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CONTENTS

M.G. Harasewych

Manuel J. Tenorio

Bianca Campagnari

Daniel L. Geiger

Richard L. Squires

Daniel Abbate

Luiz Ricardo L. Simone

Daniel C. Cavallari

Shugian Zhang

Peng Wei

Kazutaka Amano

Krzysztof Hryniewicz

Robert G. Jenkins

Omar Mejia

Benjamin Lopez

José Ma. Reyes-Gomez

Author Index ..........................

Meo riliuUs

Volume 132, Numbers 3-4

December 21, 2018

ISSN 0028-1344

The genus Cerion (Gastropoda: Pulmonata: Cerionidae) on San Salvador

[Watling Island], Bahamas: A geometric morphometric analysis of shell

TAA ONT] OL OK CH COVEY tsb acn eR heat Ao EUROS. SOR ba OE Rae RED En rr ee al

How many micromollusks are there? A case study on species richness in

Hawai'i, with the description of a new species of Murdochella (Gastropoda:

IOULOOTIICEYS)) sancneretnauhaceernnadetee ae aannene cae RE senet Netee cieheicl Ee cect ochre ato etecateceecere eae §3

Catalog of the taxonomic updates of northeastern Pacific Late Cretaceous

shallow-marine bivalves and gastropods named from 1874 to 1966 ............0.... 9]

Anatomy of Engoniophos unicinctus from Isla Margarita, Venezuela

(Gastropoda: Caenogastropoda: Nassariidae), with a discussion on the

lowmrcctinioimacsavaticl 1ElEVSIOINGLOTS) .0..2.ssqcoscenneaceeosecvossosncdx0s90aqJo00900s00Ke0e7en0G0badBENEAODDHOG 10]

Mericella zhangsupingae, a new cancellariid species from the South China

Sen (Castropocles CancalletttC®)) ccosasadonscoscenocobancscdadabéodeasodandadeanoncbosesacadosacbHGHORE 113

A newly discovered Paleocene species of Boreocomitas (Gastropoda:

Pseudomelatomidae) from eastern Hokkaido, Japan, with implications for

thesbiogeourapinyazolstnesealeocencyb eninge Stal tems eee sean en 117

Three new species of the genus Humboldtiana (Gastropoda: Pulmonata:

Helio @ | Gltsirrni Glave) efi @ Tn x © eee 124

CEC 28 7018

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THE NAUTILUS 132(3-4):71-82, 2018

Page 7]

The genus Cerion (Gastropoda: Pulmonata: Cerionidae) on San

Salvador [Watling Island],

analysis of shell morphology

M. G. Harasewych

Department of Invertebrate Zoology, MRC-163

National Museum of Natural History

Smithsonian Institution

PO Box 37012

Washington, D.C. 20013-7012, USA

Bahamas: A geometric morphometric

Manuel J. Tenorio

Departamento de CMIM y Quimica Inorganica-IN BIO

Facultad de Ciencias, Torre Norte, 1“ Planta

Universidad de

11510 Puerto Real

Cadiz, SPAIN

Cadiz

ABSTRACT

Morphometric analyses of shell shape of living specimens of

Cerion inhabiting San Salvador Island segregate samples into

two primary phenotypes, one inhabiting the westem and

southem coasts of the island, the other the eastern and much of

the northem coast. These are concordant with phenotypes

identified in prior morphometric studies. Lectotypes are

designated for Cerion watlingense Dall, 1905; C. incon-

spicuum Dall, 1905; C. inconspicuum lacunorum Dall,

1905; and C. coloni Bartsch, 1924. The lectotype of

Cerion watlingense Dall, 1905 falls within the western and

southern phenotype, and is the oldest name available for this

taxon. The lectotype of Cerion coloni Bartsch, 1924, a validly

introduced, but previously unrecognized taxon, and the holotype

of Cerion rodrigoi Gould, 1997 both fall within the group

containing the east coast populations, with Cerion coloni

Bartsch, 1924 being the oldest available name for this pheno-

type. A third, previously unrecognized phenotype, represented

by a single inland population, is morphologically similar to the

lectotype of Cerion inconspicuum Dall, 1905, which is the oldest

available name for this phenotype. The geographical distribution

and chronological succession of these phenotypes since the late

Pleistocene is reviewed in the context of both the single and

multiple colonization models for the arrival of Cerion on San

Salvador, and the evolutionary and taxonomic corollaries of each

model are discussed.

INTRODUCTION

Nearly all modern papers dealing with land snails of the

genus Cerion are prefaced by an acknowledgment of

ihe extreme morphological diversity among the many,

small populations of these snails. This is usually followed

by a lamentation (e.g., Woodruff, 1978) about the pro-

fusion of names |ca. 600] that have be en applied to the

various phenotypes, mostly in the late 19"? and early 20"

centuries. Nearly all of these named taxa are allopatric,

but many interbreed freely when brought into contact

(see e.g., Bartsch, 1920; Woodruff and Gould, 1987).

Throughout the range of Cerion (from the Keys and

barrier islands of tropical Florida, throughout the Baha-

mas, Greater Antilles, Cayman Islands, western Virgin

Islands, and the Dutch Antilles, but absent from Jamaica,

the Lesser Antilles, and coastal Central and South

America), there are but three reported cases in which two

different phenotypes co-occur sympatrically, one in Cuba

(Mayr, 1963), and two on Great Inagua (Gould and

Woodruff, 1990: Goodfriend and Gonitel 1996).

Earliest records of Cerionidae date from the Creta-

ceous of Montana (Roth and Hartman, 1998), yet the

overwhelming majority of taxa (>98%) are from late

Pleistocene, ieislbcene. and especially Recent faunas,

particularly those of Cuba (29%) and the Great Bahama

Bank (51%) (Har asewych, 2012: Table 1). Changes in sea

levels brought about by Pleistocene glaciations would

have joined populations of the numerous adjacent islands

on shallow banks such as the Great Bahama Bank during

glacial periods and isolated them during interglacial sea

level rises, facilitating hybridization and amplifying

morphological diversity over multiple eustatic cycles

(Harasewych, 2012).

Although the distribution of cerionids throughout most

of the islands of the tropical western Atlantic was the

result of overland dispersal followed by the break-up of

GAARlandia (Greater Antilles + Aves Ridge Land

Bridge) and the displacement of Antillean leaks to the

Ronheast by the advancing Caribbean Tectonic Plate

(Itturalde-Vinent, 2006: fig. 13), the lack of a land con-

nection between the islands of the Bahamas and the

Greater Antilles precluded overland colonization. Rather,

the Bahamian Islands must have been populated by

propagules dispersed from the Greater Antilles either by

rafting or by hurricanes (Harasewych, 2012: 124). Cle neh

(1 938: 495) reported that the fauna of the Bahamas

archipelago likely dates only from the Pleistocene and

has reached the Bahamas by fortuitous means, noting

(Clench, 1938: 484485) that the species of the Great

THE NAUTILUS, Vol. 132, No. 3-4

Page 72

Table 1

Pe ypulation STATION LOCATION LATITUDE

l BP Barker’s Point 24°06.61 N

2» VH Victoria Hill 24°05.83’ N

3 EA E of airport 24°03.85' N

4 Cr N of Cockburm Town 94°03.24 N

5 SU Sugarloaf 24°00.43° N

6 G1234 Sugarloaf 24°00.15° N

7 GB Grotto Bay 23°57.26° N

8 WOQ Watling’s Quarry 23°57.23' N

9 SH1 W of Sandy Hook 23°56.85° N

10 SH2 E of Sandy Hook BST NTT IN

1] rar The Thumb 24°00.S0° N

12 G193b W of Storr’s Lake 24°02.72’ N

13 G194 W of Storrs Lake 24°03.52’ N

14 @1225) Columbus Monument 24°04.06 N

15 G1240 NE of Granny Lake 24°02.89° N

16 CC Crab Cay 24°03.93° N

17 HB Hanna Bay 24°O7.14 N

18 MH Man Head Cay 924°07.54 N

19 NPI North Point 94°07.65° N

20) NP2 North Point 24°07.22’ N

21 GC Green Island (Cay) 24°08.33’ N

Bahama Bank were closely allied to those of Cuba, while

the species of the easterly islands (Crooked Island Group

to Caicos Island) were affiliated with species from Haiti.

