 mm); MNHN IM-2000-33105, station L185,

314-320 m, 46° 18’ N, 57°22’ W, 01 March 1985, 1RV,

1LV, (L=53,5, 53.3 mm); MNHN I M -2000-33 106, station

I486, 282-278 m, 46°12’ N, 57°17’ W, 01 March 1985, 2

RV, 3LV (L=65, 55.2, 52.6, 51,5, 49,7 mm); MNHN IM-

2000-33107, station L187, 323-322 m, 46°05’ N, 57°12’ W,

01 March 1985, 9 LV, 5 RV (L=56.3, 55.6, 52.8, 48.3, 47.7,

46.1, 44.6, 42.2, 42.1, 40.1, 37.9 mm); MNHN IM-2000-

33108, station L189, 332-320 m, 46°60’ N, 57°04’ W, 02

March 1985, 2 pv (L=53.4, 36.6 mm); NHMUK 20170141,

station L187, 323-322 m, 46°05’ N, 57° 12’ W, 01 March

1985, two single valves (L=46.9, 44.3 mm).

Habitat: Recorded from depths of 240-512 m near the

edge of the continental shelf off southern Newfoundland.

Rudo von Cosel (personal communication) reported that

the sediment was fine to coarse sand and the accompa¬

nying fauna comprised “...regular sea urchins, lots of

ophiurids, a few shrimps and large actinias, and among the

mollusks mostly Buccinidae of different genera and

species (17 in total).” See Cosel (1986) for a list of other

mollusks recovered from the ERHAPS cruise but without

details of individual stations.

Etymology: Derived from Latin thule for furthest

north, in reference to the northerly location of the species.

Comparison with Other Species: The three Luci¬

noma species previously described from the north¬

western Atlantic are illustrated in Figures 19-31 and

their principal features documented below. The type

localities of the northwestern Atlantic species are

plotted in Figure 33.

J.D. Taylor and E.A. Glover, 2017

Page 129

Figures 1-13. Lucinoma thula new species, all specimens from ERHAPS 851 cruise. 1-5. Holotype (MNHN I M -2000-33 102),

station L219, L = 61.3 mm. 1-2. Exterior and interior of left valve. 3—4. Exterior and interior of right valve. 5. Dorsal view. 6-7. Paratype

(NHMUK 20170141), station L187, interior and exterior of left valve, L=56.1 mm. 8-10. Paratype (MNHN IM-2000-33103), station

L181, exterior of right valve and interior of right and left valves, L=38.3 mm. 11-12. Paratype (MNHN IM-2000-33106), station L186, exterior

and interior of left vah'e, L=50.9 mm. 13. Paratype (MNHN IM-2000-33106), station L 186, exterior of left valve, L=64.1 mm.

Page 130

THE NAUTILUS, Vol. 131, No. 2

Figures 14-18. Internal drawings of valves. 14-15. Lucinoma thula new species. 14. Holotype 15. Paratype (NHMUK 201 70141 ).

16. L. filosa (ANSP 102172). 17. L. atlantis, holotype. 18. L. blakeana , holotype.

Lucinoma filosa (Stimpson 1851)

(Figures 16, 19-24, 34-35)

Brief Description: Shell length to 60 mm, subcircular,

slightly longer than high (H/L=0.88±0.02 SD, n = 12),

sculpture of regular, sharp commarginal lamellae. Slight

posterior and anterior sulci. Lunule long, slightly sunken,

Anterior dorsal margin elevated. Ligament in a groove

with a deep escutcheon. Two cardinal teeth and vestigial

anterior lateral tooth in each valve. Anterior adductor

muscle scar very long and narrow, detached from pallial

line for 4/5 of length (details below Figure 32).

Periostracum relatively thin, pale tan, or buff.

Type Material: Not located (see Britton 1970, Bretsky

1976). The type locality (Stimpson, 1851:17) is cited

as “...in 6 f sand near Pt Shirley (W.S.), Phillips Beach,

alive after a storm (Holder)”. Point Shirley is now in the

town of Winthrop, a suburb of Boston, Massachussetts.

Distribution: Lucinoma filosa is distributed from

Canada to Florida Keys and maybe into the Gulf of

Mexico. It is well represented in collections from northern

areas with recorded depths of 20-80 m but occurs in

deeper water (to 400 m) further south off Florida Keys

(Britton, 1970) (NHMUK20140794, see Figure 34-35). It

was also recorded as common on the continental shelf in

the Middle Atlantic Bight area, in less than 200 m,

between Cape Cod and Cape Hatteras (Wigley and

Theroux, 1981: 95, fig. 76).

Lucinoma atlantis (McLean, 1936)

(Figures 17, 25-27)

Brief description: Holotype is longer than high, H/

L=0.81, with widely spaced thin commarginal lamellae.

Prominent anterior sulcus and posterior sulcus with

marginal sinus. Larger of the two cardinal teeth bifid,

distinct anterior lateral teeth in both valves. Anterior

adductor muscle scar long, detached from pallial line for

% of length. Periostracum thick and olive brown.

Type Material: Holotype, MCZ 73345, PV L=58.3 mm,

H = 47.3 mm

Type Locality: Off Maryland, 216-549 m, 38°10’ N,

73°5T W (note coordinates on holotype label are in¬

correct) at edge of continental shelf.

Distribution: Known only from the holotype and we

have seen no other comparable material. Bivalves iden¬

tified as this species are widely reported from sites of

hydrocarbon seeps off Louisiana in the northern Gulf of

Mexico (Figures 39-46) (Turner, 1985: fig 2H;

MacDonald et ah, 1990; Callender et ah, 1990; Callender

and Powell, 1997) hut specimens we have examined from

|.D. Taylor and E.A. Glover, 2017

Page 131

Figures 19-31. Lucinoma filosa , L. atlantis and L. blakeana. 19-21 .Lucinomafilosa, off Martha’s Vineyard (ANSP 102 1 72), L=39.4

mm. 22-23. L. filosa (USNM 45937) exterior and interior of left valve, off Martha’s Vineyard, 144 m, USFC stn 941, L=47 mm. 24. L

filosa (MCZ 41558), interior of left valve, off New Jersey, 40° 01' N, 70°30’ W, 225 m, L=30.3 mm. 25-27. Lucinoma atlantis, holotype

(MCZ 73345) exterior of right valve and interior of right and left valves, off Maryland, 216-549 m, 38°10’ N, 75° 51’ W, L=58.3 mm.

28-29. Lucinoma blakeana , holotype (MCZ 1 19129), exterior and interior of right valve, off Cape Fear, North Carolina, 464 fms (850

m), Blake station 326, 33°42’15" N, 76 °00’50" W, L=30 mm. 30-31. L. blakeana (USNM 95694), exterior and interior of left and right

valves, albatross Station 2677, 874 m, off Gape Fear, 32°39’ N, 76°50’ W, L = 41.6 mm.

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THE NAUTILUS, Vol. 131, No. 2

= 0.5

x:

c

tJ 0.45

L. filosa

L thula

A i

A

A A

A

A A

A

35 45

shell length mm

Figure 32. Comparison between Lucinoma thula and L. filosa

in the relative length of the anterior adductor muscle scar (length

of aas/shell height) plotted against shell length.

this region differ from the holotype of L. atlantis (despite

the opinion of Turner, 1985) in lacking the anterior sulcus

and thick, dark periostracum and likely represent a sep¬

arate species.

Lucinoma hlakeana (Bush, 1893)

(Figures 18, 28 -31, 36 -38)

Type Material: Holotype MCZ 119129, single RV

L=30.1 mm, H = 26.9 mm, H/L=0.86.

Type Locality: Off Cape Fear, North Carolina, 850 m,

Blake Expedition, 1880, station 326, 33°42’ 15" N,

76°00’50’ W.

Brief Description: Small species L to 30 mm longer

than high, H/L=0.86, with a distinctive quadrate posterior

margin and shallow posterior sulcus. Sculpture of widely

spaced, thin, commarginal lamellae. Lunule long and

lanceolate. Small anterior lateral teeth present in both

valves and two cardinal teeth, larger bifid. Anterior ad¬

ductor muscle scar long, vent rally detached from pallial

line for 3A of length. Periostracum of holotype is now tan

brown hut it was originally described (Bush, 1893) as thin

and light yellow.

Remarks: A further Lucinoma species is represented

by some shells from the northern Gulf of Mexico collected

from Viosca Knoll lease block 826 (29°09’ N, 88°0T W,

depth ea. 450 m, off Alabama) and sent to Ruth Turner for

identification (MCZ). These are illustrated in Figures

47-50. This is a large (L=81 mm, H = 80 mm),

rounded, inflated species, without significant sulci and

with low commarginal lamellae that are much more

closely spaced than the other Lucinoma from the

northwestern Atlantic. The cardinal teeth are large with

a small anterior lateral tooth in the left valve and the

anterior adductor muscle scar is medium-length and

broad. This species differs from the putative L. atlantis

of the Gulf of Mexico and other western Atlantic

Lucinoma species in shape, lack of sulci and the closely

spaced lamellae. The taxonomy of all the Lucinoma

species recovered from hydrocarbon seeps in the Gulf

of Mexico remains problematic.

One of the main features that differentiates the new

species, Lucinoma thula, from L. filosa is the length and

position of the anterior adductor muscle scar; this is long

and narrow in L. filosa, lying approximately parallel with

the pallial line (Figure 16) and ventrallv detached for 4/5

of length at an angle of about 12°. By comparison, in L.

thula the adductor scar is shorter, broader, and diverges

from the pallial line at an angle of about 20°. The relative

lengths of the anterior adductor scar (as length/shell

height) were compared: mean 0.46±0.3 SD, n = 27 for

L. thula and 0.53 ± 0.2 SD, n=ll for L. filosa (Figure 32).

These were significantly different (T test, p >0.05).

Lucinoma filosa has more closely spaced commarginal

lamellae and is subcircular in outline but L. thula is

posteriorly truncate. In most shells of L. filosa the postero-

dorsal shell margin extends above the ligament (Figures

19-24), hut not in L. thula. The periostracum is thicker

and dark in L thula compared to the pale and relatively

thin periostracum of L. filosa.

The holotype of Lucinoma atlantis differs from L. thula

in its markedly longer than high shell outline, distinct

anterior and posterior sulcus (Figures 25-27), and visible

anterior lateral teeth in both valves. Lucinoma hlakeana is

a smaller species (Figures 28-29) that differs in shape

from L. thula and L. filosa with a quadrate posterior

margin, only a slight posterior sulcus and a tan brown or

yellowish periostracum compared with the darker olive

brown of L. thula and L. atlantis.

DISCUSSION

Distribution: Other specimens, also from off Cape

Fear are USNM 92670, Albatross station 2628, 966 m,

and USNM 95694, Albatross station 2677, 874 m, and

similar shells are recorded from off Havana (Figures

36-38), USNM 64435, Blake station 43, on label as

24°08’ N, 82°5F W, 449 fms (821 m); but the blake

station list in USNM records 339 fms (620 m) and 83°51’ W

for station 43. The few confirmed records available suggest

that this is a much deeper water species than L. filosa and

L. thula.

It is probable that Lucinoma thula has been confounded

with L. filosa in faunal surveys of the outer continental

shelf and upper slope of the northwestern Atlantic but we

have seen no other specimens in MCZ, ANSP, or USNM

collections. From a macrobenthie survey of the north¬

eastern USA continental shelf and slope, Theroux and

Wigley (1998) recorded L. filosa, L. hlakeana and Luci¬

noma sp. hut gave no images or details.

Ideally, the taxonomic discrimination of Lucinoma

species should be corroborated with molecular data. To

J.D. Taylor and E.A. Glover, 2017

Page 133

Figure 33. Map showing type localities of the Lucinoma

species described from the eastern Atlantic seaboard of North

America, and locations in the Gulf of Mexico of shells illustrated

in Figures 34-52. Map from Google Earth Image 2017.

date only six species of Lucinoma have been included in

molecular analyses; these originated from widely sepa¬

rated locations and depths but with none from the

northwestern Atlantic. The species group in a well-

supported clade with short branches and form a sister

group to Codakia species (Taylor et ah, 2016). Inclusion of

the type species, Lucinoma filosa, and the other species

from the western Atlantic is highly desirable. Neverthe¬

less, the new species L. thula clearly differs morpholog¬

ically from L. filosa and we are confident of its distinct

identity. The northeastern Pacific species Lucinoma

annulata (Reeve, 1850) has similar shape, sculpture, and

musculature to L. filosa and may be a sister taxon. This

species likewise occurs in shallow subtidal habitats at the

northern end of its range in Alaska and in deeper water

(665 m) at more southerly locations off Mexico (Coan

et ah, 2000). Described from a methane seep area in the

eastern Pacific Ocean off Concepcion, Chile, Lucinoma

anemiophila (Holmes, Oliver, and Sellanes, 2005) has

a general similarity to L. thula but with more prominent

anterior lateral teeth.

Recent surveys have revealed the presence of nu¬

merous sites of seafloor methane leakage along the

northern US Atlantic margin (Skarke et ah, 2014)

and cold seep communities reported at depths as

shallow as 400-430 m in the Baltimore Canyon (http://

oceanexplorer.noaa.gov/explorations/12midatlantic/

logs/aug26/aug26.html) and others off Nantucket at

1 100-1400 m (Quattrini et ah, 2016). It is quite possible

that the original specimens of L. atlantis and L. blakeana

dredged from the edge of the continental shelf off

Maryland and North Carolina (Figure 33) respectively

might have been associated with cold seeps. Certainly, the

latter species was recovered in the vicinity of extensive

methane venting from the Blake Ridge gas hydrate

province (Brothers et ah, 2013). Similar sites of methane

leakage are likely at the edge of the continental shelf off

southern Newfoundland but there have been no biological

surveys published, and, although it is possible that L. thula

samples were trawled from such a site, we have no direct

evidence. Cosel (1986) listed the macrofauna from the

ERHAPS 851 survey but gave no details of individual

stations; of 45 bivalve species recorded two other possible

chemosymbiotic bivalves are Solemija borealis Totten,

1834 and Thyasira sp. (not all species harbour symbi¬

onts) but it is uncertain whether or not they co-occurred

with L. thula.