He hypothesized that relationships among the terrestrial

molluscan faunas of this region are due to island proximity

and the paths of hurricanes.

San Salvador is a small island on the eastern margin

of the Bahamas platform that has remained isolated

by deep water, with little change in island area during

the eustatic sea level fluctuations since at least the

Pliocene. Its living and fossil Cerion fauna had been

spared some of the excesses of early taxonomists, yet has

been well studied by modern researchers, particularly

in the areas of ecology and chronostratigraphy, due

largely to the presence of the Gerace Research Centre

(formerly Bahamian Field Station) on the island. The

genus Cerion is well represented in the fossil record

aft San Salvador, with a documented dated presence

spanning the past 140,000 years (Hearty and Schellenberg,

2008).

In the present study, we analyze the morphology of

Cerion shells from representative populations spanning

the island of San Salvador in order to determine the

number of phenotypes and their distribution, and to

reconcile these with the type specimens of all taxa de-

scribed from San Salvador or Watling Island. A second

series of analyses incorporates measurements from a

small subset of well-preserved, adult fossil specimens

for which ages have been determined (Hearty and

Schellenberg, 2008). These analyses follow the chro-

nology of patterns of phenotype ‘distribution from the

Pleistocene to the Recent to the extent possible given

the sampling.

Conclusions of earlier works on the Cerion of San

Salvador are reviewed in the context of our findings.

Populations of Recent of Cerion from San Salvador (Watlings Island) included in this study.

LONGITUDE CATALOG NUMBER N GROUP

74°30.87 W USNM 1110076 10

74°31.27 W USNM 1110077 10

74°30.90° W MCZ IP 190243-190252 10

74°32.16 W USNM 590160 10

74°31.89° W USNM 1110078 8

74°32.17 W MCZ IP-190212-190221 10

74°33.63° W USNM 1110079 10

74°32.87 W USNM 1110080 10

74°29.91° W USNM 1110081 10

74°29 32) W USNM 1110082 10

74°27 .35° W USNM 1110088 10

74°27 17 W MCZ IP 190195-190204 10

74°27.19° W

74°25.80° W

MCZ IP 190205-190211 ff

MCZ IP 190264-190273 9

[NS) [82) (82) [RS) [N2) [8S) [R2) [8S) [8S) [8S) [SST Ps fe ed) Pt tt (8) 1)

74°27.19 W MCZ IP-190222-190231 10

74°25.80' W USNM 1110087 )

74°27.03° W USNM 1110085 4

74°26.97 W USNM 1110086 §

74°27.44 W USNM 1110083 8

14°27.37 W USNM 1110084 10

74°30.51° W USNM 359519 10

TAXONOMIC HISTORY

Known as Guanahani to the native Lucayan inhabitants,

the island was named San Salvador by Christopher Co-

lumbus when he first sighted land in 1492. It was settled

by John Watling in the i 7th century and became known as

Watlings Island. The name San Salvador was officially

transferred from what is now Cat Island to Watlings Island

in 1925 as historians concluded that this was the island

where Columbus first landed in the New World.

Dall (1894: 117) reported on the molluscan fauna of

Watling Island based on material sent to him. He iden-

tified as Cerion (Strophiops) glans (Kiister, 1844) a single,

rather small specimen taken from about a pint [0.473 L] of

beach drift from Watling Island lagoon during a U. S. Fish

Commission sampling. Dall (1905: 438) later wrote that

no Cerion had previously been noted from Watling Island,

and that his earlier report of C. glans from this island was

due to an error.

Pilsbry (1902: 265) described the subspecies Cerion

eximium fraternum (Figure 1) as being from San Salvador,

based on material attributed to Bland. Clench (1938: 531)

reported this taxon from Little San Salvador (now also

known as Half Moon Cay), an island between Eleuthera

and Cat Cay. He noted that the exact type locality for this

taxon is unknown, but "as Bland had received other

species of Cat Island material from near the center of the

island, it is quite probable that his fraternum material

came from the same area." As Pilsbry’s publication pre-

dated the renaming of Watlings Island, it seems clear that

Cerion eximium fraternum was described from Cat Island

(called San Salvador at the time) and does not occur on the

island that is known as San Salvador today.

In his report on land snails collected in the Bahamas,

Dall (1905: 438-439) proposed two new species, Cerion

ey)

M.G. Harasewych and M.]. Tenorio, 2018 Page

Figures 1-6.

eximium fraternum Pilsbry, 1902. Lectotype (designated by Baker, 1963: 206). ANSP ber San Salvador (Bland). 2. Cerion

(Strophiops) watlingense Dall, 1905. The specimen illustrated by Dall (1905: pl. 58, fig. 7) is here designated as the eee pe

for this taxon. USNM 132970. Watling Island [San Salvador]. 3. Cerion (Strophiops) inconspicuum Dall, 1905. The specimen

illustrated by Dall (1905: pl. 58, fig. 2) is here designated as the lectotype for this taxon. USNM 37676. Hs atling Island. 4. Cerion

(Strophiops) i inconspicuum lacunorum D: all, 1905. The specimen illustrated by Dall (1905: pl. 58, fig. 4) is here designated as

the lectotype for this taxon. USNM 127494, Watling Island, on the shores of the lagoon. 5. Cerion coloni Bartsch, 1924.

Lectotype (designated herein), USNM 359377, C ‘olumbus Point (Crab C ay), San Salvador, Bahamas. 6. Cerion rodrigoi Gould,

1997. Holotype, MCZ IP 190264. Windswept terrace of Crab Cay adjacent to the Columbus Monument, San Salvador, Bahamas.

Scale bar = | cm for all images.

Ape rtural, late ral, and dors: ul views of primary type spe cimens of Cerion de NYG ribe ole as from San Salv ador. at Cerion

watlingense (Figure 2) and Cerion inconspicuum

specimens during his two weeks on the island. This

(Figure 3), both from Watling Island. He also described

published report included a photograph (Bartsch, 1924:

the \ variety Cerion inconspicuum lacunorum (Figure 4)

from the shores of the lagoon on Watling Island.

Paul Bartsch conducte d fieldwork on San Salvador in

August of 1923, noting in a report of his visit (Bartsch,

1924) that he had gathered approximately 25,000

fig. 41) of seven specimens of Cerion on a leaf and was

captioned "Cerion coloni, new species, ... taken at the base

of the Columbus Monument, San Salvador."

sociates a name with an illustration prior to 1931, it is an

available name (ICZN 2012, Article 12.2.7). The taxon

As this as-

Page 74

THE NAUTILUS, Vol. 132, No. 3-4

Cerion coloni Bartsch, 1924, (Figure 5) was not recorded

in Clench’s (1957) catalog of the Cerionidae, nor in

Ruhoff ’s (1973) listing of Bartsch’s zoological taxa. Un-

aware of Bartsch’s publication, Gould (1997) described

Cerion rodrigoi (Figure 6) from the same locality.