Further to the south in the Gulf of Mexico several

Lucinoma species have been reported associated with

hydrocarbon seeps including putative L. atlantis

(MacDonald et ah, 1990; Conies et ah, 2000). In¬

cidentally, some Lucinoma sp. reported from the hy¬

drocarbon seeps are now assigned to other genera namely

Jorgenia and Graecina and classified in the Myrteinae

rather than Codakiinae (Taylor and Glover, 2009). In the

southern Caribbean, Gracia et id. (2012) reported three

Lucinoma species (unnamed) from cold seeps at around

500 m off Colombia and, at depths of 230-800 off

Guadeloupe, Taylor and Glover (2016) recorded two

unnamed species from dead shells in poor condition.

Globally there is a strong association of Lucinoma

species with seeps, as well as mud volcanoes and oxygen

minimum zones. Lucinoma myriamae (Coseh 2006) oc¬

curs at seep sites from 360 —425 m off West Africa (Sibuet

and Vangreisheim, 2009); Lucinoma taiwanensis was

described from an area of known hydrothermal activity off

north Taiwan 205-650 m (Cosel and Bouchet, 2008),

Lucinoma yoshidai is reported at Japanese seeps between

100-1000 m (Okutani and Hashimoto, 1997) and Luci¬

noma anemiophila from 780 m at a methane seep off

southwestern Chile (Holmes et ah, 2005). From the

eastern Mediterranean, Lucinoma kazani was described

from 1700 m deep mud volcanoes (Salas and Woodside,

2002; Olu-Roy et til., 2004) and L. asaphaeus from mud

volcanoes in the Strait of Cadiz (Oliver et ah, 2012).

Additionally two species of Lucinoma are known from

sediments in oxygen minimum zones; L. aequizonata from

400 -650 m off southern California (Caiy et ah, 1989;

Zamorano and Hendrickx, 2012; Hendriekx et ah, 2016)

and Lucinoma gagei from southern Oman at 675-967 m

(Oliver and Holmes, 2006). Lucinoma aequizonata (Dali,

1901) has a remarkable tolerance of anoxia being able to

survive 262 days without oxygen (Amdt-Sullivan et ah,

2008).

Page 134

THE NAUTILUS, Vol. 131, No. 2

Figures 34-52. Lucinoma specimens from northern Gulf of Mexico. 34-35. Lucinoma filosa (NHMUK20140794), interior and

exterior of RV, SW of Marquesas Rock, Florida Keys, 24°20.62’ N, 82°16.41’ W, 185-195 m, L=35.5 mm. 36-38. Lucinoma blakeana

(USNM 64435), exterior of LV and interior of RV and LV, off Havana, Rlake station 43, 24°08’ N, 82°5T W, 821 m, L=22.3 mm.

39-40. Lucinoma “ atlantis ” (USNM 637445), interior and exterior of RV, Gulf of Mexico, 27°50’ N, 91°1T W, 375 m, L=38 mm.

41-43. L ucinoma “atlantis” exterior of LV and interior of RV and LV, Garcia collection no 24039, off Louisiana, Rush Hill seep site in

lease block Green Canyon 185, 27°46.904’ N, 91°30.286 W, 546-555 m. L=48.6 mm. 44-46. Lucinoma atlantis exterior of LV and

interior of LV and RV, Garcia colln no 24039, Off Louisiana, Bush Hill seep site in lease block Green Canyon 185, 27°46.904 N,

91°30.286 W, 546-555 m, L=53.5 mm. 47-50. Lucinoma sp. (MCZ), off Alabama, Viosca Knoll lease block 826, 29°09’ N, 88°0L W, ca.

450 m (Figures 47-48, L = 81 mm; 49-50, L=80 mm). 51-52. Lucinoma sp. (USNM 706740), off Tortugas (no details), 347-512 m,

L=46 mm.

j.D. Taylor and E.A. Glover, 2017

Page 135

The fossil history of hucinoma also reveals an associa¬

tion with hydrocarbon seep habitats. Although Lucinoma -

like lucinids first appeared in the early Cenozoic in shallow

water Paleoeene deposits (Taylor et ah, 2011) maybe as

a sister clade to Saxolucina, they have been associated

with fossil deep-water hydrocarbon seeps since the

Oligocene; for example, L. hannibali (Clark, 1925) (Kiel,

2010; Nesbitt et ah, 2013) and in the Italian Miocene

Lucinoma perusina (Sacco, 1901) (Moroni, I960). Massive

accumulations of Lucinoma shells are recorded at sites

of fossil seeps in the Miocene and Pliocene of Japan

with inferred palaeo-depths of 50-300 m (Majima et ah,

2003, 2005).

Continuing discoveries of hydrocarbon seeps, gas hy¬

drates, pockmarks, mud volcanoes (e.g., Cunha et ah,

2013; Quattrini et ah, 2015) shows that these habitats

are abundant along the margins of continental shelves.

The biological communities associated with many of these

have yet to be investigated and the focus of the better

known sites has concerned epifaunal or shallowly infaunal

bivalves, Bathymodiolus and Vesicomyidae (Olu-LeRoy

et ah, 2007; Cordes et ah, 2009) with deeper burrowing

taxa such as Lucinoma less well sampled or studied.

Improved sampling and accompanying molecular studies

should clarify the confused taxonomy of the Gulf of

Mexico and wider Caribbean Lucinoma species and their

relationships to the more northerly species along the

eastern USA continental margin. From the research ac¬

tivity at hydrocarbon seeps in the northern Gulf of Mexico

relatively few Lucinoma samples have been archived in

museum collections despite their apparent abundance, as

evidenced for example by Callender and Powell (1997).

ACKNOWLEDGMENTS

We are grateful to our colleagues in MNHN Paris,

Philippe Bouchet, Virginie Heros and Philippe Maestrati

for access to the collections and loan of material; to Rudo

von Cosel for information about the ERHAPS 851 cruise.

We also thank Gonzalo Giribet and Adam Baldinger

(MCZ) for access to collections and for sending scaled

images of L.filosa and L. blakeana, Ellen Strong (USNM)

for access to collections and with Liz Harper (U. of

Cambridge) for images of USNM shells of Lucinoma

blakeana. Emilio Garcia kindly loaned specimens of Gulf

of Mexico Lucinoma shells illustrated in Figures 41-46.