Many eubee quent researchers (e.g., Woodruff, 1978:

Poth et al. 2010: Rose, 1990) have regarded the Cerion

fauna of San Salvador to consist of a single variable

species, while others (e.g., Baldini et al., 2007:174; Hearty

and Schellenberg, 2008) have elected to avoid species

nomenclature, "as neither proof nor consensus exists with

regard to species level taxonomy".

MATERIALS AND METHODS

A geometric Menges analysis of shell shape was

performed on 193 specimens of Cerion snails from 21

populations spanning San Salvador Island (Table 1).

Figure 7 illustrates the location and a representative

specimen from each population.

Specimens were individually photographed in an

apertural orientation using a digital camera (Nikon D300)

with the lens axis oriented perpe oacltaulle to the coiling axis

of the shell. Images were taken at a distance of 35-40 cm

in order to minimize image distortion and parallax error.

To avoid the effects of Alllennateeay only adult specimens,

free of repaired breaks and with a well- -preserved em-

bryonic shell and a fully formed terminal adult aperture

having an expanded and reflected lip indicative of terminal

growth (¢ (Gould and Woodruff, 1978: 381), were selected

for analysis.

Shells were digitized using TPSDIG2 (Rohlf, 2005). A

total of 24 points were selected to capture shell shape

(Figures 8-9). All points were treated as landmarks. These

points cover most of the ratios used for classical charac-

terization of Cerion shell morphometry (Gould et al.,

1974). Taking several photographs of the same speci-

men ensured the reproducibility of the measurements by

checking that the deviation in landmark positioning was

minimal Thus, the same Cerion shell (maximum length

25.56 mm) was photographed 10 times, even with dif-

ferent optics in the camera. The maximum error in

landmark positioning was 0.13 mm (0.51 % referenced to

the size of the shell). The mean error was 0.06 mm (0.23 %

relative to the size of the shell). Landmarks were aligned

by a standard generalized least squares (GLS) procedure.

Procrustes coordinates and centroid sizes (CS) were

generated by COORDGENG6bh (Sheets, 2003-2005). CS

provides a measure of the geometric size of each speci-

men, and is computed as the square root of the sum of

squared distances from each landmark to the centroid of

each specimen’s configuration of landmarks (Zelditch

et al., 2004). Partial warp scores (PWS) were derived

from the 24 landmarks used for capturing

the shells upon Procrustes alignment and calculation

of the consensus reference form. PWS were used as

shape variables in our analyses. PWS and their principal

components (PC, also known as _ relative warps) were

g the shape of

computed using PCAGENS, which is included in the

Integrated Morphometrics Package software suite (IMP 8,

Sheets, 2003-2014). CS and the first principal component

(PC1) of the partial warps accounted for 92.2% variance in

the dataset. Gaussian clustering of these variables (CS and

shape PC1 scores) was used to detect population clusters,

with the number of clusters determined using the Bayesian

Information Criterion (BIC) as implemented i in MCLUST

Version 5 software package for R (Scrucca et al., 2016). This

program tests different models of clustering, and the fit of

each model to the dataset is assessed by means of the

Bayesian information criterion.

In our system the best-fit model (BIC —332.56, log L

~132.0738, df = 13, for n = 193) indicated 3 compan

or clusters of variable volume and equal shape and ori-

entation (VEE model). The pairwise Procrustes distances

between groups of populations based upon shell mor-

phometry were obtained with TWOGROUPS (Sheets,

2003-2014) using resampling methods (400 bootstraps).

The significance of the Goodall’s F-test was assessed by a

bootstrappe d F-test. Discriminant Function Analysis

(DFA) using PWS+CS as variables, and morphometric

grouping as factor was performed with the programs

STATGRAPHICS CENTURION XVII and CVAGENS

(Sheets, 2003-2014). The latter program was also used for

generating the two-dimension (2D) scatter plots and the

corresponding deformation grids and vectors. Discrimi-

nation was statistically significant along both DF1 and

DF2 axes (DF 1: Wilk’s A = 0.0852, y* = 413.8103.

df=90, p < 2.22045 x 10°'°: DF 2: Wilk’s A = 0.4458 x =

IGS, Gl = WAL ip = DBT x 10").

RESULTS

A geometric morphometric analysis of the shells of 193

specimens from 21 living populations of Cerion from San

Salvador Island was conducted. A Principal Component

Avnallysi of the PWS yielded two statistically significant

components that represent 62.4 % of the overall variation

in shape in our sample. Shell height of Cerion from San

Salvador is known to exhibit a strongly bimodal distri-

bution (Fronabarger et al., 1997). Gaussian clustering of

CS and shape PC] data was used to determine the dif-

ferent morphometric groups in our sample according to

size and shape by the program MCLUST in R. The best-fit

model indicated three components or clusters of variable

volume and equal shape and orientation (Figure 10).

This analysis indicates that the pattern of dtsucbntion of

size and shape in our sample of Cerion shells is best

represented by three separate groups, partitioned among

three different phenotypes. We obtained the initial com-

position of each of the groups directly from MCLUST,

taking into consideration that the group assignment for

specimens falling in the overlap region among clusters

may be subjected to increased uncertainty (Figure 11).

Final assignment of individuals to each of the three groups

was determined by checking the composition of each of

the populations against the suggested grouping resulting

M.G. Harasewych and M.]. Tenorio, 2018 Page

ea bank edge and lakes

fe] present shoreline

al +6 m shoreline

LENE

Dy. \ i

Figure 7. Map of San Salvador, Bahamas. Locations of sampled populations (solid circles) are identified by alphanumeric station

designations (see Table 1). A representative specimen from each station is shown.

from MCLUST Gaussian clustering. Individuals from Group 2, and thus had a higher uncertainty in group

each of the populations studied were all members of the classification.

same group with the exception of populations VH and CT, Phenotype 1 (Group 1) includes all individuals from

which fell into the region of overlap between Group | and populations: BP, VH, CT, SU, G1234, GB, WQ, SH1, and

Page 76

THE NAUTILUS, Vol. 132, No. 3-4

Figures 8-9.

coloni Bartsch, 1924, showing 9. positions of the landmarks used

in the morphometric analyses.

Position of landmarks. 8. A specimen of Cerion

SH2. Phenotype 2 (Group 2) includes all individuals from

populations: TT, G193b, G194, G1225, G1240, CC, HB,

MH, NP1, NP2, and GC. Phenotype 3 (Group 3) includes

all individuals from a single population, EA.

Phenotype 1 occurs exclusive ly along the western and

southern coasts of San Salvador to the arom of the lagoon

(Figure 7, BP to SH2). Phenotype 2 occurs along the east

coast of the island, from the mouth of the lagoon

northward to the base of the North Point Peninsula (Fig.

7, TT to NP2), as well as at several locations inland from

the east coast (Fig. 7, G193b, G194, G1240). Populations

from an isolated cay (Beane 7 7, C C) off the north coast as

well as from the tip of the North Point Peninsula

(Figure 7, NP1) include phenotypes primarily from Group

2 but ih several individuals from Group 1, indicating

areas of potential overlap or hybridization. Populations

VH and CT appear on the plot clustered in the area of

overlap of between Phenotypes | and 2. The geographic

distribution of these populations in the western coast of

the island led to their final classification as members of

Phenotype 1. Phenotype 3 is known only from a single

population inland from the west coast (Figure 7, EA).

Confirmation of the composition of these morphom-

etric groups was accomplished by means of Discriminant

Function Analy sis (DFA). Discrimination was See

significant along both DF1 and DF2 axes (Figure 12). A

DFA using the above group composition as a factor and

PWS+CS as variables classified correctly 190 out of 193

specimens (98.4 %). In the jack-knifed assignment test,

the rate was 90.7 % (175 oe out of 193). Using shape-

only variables (i.e. leaving CS out of the anelyai) the

correct classification rate was 97.4 % (89.1 % in the jack-

knifed test). Figures 13 to 24 represent the relativ e po-

sition of the specimens from each of the populations in

discriminant space.

The three morphometric groups resulting from this

analysis differ statistically in size and shape. ANOV A in-

dicates that there are significant differences in CS be-

tween Group | (mean CS = 35.79) and Group 2 (mean

CS = 43.05) (F = 311.37, p = 0.0000), and between

Group 2 and Group 3 (mean CS = 34.98) (F = 79.85, p =

0.0000). However, there is no significant difference in size

between Group | and Group 3 (F = 080; p = 0!3735):

The three groups exhibit significant differences in mean

shell shape, as inferred from the calculated pairwise Pro-

crustes distances between means and the corresponding

results of bootstrapped Goodall’s F-tests (Table 2).

Thus, individuals from groups 1 and 3 do not differ in

shell size, but differ greatly in shell shape. The dis-

tinguishing features of each of the groups can be extracted

fen the shape deformation implied by DF1 and DF2

(Figure 12). Group | is separated from Group 2 along the

DF] axis, while € Group 3 is separated from group 1 anal D 2

along the DF 2 axis. Members of Group 2 can be separated

from Group | by the larger size of their shells, which are

more elongated with a projecting apex and a relatively

narrower aperture. Group I is Eiamcrereed by smaller

shells, with broadest whorls at mid-length, and a more

depressed apex, giving a general oval appearance and with

a relatively wider aperture. Members of Group 3 exhibit

shell features similar to those of Group 1. However,

individuals in Group 3 display a less inflated shell shape,

with parallel sides slightly narrowing at the mid-length

and with a convex profile towards the apex. She Ils of

individuals from Group 1 are relatively broader at the

center, with a straight profile towards the more pro-

truded apex. The aperture in Group 3 is narrower than in

Group 1. Apart from these differences in size and shape,

the shells of individuals in groups 1 and 3 also exhibit a

more developed sculpture of axial ribs compared to

members of group 2. In the latter, the axial ribs may be

reduced or even anes nt in certain specimens. This fea-

ture provides additional support for the assignment of

populations VH and CT to Group 1, given their strong

axial ribbing.

Using this morphometric framework, primary type

specimens of each of the named taxa from San Salvador

(Lectotypes de ssignate od in captions to Figure s 2-5) were

digitized and analy zed by means of DFA (Figure 25). The

lectotype of Cerion watlingense clearly falls within

Group 1 (probability 98. 8%), and is the oldest name

available for this phenotype. The lectotype of Cerion

coloni and the holotype of C. rodrigoi both fall within

Table 2. Comparison of pairwise Procrustes distances be-

tween groups of specimens as defined by shell phenotype, based

upon shell morphometry. The significance of the Goodall’s

F-test was assessed by a bootstrapped F-test (400 replications).

Group Group Procrustes distance | Goodall’s-F )

I i

] ») 0.0361 30.03 0.0025

1 3 0.0823 31.25 0.0025

2 3 0.0577 16.26 0.0025

M.G. Harasewych and M.]. Tenorio, 2018 Page 77

e =Group1

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Figures 10-12. Plots of the first principal component (PC1) versus Centroid size (CS). 10. Symbols indicate classification corresponding

to the best-fit model as determined by MCLUST. Component means are marked, and ellipses with axes are drawn corresponding to their

covariances. 11. Classification uncertainty as determined by MCLUST. Symbol size proportional to degree of uncertainty. 12. Discriminant

Function Analysis (DFA) of the shapes and sizes of the shells of 193 specimens corresponding to 21 populations of Cerion snails from San

Salvador Island, using Partial warp scores (PWS) + Centroid size (CS) as variables and considering morphometric group as classification

factor. The 2D scatter plot of the DF1 and DF2, and the corresponding deformation grids and vectors are shown.

Group 2 (probabilities > 99.9%), with C. coloni being the Group | and Group 3 with the algorithm assigning them

oldest available name for this phenotype. The lectotypes to Group 3 at 95 % probability for C. inconspicuum, and

of Cerion inconspicuum and Cerion inconspicuum lacu- 51.4 % for C. inconspicuum lacunorum. Cerion incon-

norum have morphologies that are intermediate between spicuum is the oldest available name for this phenotype.

Page 78

THE NAUTILUS, Vol. 132, No. 3-4

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Figures 13-24. Positions of specimens from individual living (Figures 13-23) and fossil (Figures 22-24) populations in the dis-

criminant space.

A number of well-preserved fossil specimens that had

been dated by amino acid racemization in a previous study

(Hearty and Schellenberg, 2008) were digitized, scored,

and classified by DFA as had been done with the type

material. The oldest well-preserved specimens suitable

for morphometric analyses were from French Bay (be-

tween WQ and SHI on Fig. 7). All three specimens from

SFB4b(1) (Figure 26) (Hearty and Schellenberg, 2008:

M.G. Harasewych and M.]. Tenorio, 2018

a @ =Group 1

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Page 79

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TYPE SPECIMENS |

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Figure 25. Positions of primary type specimens in the discriminant space.

Fig. 2, Table 2) fell well within Group 1 (Figure 22)

(probabilities > 99 %), indicating that this phenotype has

been represented on the southern coast of San Salvador

between 125,000 and 140,000 years before present. Three

fossil specimens from Man Head Cay SMH 1b (Figure 27)

(Hearty and Schellenberg, 2008: Fig. 2, Table 2) also fell

well within Group 1 (Figure 23) (probabilities > 99.8 %),

indicating that this was the only phenotype occurring on

Page 80

THE NAUTILUS, Vol. 132, No. 3-4

Figures 26-29.

Fossil specimens of San Salvador Cerion used

in morphometric analyses. 26. Three specimens from the

French Bay Member of the Grotto Beach Formation [station

SFB4b(1), Hearty and Schellenberg, 2008: Fig. 2, Table 2],

(125,000— ie 000 years before present). 27. Three specimens

non Man Head Cay {station SMH 1b, Hearty and Schellenberg,

2008: Fig. 2, Table 2], (90,000-110,000 years before present).

28. Five specimens from The Gulf [Station STGIcl, He arty and

Schellenberg, 2008: Fig. 2, Table 2], (80,000-90,000_ years

before pre sent). 29. Thee specimens from Crab Cay [SCC4d

(1-2), Hearty and Schellenberg, 2008: Fig. 2, Table 2],

(70,000-80,000 years before present).

Man Head Cay between 90,000 and 110,000 years before

present. Modern Cerion on Man Head Cay are all of

Group 2 (Figure 20). Fossil samples STGlel (Figure 28)

(Hearty and Schellenberg, 2008: Fig. 2, Table 2 \ ron TN

Gulf (very near SH1 on Figure ays were morphologically

ai ‘rse (Figure 24), with three falling within Group

1 (probabilities > 99 %), 1 within Group 2 (probability

93 %) and one questionably within Group 3 (probability of

only 61.9 %), indicating that in addition to phenotype 1,

phe notypes 2 and possibly 3 were present on San Salvador

between 80,000 and 90,000 years before prese! nt, al-

though in a region of the island where phenotypes 2 and 3

do not occur Roakey, Three fossil samples from Crab Cay

(Figure 29) [SCC4d(1-2), Hearty and Schellenberg,

2008:Fig. 2, Table 2] near the geographic center for

Group 2 in the modern fauna, reveal that both Group 1

(probability > 95 %) and Group 2 (probabilities > 99 %)

(Figure 24) were present at this location between 70,000

and 80,000 years before present.