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THE NAUTILUS 131(2):138-146, 2017

Page 138

Bayerotrochus belauensis , a new species of pleurotomariid from the

Palau Islands, western Pacific (Gastropoda: Pleurotomariidae)

Patrick Anseeuvv

Mispelstraat, 18

9820 Merelbecke, BELGIUM

Lori J. Bell

Coral Reef Research

Foundation Box 1765

Koror, PW 96940, PALAU

M.G. Harasewych1

Dept, of Invertebrate Zoology, MRC-163

National Museum of Natural History

Smithsonian Institution

PO Box 37012

Washington, DC 20013-7012 USA

ABSTRACT

A new pleurotomariid species, Bayerotrochus belauensis new

species, collected from the Palau Islands, western Pacific, is

described and illustrated. This new species is most similar in shell

moqihology to B. teramachii (Kuroda, 1955), from which it may

be distinguished by its thinner, lighter shell with a taller, more

stepped spire and lack of pronounced spiral sculpture along the

shell base. Molecular data (COI) show B. belauensis new species

to be more closely related to B boucheti from New Caledonia

and B delicatus from Yap, than to B. teramachii. Bayerotrochus

boucheti (Anseeuw and Poppe, 2001) differs in having a broader,

more conical spire, a more depressed aperture, and a more

darkly pigmented shell with spiral sculpture on the shell base.

The recently described B. delicatus S.-P. Zhang, S.Q. Zhang, and

Wei, 2016 is easily distinguished by its much smaller size and

distinctive shell profile.

Additional Keywords: Slit shell, COI. phylogeny, ecology.

INTRODUCTION

Bayerotrochus Harasewych, 2002 is the most widely

distributed of the Recent pleurotomariid genera. It in¬

cludes 12 living species, two inhabiting bathyal depths

of the temperate and tropical western Atlantic, and ten

from similar depths along the margins of the Indian and

western Pacific Oceans. Ongoing sampling of deep-sea

biota using dredges and trawls, as well as manned and

remotely operated research submersibles, in addition to

analyses of relationships among samples using molecular

techniques, continue to expand and refine our knowledge

and understanding of the biodiversity and biogeography of

the deep oceans and their faunas.

Two samples of living Bayerotrochus were first ob¬

served and collected off Palau at depths around 200 m

by the Japanese research submersible HAKUYO in

1 Author for correspondence: [email protected]

October of 1996 (Okutani and Kurata, 1998). They were

provisionally identified as Perotrochus africanus ter¬

amachii (Kuroda, 1955) based on shell morphology.

During subsequent explorations at similar depths in Palau

in 2001 using the research submersible Deepworker

2000, Coral Reef Research Foundation (CRRF) was able

to observe, photograph, and collect additional specimens

of that species of Bayerotrochus. Shells and tissues of

several of these samples were deposited in the collections

of the National Museum of Natural History, Smithsonian

Institution (USNM). Most recently, Zhang et al. (2016)

described Bayerotrochus delicatus from Yap Seamount

(8°5L N, 137°47’ E), SE of Yap near Nguluu Atoll, col¬

lected at slightly greater depths (255-289 m). A phylo¬

genetic analysis based on partial COI sequence data that

was included in their description showed that B. delicatus

was more closely related to one of the specimens from

Palau (USNM 905395) than to B. teramachii.

These results prompted our re-examination of the

Palau specimens, which were compared morphologically

against all species of Bayerotrochus known from the

Indian and Pacific Oceans. Results of these morphological

comparisons as well as a phylogenetic analysis of COI

sequences derived from three of the Palau specimens and

additional Bayerotrochus sequences, both published and

previously unpublished, indicate that the Palau samples

differ from all other species of Bayerotrochus , and are

here described as a new species.

MATERIALS AND METHODS

Radular Morphology: The radula was dissected from

die alcohol-preserved holotype, cleaned in 10% NaOH

overnight, rinsed in distilled water, cleaned in an ultrasonic

cleaner, air (hied, coated with gold, and photographed using

a Leica StereoSean 440 Scanning Electron Microscope.

Molecular Procedures: Genomic DNA was extracted

from alcohol-preserved tissue samples (e. 25 mg buccal

muscle) using the DNeasy Tissue Kit (Qiagen) following

P. Anseeuw et al., 2017

Page 139

the maufaeturer’s animal tissue protocol. A 658 bp portion

of the cytochrome c oxidase I gene was amplified using the

primers of Folmer et al. (1994). PCR amplifications used

the Promega GoTaq hot start master mix (Promega

M7132) according to manufacturer’s instructions, hut

modified to reduce the reaction volume to 20 |jlL.

Cycling parameters: initial denaturation at 95°C for

3 minutes, followed by 45 cycles of denaturation at

94 °C for 30 seconds, annealing at 45°C for 45 seconds

and extension at 72 °C for 2 minutes, with final extension

at 72°C for 5 minutes. Resulting PCR products were

visualized by agarose gel electrophoresis (1.5% garose)

and purified with ExoSAP-IT (Affymetrix). Sequencing

reactions were performed using 1 piL of purified PCR

product in a 10 qL reaction containing 0.5 |xL primer,

1.75 p-L Big Dye buffer and 0.5 p,L Fig Dye (Life

Technologies). Reactions were run for 30 cycles of

95°C for 30 seconds 50°C for 30 seconds and 60°C for

4 minutes then held at 12°C. Reactions were purified

using Millipore Sephadex plates and sequenced on an ARI

3730XL automated DNA analyzer. Sequeneher v. 4.7

(Gene Codes) was used to visualize, trim and assemble

contigs from forward and reverse sequences. The

sequences have been deposited in GenBank (NCBI).

Accession numbers are provided in Table 1.

The COI sequences were aligned using MUSCLE

within Geneious (v. 9.1.6;http://www.geneious.com,

Kearse et al., 2012) and trimmed to the 658 bp region

flanked by the Folmer primers. The phylogenetic tree

(Figure 28) was produced using the RAxML 7.2.8 (GTR

Gamma nucleotide model) plugin in Geneious 9.1.6.

Institutional acronyms are listed on Table 1.

SYSTEM ATICS

Class Gastropoda Cuvier, 1795

Order Vetigasgtropoda Salvini-Plawen, 1980

Family Pleurotomariidae Swainson, 1840

Genus Bayerotrochus Harasewych, 2002

Type Species: Perotrochus midas Bayer, 1965, by orig¬

inal designation.

Description: Shell large, thin, turbiniform, lacking an

umbilicus, with inflated, rounded whorls. Slit, broad, short

(<90°), forming a selenizone slightly below mid whorl.

Outer prismatic layer of shell thin, translucent, sculpture

usually limited to spiral threads or weak cords and axial

riblets that may produce weakly reticulate sculpture on

early whorls. Aperture large, ovate. Operculum small

relative to aperture.

Remarks: Bayerotrochus is easily distinguished from

Entemnotrochus , which is characterized by having a large,

conical shell with a narrow, long (>120°) slit and a deep

Table 1. Sources for the cytochrome e oxidase 1 sequence data used to produce the phylogenetic tree in Figure 28.

Page 140

THE NAUTILUS, Vol. 131, No. 2

Figures 1-10. Bayerotrochus belauensis new species. 1. Apertural, 2. lateral, 3. dorsal, 4. apical, and 5. basal views of holotype

(USNM 905395). Palau Islands, North Turtle Cove, 219 m, 26 March 2001. 6. Detail of early whorls. 7. Outer surface of damaged

operculum of holotype. 8. Apertural and 9. apical views of Paratype 1 (USNM 905397). Palau Islands, Mutremdiu 3, in 236 m, March 24,

2001. 10. Outer and inner surfaces of the operculum of Paratype 1. Scale bar = 5 cm for all shells and opercula, 1 cm for figure 6.

P. Anseeuw et ah, 2017

Page 141

Figures 11-18. Apertural and basal views of: 11-12. Bayerotrochus teramachii, SAV. of Makurazaki, Kagoshima, Japan trawled 350 m,

13-14. B. cf. teramachii, off Panglao, Bohol, Philippines, by native fishermen using tangle nets set at 100 m., 15-16. B. westralis, Rowley

Shoals, N.W. Australia, Timor Sea, by shrimp trawler in 580 m. 17-18. B. houcheti, S.E. New Caledonia, trawled in 550 m. Scale bar = 5 cm

for all specimens. All specimens in Anseeuw Collection.

Page 142

THE NAUTILUS, Vol. 131, No. 2

Figures 19-24. Radula of holotype of Bayerotrochus belauensis, new species. 19. Extended radula. 20. Rachidian, inner lateral and

outer lateral teeth. Teeth to the left of the rachidian are further anterior than their homologues to the right. 21. Sickle teeth. 22.

Transition f rom sickle teeth to filament-tipped teeth marked by onset of bristles. 23. Detail of distal ends of filament-tipped teeth near

the outer margin of the radula. 24. Paddle-shaped teeth, showing transition from filament-tipped teeth. Abbreviations: b, bristle; f,

filament: il, inner lateral teeth; ol, outer lateral teeth; p, paddle-shaped teeth; r, rachidian; s, sickle teeth.

P. Anseeuw et al., 2017

Page 143

umbilicus. It may be distinguished from Perotrochus and

Mikadotrochus by its larger, thinner, more rounded shell.

Sequence data from several genes (18S and COI) also

readily distinguish among the genera of li\ing Pleuro-

tomariidae (e.g., Harasewych, 2002: 269; Zhang et ah,

2016: 258).

Bayerotrochus helauensis new species

(Figures 1-10, 19-27)

Perotrochus africanus teramachii: Okutani and Kurata,

1998: 11, figs. 2, 3. (not of Kuroda, 1955)

Description: Shell (Figures 1-6, 8, 9) large for genus

(to 114.6 mm diameter), thin, turbiniform, with conical

spire and rounded base, non-umbilieate, consisting of 8 +

whorls. Spire coeloconoid (spire angle = 87°-99°), with

shoulder increasingly pronounced after filth whorl.

Protoconch unknown. Selenizone narrow, situated at

mid-whorl in early whorls, becoming broader,

descending to just below the shoulder by fifth whorl.

Axial sculpture of prosoeyrt riblets on early whorls

(Figure 6) that form beads at intersections with spiral

cords above and below selenizone. Ribs, cords, and beads

become reduced above the selenizone by fourth

teleoconch whorl. Spiral cords below selenizone remain

more pronounced, with 14-16 weak cords persisting onto

the last whorl between the selenizone and shell periphery.

Selenizone with opisthocyrt ribs on early whorls that

transform to spiral cords by fourth whorl and subsequently

decrease in prominence. Selenizone on last whorl with

coarse, prosoeyrt growth striae. Slit broad (to 7.5 mm),

short (75°-79° between limit of suture and rear of slit).

Suture adpressed. Basal disc strongly and evenly convex,

with sigmoidal growth striae more pronounced than weak

spiral threads (Figure 5).

Aperture broadly elliptical (w/h = 1.3-1 .5), deflected

from coiling axis by ~81°-85°. Outer lip thin, smooth,

portion below slit offset from portion above slit by

36°-51°. Columella spirally coiled, with sigmoidal flexure

near the adapical margin. Aperture nacreous, columella

porcellaneous. Base color of prismatic layer of shell ex¬

terior white, with broad, irregular axial bands of light-

orange tan and narrower bands of darker reddish -orange.

Shell may appear lustrous to varying degrees, depending

on thickness / translucence of the prismatic layer.

Operculum (Figure 10) amber-colored, multispiral, cor¬

neous, small (spanning 0.5 of minor axis of aperture).

Radula: Radula (Figures 19-24) of holotype

(shell= 114.6 mm maximum diameter) 97.7 mm long,

6.7 mm wide, asymmetrical, left-skewed, bifid posteriorly,

composed of 148 inverted V-shaped teeth rows. Following

Hickman’s (1984) terminology, each row consists of

a single raehidian tooth, 4 inner lateral, 30 outer lateral,

13 sickle, ~58 filament-tipped, and 8-9 paddle-shaped

teeth. Transitions between adjacent tooth types are

gradual, and may span 2-6 teeth. Tooth morphologies

are similar to those of other species of Bayerotrochus and

Perotrochus.

Type Locality: Palau Islands, Peleliu State, North

Turtle Cove, 07°05.12 N, 134°15.61 E, 219 m.

Type Material: Holotype, USNM 905395 (shell and

preserved animal) from the type locality, collected

26 March 2001 by Patrick L. Colin; Paratype 1, USNM

905397 (shell and preserved animal), Palau Islands, Koror

State, Mutremdiu 3, Uchelbeluu Reef, 07° 16.27 N,

134°31.37 E, in 236 m, collected 24 March 2001 by

Patrick L. Colin; Paratype 2 USNM 905396 (preserved

animal only), shell in the Patrick Anseeuw Collection,

Palau Islands, Koror State, Mutremdiu 3, Uchelbeluu

Reef, 07°16.27 N, 134°31.37 E, in 232 m, collected

21 March 2001 by Patrick L. Colin; Paratype 3, USNM

905393 (preserved animal only), shell in the Etpison

Museum, Palau Islands, Koror State, Mutremdiu 2,

Uchelbeluu Reef, 07°16.41 N, 134°31.43 E, 212 m,

collected 12 March 2001, by M.N. Dawson.

Etymology: Belau , the name of the Palau Islands in the

Palauan language.

Ecology: In Palau this new slit shell was seen and

collected on four out of over 60 submersible dives from

both rock anti sediment substrates at depths from

210-235 m (Figures 25-27). Submersible operations were

conducted for many hours both above and below these

depths, but no pleurotomariids were observed outside this

depth range. Both of Okutani and Kurata’s (1998) spec¬

imens were from 200 m, implying perhaps a limited depth

range for this species along the outer reef slopes of the

Palau Islands.

Downwelling light at these depths was relatively low,

although there was still sufficient light to maneuver the

submersible around large objects without additional

lighting. The water is extremely clear. Temperatures at

200-250 m depth along Palau’s outer reefs are generally

about 10-13°C.

Overall the outer island slopes in Palau are steep, av¬

eraging about 30°-45°. The geomorphology of the outer

slope has vertical escarpments, steep rocky slopes and

more gentle sediment slopes. No slit shells were seen on

any vertical to very steep slopes, although such profiles

occurred in the species’ depth range. Where seen, ob¬

servations of slit shells were on both sediment and hard

bottoms with slopes from about 25° to 45”.