DISCUSSION

Despite minor differences in the number and types of

measurements and analytical algorithms, our results

concur with prior morphometric analyses (e.g. Hearty

et al., 1993; Fronabarger et al., 1997: Hearty and

Schellenberger, 2008) in identifying two primary shell

phenotypes of Cerion living on the idbarn of San Salvador,

one centered along the w estern coast (Group 1), the other

along the eastern coast (Group 2) of the island. The

presence of a third phenotype (Group 3) with a very

restricted inland distribution has not been previously

reported, since prior studies had not sampled in the area

in which it occurs.

A number of these studies had dealt with morphological

changes in Cerion shells throughout the continuous fossil

record that spans 140,000 years. Hearty et al. (1993: Fig.

6) had reported that a single phenotype was present on

San Salvador in the Grotto Beach Formation (MIS 5e),

the Gulf Unit (MIS 5a), and the Man Head Unit (MIS5a),

but that a different phenotype was present in the later

Almgreen Cay Formation (MIS5a). They noted that the

more recent Holocene Rice Bay Formation contained

shells with both phenotypes. They had also documented

(Hearty et al., 1993: Fig. 7) a significant increase in shell

length ‘and especially in shell mah within the Almgreen

Cay Formation (70,000—90,000 years before present).

Hearty and Sche llenberg (2008) ‘subseque ently reported

that change 5 in the morphology of Cerion shells on San

Salvador over the past 140,000 years appear to be con-

tinuous and generally directional, and that a trend of

increasing gross she ll size characterizes each of the in-

terglacial “phase s (MIS 5e, MIS 5a and MIS 1). They also

commented that live-collected Cerion have a range in

gross morphology that nearly encompasses that of the

entire fossil sequence.

In morphometric analyses of living and fossil Cerion

shell morphology, Fronabarger et al. (1997) concluded

that variation is dependent on ies and suggested

that environment affects shell shape. They also noted that

variation seen between living and fossil populations is the

same as the variation seen in the geogr aphic distribution

of living samples.

These studies interpret the considerable variation in

shell morphology, both geographic and chronostrati-

graphic, based on the premise that there was but a single

colonization of San Salvador by Cerion, and that all var-

iation resulted from subsequent differentiation within a

single lineage. Rose (1990) attributed the different

morphologies to intraspecific variation, with the larger,

thicker ribs found on shells that inhabited the canton

coast being a response to predation by crabs and rodents.

Hearty and Schellenberg (2008) attributed rapid changes

M.G. Harasewych and M.]. Tenorio, 2018

Page 81

in shell size during MIS 5a as responses to environmental

factors due to climate change during this interglacial

period with its warmer and wetter aiimate: aBuew con-

cluded that multiple Cerion extinctions and recoloniza-

tions during the past 140,000 years are improbable due to

the continuous fossil record, and that multiple species are

unlikely to occur on San Salvador.

Gould (1997) noted the intermediate position of San

Salvador he tween the "two major geographic domains of

Bahamian Cerion" and concluded that the Cerion fauna of

the island is descended from propagules from both

geographic regions. He considered Cerion watlingense

(Group 1) to be derived from the "ribby" bank- “edge

phenotype (see Gould and Woodruff, 1986) of the

Great Bahama Bank, while the "triangular" phenotype,

which he named Cerion rodrigoi (a junior synonym of C.

coloni Bartsch, 1924) (Group 3) to have originated in the

southeastern Bahamian islands. He speculated that a third

phenotype, corresponding to the "mottled", bank interior

phenotype of the northern Bahamas, would either be

absent or restricted to the interior of San Salvador because

the island has no bank-interior coast (a coast that was

inland when eustatic sea level fell, exposing larger banks

during glacial periods). He commented that it was not

known whether the "ribby" and "mottled" phenotypes

represent distinct, monophyletic lineages throughout the

northern Bahamas, or whether the plienotypes arise in-

dependently and convergently as adaptations to common

environments.

Our findings of three distinct Cerion phenotypes on San

Salvador are congruent with Gould’s hypotheses. Phe-

notype | corresponds to Cerion watlingense, derived from

a propagule originating on the Great Bahama Bank, has

been present on San Salvador since at least 125,000 to

140,000 years before present, and was widespread

throughout the island, occurring at Man Head Cay By

90, 000 to 110,000 years before present. Phenotype 2

corresponds to Cerion coloni, and is believed to be de-

rived from a propagule originating in the southern

Bahamas. Based on our limited sampling of fossil speci-

mens, earliest records are known from the southeastern

corner of San Salvador between 80,000 and 90,000 years

before present, although in a region of the island where

this phenotype does not occur today, This phenotype

appears to have expanded its range northward along the

eastern coastline of San Salvador, here it likely replaced

or hybridized with Phenotype 1. It co- occurred with

phenotype 1 at Crab Cay between 70,000 and 80,000

years before present, but is the only phenotype present

along the east coast in the modern fauna, including at Man

lead Cay, which was inhabited exclusively by phenotype

1 between 90,000 to 110,000 years before present.

Phenotype 2 continued to expand westward along the

northern coast of the island, including to several of the

offshore cays. Populations at the northern tip of North

Point Peninsula and Green Cay are dominated by Phe-

notype 2, although vestiges of Phenotype 1 still remain

at these localities. The presence of a third, morpho-

logically distinct phenotype (Group 3) on San Salvador,

corresponding to the "mottled" phenotype of the northern

Bahamas, is presently known from a single population in

the interior of the island, in agreement with Gould’s

prediction.

Analyses of shell morphology of living and fossil Cerion

occurring on San Salvador may be interpreted in the

context of several bioge -ographic scenarios, each with

different evolutionary aad systematic consequences. The

single introduction hypothesis predicts that all Cerion

living on San Salvador are part of a monophyletic clade,

and all have a most recent common ancestor not shared

with Cerion from other islands. As such, they may all be

members of a single species, Cerion watlingense, or the

phenotypes may have differentiated sufficie sntly to com-

prise either separate species or subspecies of C. watlin-

gense. If the multiple introduction hypothe sis 18 correct,

Cerion watlingense as well as Cerion inconspicuum would

be more blosely related to species from the Great Bahama

Bank, while Cerion coloni would be closer to species from

the southern Bahamas. A molecular phylogeny with ap-

propriate sampling should resolve between these two

hypotheses and provide insights into the processes by

which Cerion achieve and modify their patterns of ex-

treme morphological diversity.

ACKNOWLEDGMENTS

We are grateful to Dr. Paul Hearty for his extensive

ltdkaordk on San Salvador [supported by NSF award EAR

0106936 (Goodfriend, Gould, and Harasewych PI’s)

through a subcontract to Dr. Hearty], and for depositing

the wacker material for his published study, including

dated fossil specimens, in the collections of the Netiomell

Museum of Natural History, Smithsonian Institution. We

also thank Jessica Cundiff and Adam Baldinger of the

Museum of Comparative Zoology for alan available

specimens in their care. We chen the reviewers, Dr.

James Carew and Dr. Timothy Pearce, for their helpful

comments and suggestions.