The new slit shell was observed once apparently

feeding on the sea pen Anthoptilum grandiflorum at

212 m depth during the day (Figure 25). The snail was

on the downslope side of the sea pen with its foot ex¬

tended and wrapped around the exposed base of the sea

pen. Potentially it was digesting tissue around the

central pen; the same sea pen has an area slightly above

that shows damage, perhaps due to earlier feeding by

the mollusk. These fleshy whip-like sea pens occurred

only on sediment bottoms in clusters of individuals,

their basal bulb buried in the gentlv sloping bottom.

Page 144

THE NAUTILUS, Vol. 131, No. 2

Figures 25-27. Bayerotrochus belauensis, new species, in

situ photographs. 25. Paratype 3, station Mutremdiu 2, in 212 m.

26. Paratype 2, station Mutremdiu 3, in 232 m. 27. Holotvpe,

station North Turtle Cove, in 220 in.

This sea pen was only observed at depths between 190

and 240m, coinciding with the observed range of the

snail.

Comparative Remarks: Morphological similarities

among many of the Indo-Pacific species of Bayerotrochus

have led to the reluctance of some researchers to

distinguish populations from throughout the broad

range of the genus as separate species (e.g., Abbott

and Dance, 1982; Okutani and Goto, 1984; Okutani and

Kurata, 1998; Williams and Ozawa, 2006). More recent

studies, some based on larger sample sizes and

supplemented with molecular data, have led to the

recognition of ten Indo-Pacific species within Bayerotrochus

(Bouehet, 2016).

Among its geographically proximal congeners, Bayer¬

otrochus belauensis new species more closely resembles

B. teramachii, B. westralis, and B. boucheti than the

smaller species B. poppei, B. philpoppei, and B. delicatus,

which can be readily distinguished on the basis of size,

spire profile, and surface sculpture. This new species

is conchologically most similar to the widespread B.

teramachii , which ranges from Honshu, Japan through the

East and South China Seas to the Sulu Sea (Anseeuw &

Goto, 1996), and particularly to specimens attributed to

this species from Panglao, Bohol, Philippines.

Japanese specimens of B. teramachii (Figures 11-12)

are similar to B. belauensis new species in adult size, but

have a less stepped shell profile, thicker and heavier shells

that are less lustrous, have more pronounced spiral ribs

on the base, and weaker axial growth lines on the apical

whorls. Specimens from Panglao that have been pro¬

visionally identified as B. cf. teramachii (Figures 13—14)

more closely resemble B. belauensis new species than

Japanese B. teramachii in spire angle and profile, color and

luster of the shell surface and aperture shape. Additional

research will be required to more precisely determine the

relationships of this population from Panglao.

Bayerotrochus westralis (Figures 15-16) resembles B.

belauensis new species in adult size and in having a thin,

light shell with a short slit and similar sculpture on its early

whorls. However, B. belauensis new species differs in

having a narrower, more stepped spire, a smoother, more

lustrous surface and a broader selenizone. Bayerotrochus

boucheti (Figures 17-18) is also comparable to B.

belauensis new species in size, but differs in having a more

heavily pigmented shell with a more conical, less stepped

spire, a more depressed aperture shape, and in having

more pronounced spiral sculpture on the basal disk.

Molecular Analysis: Partial sequences of the cyto¬

chrome c oxidase I gene spanning 658 bp were obtained

from the holotvpe and two paratypes of B. belauensis new

species, and a maximum likelihood tree (Figure 28) was

produced that included the samples itemized in Table 1.

This analysis resulted in a single, fully resolved tree that

recovered the monophyly of Bayerotrochus, segregated

the Atlantic from the Pacific clades, and differentiated

the species B. teramachii , B. delicatus , B. boucheti, and

P. Anseeuw et al., 2017

Page 145

Calliostoma torrei DQ314293

Lischkeia imperialis KY426958

86

100

o 10

0 05

Entemnotrochus rumphii L7891 1

Entemnotrochus adansonianus KY435216

— Entemnotrochus adansonianus KY435217

100

99

Entemnotrochus adansonianus KY435215

— Perotrochus quoyanus KY432518

— Perotrochus amabilis KY432519

- Perotrochus gotoi KY432520

97

94

99

c,

Perotrochus caledonicus KR087190

100

97

89

Perotrochus deforgesi KR087208

Mikadotrochus beyrichii EU530109

Mikadotrochus beyrichii AM049331

6. midas KY432521

6. midas KY432523

B. midas KY432524

100

98

74^

97 f 8. teramachu AM049331

8. teramachii KY432524

r- 8. dehcatus KU 759008

r 8. boucheti KU693174

B. boucheti KU693173

8. boucheti KY432525

- 8. beiauensis n. sp. KY432528

gg j B. beiauensis n. sp. KY432526

100 ' B beiauensis n. sp. KY432527

Figure 28. Maximum likelihood (RAxML) analysis of phylogenetic relationships among pleurotomariid taxa based on partial COl

sequences. Numbers adjacent to nodes indicate bootstrap values > 50.

DO

CD

O

=r

e:

Co

B. beiauensis new species, all with high levels of support.

These data indicate that B. beiauensis new species is more

closely related to B. boucheti and B. delicatus than to B.

teramachii.

DISCUSSION

Based on morphological criteria, specimens of Bayero-

trochus collected off Palau had been identified as being

nonspecific with the Japanese species, B. teramachii

(Okutani and Kurata, 1998). Molecular data from

a specimen from this population (USNM 905395, as

Bayerotrochus sp.) was used in a more recent study de¬

scribing a new species of Bayerotrochus from Yap (Zhang

et al., 2016) and revealed that the Palau specimens were

not closely related to B. teramachii and likely represented

a new species. Sequence data from additional Palau

specimens as well as from several other Indo-Pacific

Bayerotrochus confirm that this population from Palau

represents a distinct species that is more closely related to

B. boucheti and B. delicatus that any of these species are to

Japanese B. teramachii.

Specimens from a population of Bayerotrochus from

Panglao (Figures 13-14) that were previously identified as

B. ef. teramachii, are morphologically very similar to, and

possibly conspecific with the Palau species described here.

Additional sampling and molecular studies wall be needed

to determine the relationships of this population and

perhaps revise the range of B. teramachii.

Transects along the tropical western Atlantic conducted

using submersibles revealed that pleurotomariids spe¬

cies and genera are not sympatric, but inhabit well de¬

marcated, non-overlapping bathymetric zones, each

corresponding to a specific bottom topology and habitat

(Harasewych, 2002: fig. 12). Entemnotrochus species are

the shallowest and those of Bayerotrochus the deepest

dwelling taxa along any transect. Similar data is not yet

available for Indo-Pacific species. Although the specimens

of Bayerotrochus beiauensis new species occur at shal¬

lower depths than their western Atlantic and most Indo-

Pacific congeners (Harasewych, 2002: fig. 12A), their

Page 146

THE NAUTILUS, Vol. 131, No. 2

habitat, including inclination of the slope and the sub¬

strate, appear similar.