LITERATURE CITED

Baker, H.B. 1963. Type land snails in the Academy of Natural

Sciences of Philadelphia part IT. Land Pulmonata, exclusive

of North America north of Mexico. Proceedings of the

Academy of Natural Sciences of Philadelphia 115: 191-25

Baldini, L.M., S.E. Walker, B. Railsback, ae Baldini, and

D.E. Crowe. 2007. Isotopic ecology of the modern land

snail Cerion, San Salvador, Belhanmer: preliminary advances

toward establishing a low-latitude island paleoenvir-

onmental proxy. Palaios 22: 174-187.

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

Papers of the Department of Marine Biology, Carnegie

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

Bartsch, P. 1924. Molluscan studies se the Florida

Keys, Bahamas, and West Indies. In: Explorations and

Field-Work of the Smithsonian Institution in 1923.

Smithsonian Miscellaneous Collections 76(10): 35-43,

figs. 36-41.

Page 82

THE NAUTILUS, Vol. 132, No. 3-4

Clench, W.J. 1938. Origin of the land and freshwater mollusk

ae of the Bahamas, with a list of the species occurring on

Cat and Little San Salvador Islands. Bulletin of the Museum of

Comparative Zoology 80(14): 481-541, pls. |= 3:

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

Pulmonata). Bulletin of the Museum of Comparative Zo-

ology 116: 121-169.

Dall, W.H. 1894. Cruise of the Steam Yacht “Wild Duck” in the

Bahamas, January to April, 1893, in charge of Alexander

Agassiz. II. Notes on the shells collected from the Shores of

the Great Lagoon, Watling Island, Bahamas. Bulletin of the

Museum of C Jomparative Zoology 25(9): 113-124, 1 plate.

Dall, W.H. 1905. Report on land and eh water shells “allie

in the Bahamas in 1904, by Mr. Owen Bryant and others.

Smithsonian Miscellaneous Collections 47(4): 433-452, pls.

58-59.

Fronabarger, A.K., J. L. Carew, and kK. Harborak. 1997. Mor-

phology of fossil and moder Cerion from San Salvador

Island, Bahamas. pp- 59-72. In: Carew, J. (ed.) Proceedings

of the 8" Symposium on the Geology of the Bahamas andl

other Carbonate Regions. Bahamian Field Station, San

Salvador, Bahamas.

Coordin nd, G.A. and S.J. Gould. 1996. Paleontology and chro-

nology of two evolutionary transitions by hybridization i in the

B: slnewratera land snail Cerion. Science 274: 1894-1897.

Gould, S J. 1997. The taxonomy and geographic variation of

Cerion on San Salvador (Bahama Islands) . pp. 73-91. In:

Carew, J. (ed.) Proceedings of the ger Symposium on the

Geology of the Bahamas and other Carbonate Regions.

B: Ahonen Field Station, San Salvador, Bahamas.

Gould, S.J., D.S. Woodruff, and J.P. Martin. 1974. Genetics and

morphometrics of Cerion at Pongo Carpet: A new sys-

tematic approach to this enigmatic land snail. Systematic

Zoology 23(4): 518-535.

Gould, S.J. and D.S. Woodruff. 1978. Natural History of the

genus Cerion VIII: Little Bahamas Bank—a revision based

on genetics, morphometrics, and geographic distribution.

Bulletin of the Museum of Comparative Zoology Harvard

148: 371-415.

Gould, S.J. and D.S. Woodruff, 1986. Evolution and systematics

of Cerion (Mollusca: Pulmonata) on New Providence Is-

land: A radical revision. Bulletin of the American Museum

of Natural History 182(4): 389-490.

Gould S.J. and D.S. Woodruff. 1990. History as a cause of area

effects: an illustration from Cerion on great Inagua, Bahamas.

Biological Journal of the Linnean Soest 40: 67-98.

Harasewych, M.G. 2012. The fossil record and phylogeography

of the family Cerionidae (Gastropoda: Pulmonata), with the

description of a new species from the Pleistocene of

Florida. The Nautilus 126: 119-126.

Hearty, P.]., P. Kindler, and S.A. Schellenberg, 1993. The Late

Quatermary Evolution of Surface Rocks on San Salvador,

Bahamas. pp. 205-222. In: White, B. (ed.) Proceedings of

the Sixth Symposium on the Geology of the Baharia

Bahamian Field Station, San Salv alo, Bahamas.

Hearty, P.J. and S.A. Schellenberg, 2008. Integrated late

Quatemary chronostratigraphy for San Salaclor Island,

Bahamas: Patterns and trends of morphological change in

the land snail Cerion. Palaeogeography, Palaeoclimatology,

Palaeoecology 267: 41-58.

ICZN, 2012. intentional Code of Zoological Nomenclature,

Fourth Edition, The Code Online. International Trust for

Zoological Nomenclature, London. http://www.iczn.org/

iczn/index.jsp#

Iturralde-Vinent, M.A. 2006. Meso-Cenozoic Caribbean

paleogeography: implications for the historical bio-

geography of the region. International Geology Review

48: 791-827

Mayr, E. 1963. Avstianel Species and Evolution. The Belknap

Press of Harvard University Press, Cambridge, xiv + 797 pp.

Pilsbry, H. A. 1901-1902. Family Cerionidae. Menwel of Con-

chology. Ser. 2. Pulmonata. Acade »my of Natural Sciences of

Philadelphia 14: 174-286, pls. 27-AT.

Poth, K.A., P.L. Gnivecki, M.J. Berman, and A. Michelson. 2010.

Morphometric comparison of prehistoric and modern

Cerion at Pigeon Creek Site, San Salvador, Bahamas.

cg Society of America, Abstracts with Programs,

- 60.

Roll oT 2005. tpsDig version 2.04. Department of Ecology

and Evolution, State University of New York at Stony Brook.

Available from: http://Life. Bio. SUNY SB.e dw/morph/morph.

html.

Rose, J.A. 1990. Cerion on San Salvador, Bahamas: Ecology

and Intraspecific Variation. Ph. D. Dissertation, Harv al

University.

Roth, B. and J.H. Hartman, 1998. A probable Cerion (Gas-

tropoda: Pulmonata) from the uppermost Cretaceous Hell

Creek Formation, Garfield County, Montana. Paleobios 18:

16-20.

Ruhoff, F.A. 1973. Bibliography and Zoological Taxa of Paul

ee Smithsonian Contributions to Zoology 142: 1-166.

Serucca, L., M. Fop, T.B. Murphy, and A.E. Raftery. 2016.

MCLUST 5: clustering, classification and density estima-

tion using Gaussian aati mixture models. The R Journal

8/1: 289-317.

Sheets H.D. 2003-2005. IMP-Integrated Morphometrics

Package, Buffalo. Available at: http:/Avww3.canisius.edu/

~sheets/morphsoft.html.

Sheets H.D. 2003-2014. IMP 8, Buffalo. Available at: http://

www3.canisius.edu/~sheets/IM P%208.htin.

Woodruff, D.S. 1978. Evolution and adaptive radiation of

Cerion: a remarkably diverse group of West Indian land

snails. Malacologia 17: 223-239.

Woodruff, D.S. and SJ Gould. 1987. Fifty years of interspecific

hybridization: genetics and morphometrics of a controlled

experiment on wine land snail Cerion in the Florida Keys.

Evolution 41: 1022-1045.

Zelditch, M.L., D.L. Swiderski, H.D., Sheets, and W.L. Fink.

2004. Geometric Morphometrics for Biologists: A Primer.

Elsevier Academic Press, New York and London, 437 pp.

THE NAUTILUS 132(3-4):83-90, 2018

Page 83

How many micromollusks are there? A case study on species

richness in Hawai‘i, with the description of a new species of

Murdochella (G astropoda: Epitoniidae)

Bianca Campagnari

Quasars to Sea Stars Teen Program

Santa Barbara Museum of Natural History

2559 Puesta del Sol, Santa Barbara CA 93105 USA

[email protected]

Daniel L. Geiger

Department of Invertebrate Zoology

Santa Barbara Museum of Natural History

2559 Puesta del Sol, Santa Barbara CA 93105 USA

[email protected]

ABSTRACT

Four statistical metrics (Chaol, Chao2, ACE, and rarefaction)

were used to estimate the total number of species of shelled

micromollusks based on composition of nine benthic samples

collected from the sublittoral zone off Wailea, Maui, Hawai'i.

There was a total of 250 species in the nine samples analyzed,

and the estimated total number of species of shelled micro-

mollusks in this area, based on the. statistical metrics, was

317-375 species. Murdochella eee new species is de-

scribed and Pelycidion habei (Kay, 1979) is discussed.

Additional Keywords: Biodiversity, endemism, oceanic island,

Indo-West Pacific

INTRODUCTION

This study estimates the species richness of shelled

marine micromollusks off of Wailea, Hawai'i. Micro-

mollusks are defined as having shells smaller than 5 mm

as adults (Geiger et al., 2007). There are only 1,300

named species of shelled marine mollusks in Hawai'i

(Severns, 2011), out of an estimated 37,000 species in

the entire Indo-West Pacific (Contrafatto and Minelli,

2011). This difference is likely because of Hawaii's

geographic isolation. Biodiversity in the Indo-West

Pacific peaks around the islands of Southeast Asia,

and decreases outward from there (Severns, 2011).

Endemism of Hawaii's malacofauna is 21%, which is the

world’s highest. Severns has recorded 394 micromollusk

species (Severns, 2011). Estimates for the total number

of named mollusk species in the world reach

70,000-76,000, while estimates including undescribed

species go as high as 200,000 (Rosenberg, 2014).

The shells in this study were part of a thanatocoenosis,

or death assemblage; the composition of the samples is

assumed to represent the composition of the surrounding

areas. We used non-parametric calculations of species

richness, which estimated the total number of species

based on the relative abundance of rare and common

species in the sample. An area from which a sample with a

relatively high number of rare species was taken is esti-

mated to ene ve a high number of unfound species, because

a high number a rare species indicates high species

heterogeneity and low sampling intensity (Gotelli et al.,

2007).

MATERIALS AND METHODS

Nine dredged samples from six locations off Wailea, Maui,

Hawaii, at depths of 85-140 meters were collected and

kindly made available by M. Severns (Figure 1, Table 1).

The bulk sediment samples were sorted, and all speci-

mens were deposited at the Santa Barbara Museum of

Natural History (SBMNH: Table 1); some paratypes were

also deposited in the Bernice P. Bishop Museum, Hon-

olulu, Hawai'i, USA (BPBM). Identifications were based

primarily on Kay (1979), Okutani (2000), Severns (2011),

and Okutani (2017).

A rarefaction curve was generated to display the av-

erage species accumulation of each sample and was ex-

trapolated to an estimate for the species richness of the

entire area using the iNEXT function from the R package

(Chao et al., 2016; Hsieh et al., 2016). Species richness

estimators used in this study were ACE (abundance-based

coverage estimate) (Chazdon, 1998), as well as Chao] and

Chao2 (Chao, 1984). ACE and Chaol combine all the

samples, while Chao2 considers each sample in the study

separ ately and constructs an estimate by comparing the

samples. ‘In the R package Vegan (Oksanen et al., 2017)

the estimateR and spe cpool functions estimated C heal,

ACE, Chao2 and their standard errors, respectively.

Images of select shells were taken by scanning e Jectron

microscopy (SEM). Standard met thods were applied for

uncoated specimens (Geiger et al., 2007). Light micro-

scope z-stack images were taken using a Zeiss Axioskop

2plus compound microscope (Zeiss, Oberkochern,

Germany). A Cognisys (Traverse City, Michigan, USA)

DF > 2

Page 84

THE NAUTILUS, Vol. 132, No. 3-4

Figure 1

ste pping motor connected to a C ognisys StackShot con-

troller and programmed through ZereneStacker (Rich-

land, Washington, USA) was couple d to the fine focus of

the microscope with a custom adapter (Aben Machine

Products, Canoga Park, California, USA). Step size was

calculated based on depth of field of numerical aperture

(2.4-152 um). Images were acquired with a Canon 5DsR

digital SLR camera; RAW files were processed with DxO

Optics Pro (DxO Labs, Paris, France), and stacked with

ZereneStacker using the Pmax algorithm. Further digital

imaging was carried out in AffinityPhoto (Serif Ltd.,

Nottingham, Uk).

RESULTS

Diversity Estimates: A total of 4,233 shells belonging to 250

discrete morphospecies were recovered from the samples.

The estimates that used abundance data for the total number

of species in the area encompassed a 95% confidence interval

of 288-404 species (rarefaction, ACE, Chaol: Figure 2,

Table 2). All three abundance estimates had overlapping 95%

confidence intervals, which suggests that the different

methods of estimating species richness were consistent, and

Map of Maui. Markers indicate locations of samples.

thus more likely to be accurate. On the order of three-

quarters of species of micromollusks in the overall sampled

area were represented in the individual samples.

To explore the possible correlation between number of

specimens and species in the sample and diversity esti-

mates, _ E and C haol were applie .d to each of the nine

samples (Table 3). Estimated species richness correlated

positively with aan size, but that correlation was in-

significant C < 088. p >> 0.1: Figure 3). Similarly,

de »pth was found to have no effect on number of speci-

mens found (r = 0.06, p >> 0.1: Figure 4).

New Records: The Hawaiian malacofauna has recently

been treated by Severns (2011), including 394 micro-

mollusks. Although Hawaii is among the best studied

tropical areas aril despite it being a rather low diversity

region relative to its tropical keine, the faunal inventory

is far from complete. Exciting finds such as the highly

distinctive Severnsia strombe vlna Geiger, 2016 highlight

the potential for new species discoveries, particularh ly in

micromollusks (Geiger, 2016). Some additions to the

Hawaiian malacofauna stemming from the present study

are detailed below.

B. Campagnari and D.L. 2018

Geiger,

Page 85

Table 1

waypoints.

Coordinates

Sample site Location

WP267-424 2.5 mi W off Wailea 20.673° N, 156.482° W

WP502-506 3.6 mi W off Wailea 2(0).689° N, 156.500° W

WP714-716 1.5 mi NE of Molokini 20.651° N 156.479° W

WP384-394 1.5 mi W of Wailea 20.670° N 156.467° W

WP267-468 2.3 mi W of Wailea 20.673° N 156.482° W

WP267-496 2.3 mi W of Wailea 20.673° N 156.482° W

20.660° N 156.494° W

20.673° N 156.482° W

mi W of Wailea

WP404-640

WP404-713

WP641-642

CO OO UW

OV Ot Ot

SYSTEMATICS

Epitoniidae Berry, 1910 (1812)

Murdochella Finlay, 1926

Diagnosis: Shell high-spired, many fine, low, axial la-

mellae, crossed by few spiral cords. Base flattened. Pro-

toconch bulbous, smooth.

Type Species: Scala levifoliata Murdoch and Suter,

1906, by original designation.