Th ree of the four individuals collected had most of the

shell covered with an unidentified zoanthid, Epizoanthus

sp. (Figures 8, 9, 26, 27). The specimen observed feeding

on the sea pen lacked zoanthids. Anseeuw and Goto

(1996) also report the growth of zoanthids on shells of

some species, but attribute no benefit to this relationship,

describing it as parasitic growth. Zoanthids are known to

possess some of the most potent toxins found in marine

organisms and it is likely the covering of toxic zoanthids

may provide some protection from predation. Large

fishes may be deterred from ingesting whole or crushing

the shell by the living sheath of zoanthids. Despite having

had an overgrowth of zoanthids (Figure 27), the holotype

of B. belauensis new species (Figures 1-6) shows evi¬

dence of numerous repaired breaks indicating multiple

episodes of unsuccessful predation by shell-breaking

predators, possibly large crustaceans. Larger sample

sizes would help to clarify potential correlations between

zoanthid overgrowth and incidence of unsuccessful

predation. Bayerotrochus belauensis was the largest

gastropod found during more than 60 submersible dives

off Palau.

ACKNOWLEDGMENTS

Specimens were collected by the Coral Reef Research

Foundation through contract number N02-CM-77249

with the US National Cancer Institute. Dr David

Newman, NCI Project Officer, is gratefully acknowledged

for facilitating the submersible project. Patrick L. Colin,

Michael Dawson and Laura Martin were integral to the

success of the collections. The Palau Bureau of Marine

Resources and Koror and Peleliu state governments are

thanked for permission to operate the submersible and

carry out specimen collections. The assistance of Yolanda

Villacampa in producing the Scanning Electron Micro¬

graphs, and of Robin Turner in obtaining the sequence

data, is gratefully acknowledged. We thank Dr. James

Reimer for his identification of the zoanthids based on

specimens taken from the dorsal surfaces of paratypes 1

and 2.

LITERATURE CITED

Abbott, R.T. and S.P. Dance. 1982. Compendium of Seashells.

E.P. Dutton, Inc. New York, 410 pp.

Anseeuw, P. 2016. Two new pleurotomariid subspecies from the

South Pacific (Gastropoda: Pleurotomariidae). Visaya 4:

43-57.

Anseeuw, P. and Y. Goto. 1996. The Living Pleurotomariidae.

Elle Scientific Publications, Osaka, 202 pp.

Anseeuw, P., N. Puillandre, J. Utge, and P. Bouchet. 2015.

Perotrochus caledonicus (Gastropoda: Pleurotomariidae)

revisited: descriptions of new species from the South-West

Pacific. European journal of Taxonomy 134: 1-23.

Bouchet, P. 2016. Pleurotomariidae Swainson, 1840. In:

MolluscaBase (2016). Accessed through: World Register

ol Marine Species at http://www. marinespecles.org/aphia.

php?p=taxdetails&id= 196325 (accessed 11 January 2017).

Folmer. O., M. Black, W. Hoeh, R. Lutz, and R. Vrrijenhoek.

1994. DNA primers for amplification of mitochondrial

cytochrome c oxidase subunit I from diverse metazoan

invertebrates. Molecular Marine Biology and Biotechnology

3: 294-299.

Harasewych, M.G., S.P. Adamkewicz, J.A. Blake, D. Saudek,

T. Spriggs, and C.J. Bult. 1997. Phylogeny and relationships

ol pleutoromariid gastropods (Mollusca: Gastropoda): an

assessment based on partial 18S rDNA and cytochrome

oxidase I Sequences. Molecular Marine Biology and

Biotechnology 6: 1-20.

Harasewych, M G. and G.R. Sedberry. 2006. Rediscover)',

Range Extension, and Redescription of Calliostoma torrei

Clench and Aguayo, 1940 (Gastropoda: Vetigastropoda:

Trochidae). The Nautilus 120: 39—44.

Hickman, C.S. 1984. Form and function of the radulae of

pleurotomariid gastropods. The Veliger 27: 29-36.

Kearse, M., R. Moir, A. Wilson, S. Stone-Havas, M. Cheung, S.

Sturrock, S. Buxton, A. Cooper, S. Markowitz, C. Duran, T.

Thierer, B. Ashton, P. Mentjies, and A. Drummond. 2012.

Geneious Basic: An integrated and extendable desktop

software platform for the organization and analysis of se¬

quence data. Bioinformatics 28: 1647-1649.

Okutani, T. and Y. Goto, 1998. Perotrochus africanus teramachii

from the Andaman Sea, a new locality. Chiribotan 59:

55-56.

Okutani, T. and Y. Kurata. 1998. An unusual molluscan as¬

semblage containing Perotrochus africanus teramachii on

the insular shelf of Palau Islands. Venus 57: 1 1-16.

Williams, S.T. and T. Ozawa. 2006. Molecular phylogeny sug¬

gests polyphyly of both turban shells (family Turbinidae)

and the superfamily Trochoidea (Mollusca: Vetigas¬

tropoda). Molecular Phylogenetics and Evolution 39:

33-51.

Williams, S.T., S. Karube, and T. Ozawa. 2008. Molecular

systematics of Vetigastropoda: Trochidae, Turbinidae and

Trochoidea redefined. Zoologica Seripta 37: 483-506.

Zhang, S.P., S.Q. Zhang and P. Wei. 2016. Bayerotrochus

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tomariidae). Zootaxa 4161: 252-260.

THE NAUTILUS 131(2): 147-149, 2017

Page 147

A redescription of Rissoina mayori Dali, 1927, a junior

subjective synonym of Opalia pumilio (Morch, 1875)

(Gastropoda: Epitoniidae)

Leonard G. Brown

5 Vuinbaco Dr.

Wallingford, CT 06492 USA

[email protected]

Bruce D. Neville

2700 Sandy Circle

College Station, TX 77845-5309 USA

[email protected]

ABSTRACT

An examination of the type material of Rissoina mayori Dali,

1927, confirmed that Ponder (1983) was correct in allocating the

species to the family Epitoniidae. It is here redescribed and

a lectotype is designated. Because of the similarity in shell

characters and the overlap in geographic range, we consider R

mayori and Opalia pumilio (Morch, 1875) to be synonyms.

Additional Keywords: New synonym, western Atlantic, Rissoidae

INTRODUCTION

In 2015, Marien Faber, who has been researching the

Rissoininae, contacted us regarding Rissoina mayori Dali,

1927. He was looking for our opinion on whether tins

species was referable to the Rissoininae as proposed by Dali,

or to the Epitoniidae, as proposed by Ponder (1983: 90).

Neither Dali nor Ponder provided figures of the spe¬

cies, nor, apparently, has it been critically discussed or

figured elsewhere in the literature. To resolve this

question, we contacted Yolanda Villacampa and Dr. Ellen

Strong at the Smithsonian Institution’s National Museum

of Natural History (USNM), requesting photographs of

material of this species in the USNM type collection. They

provided the scanning electron microscope (SEM) pho¬

tographs reproduced here.

After reviewing the SEM photographs and the type

material, it became apparent that Ponder was correct in

his placement. This species indeed seems to belong to the

Epitoniidae. Because this species has, to date, been

overlooked in reviews of western Atlantic epitoniids, we

herein redescribe and illustrate this species.

SYSTEMATICS

Family Epitoniidae S. S. Berry, 1910

Genus Opalia H. and A. Adams, 1853

Type Species: Scalaria australis Lamarck, 1822, type

of Clathrus Gray, 1842.

Opalia pumilio (Morch, 1875)

Scala (Opalia) subvaricosa var. pumilio Morch, 1875: 268

Rissoina mayori Dali, 1927 new synonym (Figures 1,

2, 3)

Description of Rissoina mayori: Lectotype (Figures

1, 2, 3) 4.2 mm in length (apex missing), 1.8 mm in

diameter, imperforate and sculptured. Color white.

Five+ convex teleoconch whorls; suture strongly

crenulated, aperture obliquely ovate, outer lip

thickened. Costae strong on the early whorls, but

weaker on the rounded base of the body whorl; 15

costae on the body whorl, including one that is

thickened and varix-like. Intervals with minute spiral

striae, separated by rows of punctae. Operculum

unknown.

Type Material: Lectotype of Rissoina mayori Dali,

1927, USNM 108371, here designated.

Type Locality: Off Georgia, 30°44’ N, 79°26" W, 440

fms [= 805 m], Albatross station 2415 (Dali 1927: 1, 105).

Remarks: Dali (1927: 105) referred to Rissoina

mayori one specimen from Station 2415 which is due

east of Cumberland Island, Georgia, as well as material

collected off Miami in 58 fathoms. There is no in¬

dication on the label that Dali designated the specimen

collected from Station 2415 to be the holotype. The

material collected off Miami was not located in the

USNM collection. (Y. Villacampa, personal comment,

15 April 2015). Therefore, we consider these specimens

to be syntypes and are designating USNM 108371 to be

the lectotype.

Because Rissoina mayori has crenulated sutures and

a pitted intritacalx, we are referring this species to the

genus Opalia. This is consistent with Bouehet and

Waren (1986: 54), who referred to the genera Gre-

goroioi.scala Cossmann, 1912, and Punctiscala de

Page 148

THE NAUTILUS, Vol. 131, No. 2

Figures 1-6. Opalia species. 1-3. Rissoina maijori , leetotype USNM 108371, 105 miles off Georgia, 30°44’ N, 79°26’ W, 805 m,

Albatross station 2415. 4-6. Scala (Dentiscala) hellenica nodosocarinata Dali, 1889 [= Opalia pumilio (Morch, 1875)], holotype

USNM 82955, 5 miles off Cape Florida, Florida, 15 m, 3.9 mm.

Boury, 1890, species with non-crenulated sutures and

a strong basal disk over which the costae do not extend

and who referred to the genus Opalia species with

crenulated sutures that may or may not have a basal

disk.

Based on the material at hand, Rissoina mayori

appears to be indistinguishable from Opalia pumilio

(Morch, 1875). The shell characters cited in Clench

and Turner (1950: 238) for O. pumilio are certainly

applicable to Dali’s species. While the holotype of

Scala nodosocarinata Dali, 1889 (a synonym of O.

pumilio [Clench and Turner, 1950: 237]) (Figures 4, 5,

6) appears to differ from R. mayori in having finer

pitting and raised ridges on the teleoeoneh whorls, it is

not clear that these differences are taxonomically

significant, given the fact that O. pumilio is “...an

exceedingly variable species” (Clench and Turner,

1950: 238). Furthermore, the known range of R.

mayori, from Cumberland Is., Georgia, south to Mi¬

ami, Florida, is well within the range of O. pumilio. In

fact, the type locality for Scala ( Dentiscala ) nodoso¬

carinata Dali, 1889, is 5 miles off Cape Florida,

Florida, a locality that could be considered to be

Miami, Florida.

ACKNOWLEDGMENTS

We want to thank Marien Faber for bringing this prob¬

lematic species to our attention. We also want to thank

Dr. Ellen Strong and Ms. Yolanda Villacampa at the

USNM for providing us with the photographs of the type

material for R. mayori and S. hellenica nodosocarinata

and for allowing us to examine the type material of these

two species. We wish to thank Patrick Zinn at Texas A&M

University for creating the plate.

EG. Brown and B.D. Neville, 2017

Page 149

REFERENCES

Adams, H. and A. Adams. 1853, in 1853-1858. The genera of

Recent Mollusca, arranged according to their organization.

London: Van Voorst.

Bouchet, P. and A. Waren. 1986. Revision of the northeast

Atlantic bathyal and abyssal Aclididae, Eulimidae, Epito-

niidae (Mollusca, Gastropoda). Bollettino Malacologico,

supp. 2: 300-576.

Clench, W.J. and R.D. Turner. 1950. The genera Sthenonjtis,

Cirsotrema, Acirsa, Opalia and Amaea in the Western

Atlantic. Johnsonia 2: 221-246.

Dali, W.H. 1927. Small shells from dredgings off the southeast

coast of the United States by the United States Fisheries

Steamer "Albatross” in 1885 and 1886. Proceedings of the

United States National Museum 70(18): 1-134.

Dali, W.H. 1889. Reports on the results of dredging, under the

supervision of Alexander Agassiz, in the Gulf of Mexico

(1877-78) and in the Caribbean Sea (1879-80), by the U.S.

Coast Survey steamer “Blake,” Lieut. -Commander C. D.

Sigsbee, U. S. N., and Commander J. R. Bartlett, U. S. N.,

commanding. XXIX. Report on the Mollusca. Part II.

Gastropoda and Seaphopoda. Bulletin of the Museum of

Comparative Zoology at Harvard College 18: 1-492, pis.

410-440.

Morch, O.A.L. 1875. Synopsis familiae Scalidarum Indiarum

occidentalium = Oversigt over Vestindiens Sealarier.

Videnskabelige Meddelelser Ira Naturhistorisk Forening i

Kjobenhavn, for 1874: 250-268.

Ponder, W.F. 1983. Reclassification of some American species

assigned to the Rissoidae (sensu Into). The Nautilus 97:

90-91.

THE NAUTILUS 131(2):150, 2017

Page 150

Erratum

The recent article by Miyajima et al. (2017) was published with an earlier, incorrect version of Figure 55. The correct

version of that illustration is included here. The new version includes the corrected species name Adulomya sp. 1 in the

figure legend, replacing the originally submitted name Adulomya sp. A.

Figure 55. Relationships between shell length and height with growth of Adulomya sp. 1 from Loc. 2, A. uchimuraensis Kuroda,

and A. chitanii Kanehara. Measurements of A. uchimuraensis and A. chitanii are from Kanno et al. (1998) and Amano and Kiel (2011),

respectively.

LITERATURE CITED

Miyajima, Y., T. Nobuhara, and 11. Koike. 2017. Taxonomic reexamination of three vesicomyid species (Bivalvia) from the middle

Miocene Bessho Formation in Nagano Prefecture, Central Japan, with notes on vesicomyid diversity. The Nautilus 131: 51-66.