Remarks: The genus is currently known from New

Zealand (M. alacer Finlay, 1926, M. levifoliata) Australia

(M. macrina Iredale, 1936), Antarctica (M. antarctica

Dell, 1990), South Africa (M. crispata Kilburn, 1985, M.

lobata Kilburn, 1985), and Alaska (M. turritelliformis

Brown, 2018).

400 400

= 300 300

bent

% 200 200

—_

—

100

0 5000 10000 15000 20000 25000 300000

Number of Individuals

Figure 2. Interpolated (solid line) and extrapolated (dashed

line) rarefaction curve of all samples. The shaded region is 95%

confidence interval. The asymptote is at 350 species, 95%

confidence interval covers 305-394 species.

Depth SBMNH

101 m 424489424607, 424609, 453258

$588 m 265846—-265857, 265859—265934

120-125 m 266363-266380, 452777-452856

92m 26625 1-266296

88 m 266081—266181

88-92 m 266008, 265984—266007, 266009—266046,

966048—266054

139-141 m 266297—266333

139-140 m

266334—266362, 424916, 424917,

453262

Sample sites of this study. SBMNH: range of SBMNH registration numbers for the particular site. WP: Severns GPS

124-132 m

454728454731, 454733454738, 454740—454750,

454752454762, 454764, 454765, 454767—4548 15,

454817454819, 454859454915, 457572

Murdochella tricingulata new species

(Figures 5-14)

Description: Shell high-spired, holotype 2.7 mm

(Figures 5-6), to 4.5 mm (Figure 9). Shell off-white to

dark-tan. Protoconch bulbous, smooth. Up to ten tele-

oconch whorls. First whorl with more widely spaced,

stronger axial cords, no spiral sculpture (Figures 6, 8).

Second whorl with spacing and strength of axial lamellae

as on remainder or teloconch, peripheral spiral cord

distinct, adsutural spiral cord indistinct (Figures 5, 7,

9-11, 13, 14). Subsequent whorls with three spiral

cords: strongest to just below periphery, second at two-

thirds towards suture, last just above suture. Height of

lamellae variable, some specimens with irregular distal

horizontal extensions towards anterior (Figure 12). Base

concave with pericolumellar cord. Aperture subquadratic.

Type Material: Holotype SBMNH 266361; Paratypes:

SBMNH 266300 (two): Severns WP 404-640 (one) (all

from type locality); SBMNH 452826 (nine); BPBM

284613 (two): Severns WP 714-716 (three), all from

2.78 km NE Molokini, Maui, Hawai‘i, USA, 20°39.047' N,

156°28.759' W, 131-136 m; SBMNH 454728 (five):

Severns WP 641-642 (two), all from off Wailea, Maui,

Hawai'i, USA, 20°39.29' N, 156°29.56’ W, 136-144 m.

USA,

Type Locality: Off Wailea, Maui, Hawai‘,

20°29'38" N, 156°2938" W, 152-154 m.

Etymology: Latin tri- = three, -cingulata referring to the

spiral cords; noting the three spiral cords.

Remarks: Murdochella tricingulata is most similar to M.

levifoliata with the very tight axial lamellae and strong,

stepped spiral cords in the lower half of the whorl. The

Hawaiian species is overall more slender, and has three

distinct spiral cords, as compared to two in M. levifoliata.

Both M. antarctica and M. macrina have more rounded

whorls with less prominent spiral cords (Dell, 1990;

Page 86

THE NAUTILUS, Vol. 132, No. 3-4

Table 2.

Species richness estimates (95% confidence intervals) based on rarefaction curve, ACE, Chaol, and Chao2.

All specimens combined

Separate samples

Species in samples Rarefaction curve

250 350 (305-394 ) 343

ACE Chaol Chao2

(323-363) 346 (288-404)

425 (341-509)

Beechey, 2008; Engl, 2012). The South African (Transkei)

M. crispata has Sea nette sculpture with spiral cords well

above the periphery of the whorls, while M. lobata has

fewer but strong axial lamellae but no spiral cords

(Kilburn, 1985). Murchochella turritelliformis Brown,

2018 from the Aleutian Islands, Alaska (Brown, 2018),

has a proportionally wider shell and more numerous spiral

cords.

The new species exhibits a moderate degree of vari-

ability in the sculpture, which appears pradual/clinal

among the 19 specimens available. Acoondting, no fur-

ther taxonomic separation is justified. The lamellae can be

rather low (height less than half distance between la-

mellae) to rather high (height greater than distance be-

tween lamellae). The specimens with higher lamellae have

trapped debris between the lamellae, which cannot be

removed by sonication. The specimens with high lamellae

tend to deve ‘lop extremely fine and irregular etl hori-

zontal extensions towards the anterior part of the shell.

Those extensions are always pure white in color, con-

trasting with the off-white to dark-tan background color.

The specimens with higher lamellae also appear slightly

broader. As this difference in shape can be accounted for

by the projecting sculpture, the shape of the main body of

the shell is ve ry similar in all specimens.

Pelycidion habei (Kay, 1979)

(Figures 15-19)

Remarks: Severns (2011: pl. 36, figs 8-9) showed two

putative species of Pelycidion from Hawai'i i, the holotype

of P. habei and Pelycidion sp. They differ nominally in the

glassy (P. habei) vs. opaque white (P. sp.) shell color, the

degree the whorls are inflated or incised at suture, re-

spectively, and the apparent smooth (P. habei) vs. finely

clathrate (P. sp.) shell sculpture. The illustrations in Kay

(1979: 27, i-j) reveal fine clathrate sculpture, which

Table 3.

suggests that the apparent differences in the images in

Severns (2011) are either due to specimen condition or

photographic technique, but that they are not systemat-

ically relevant. The three specimens illustrated by SEM

here ( Figures 15-17) show quite a bit of variability with

respect to degree of incision of the whorls are in-

termediate between the specimens figured by Severns

(2011). It suggests that there is only a single species of

Pelycidion in Hawaii.

DISCUSSION OF DIVERSITY

All diversity measures are estimates, and one needs to be

aware of their inherent limitations. While an estimator

based on incidence data such as Chao2 can be useful

because it takes distribution heterogeneity into account, in

this study estimators using lnmadenee data such as ACE

and Chaol were more use ‘tal because of the small number

of samples analyzed and the fact that the samples in this

study were of different sizes. The Chao2 estimate is

known to stabilize above 30 samples (Gotelli, 2011), while

this study was based on only nine samples. Regardless,

even though the Chao2 estimate is much larger Siva the

others, the Chao2 confidence interval includes the other

estimates.

All models used in this study are minimum richness

estimators (O’Hara, 2005). Richness estimates of a par-

ticular area have been found to be correlated with sample

size, suggesting underestimation due to low sample size

Cmitroras et ail, 201 3). Here, this correlation was positive

but insignificant (r 4 & (N.S. 1) >> 0.1, Figure 3). Because

of the correlation between richness ential and sample

size, estimates are ideally compared between samples of

equal numbers of individuals from different locations to

compare species richness (Chao, 2016). The sample s in

this study had unequal numbers of specimens (66-650),

though the raw sediment samples had approximately the

ACE and Chaol species richness estimates and standard error for each sample. WP: Severns GPS waypoints.

Number of specimens Number of species ACE ACE 95% Chaol Chaol 95%

Sample Site in sample in sample estimate confidence interval estimate confidence interval

WP267-624 196 UU 85 75-95 fol 73-89

WP502-506 102 56 123 109-137 110 54-166

WP714-716 432 64 87 77-97 90, 60-120

WP384-394 195 37 i, 62-82 61 33-89

WP267-468 617 $4 129 117-141 131 87-175

WP404-640 66 28 40) 36-46 39 23-55

WP404-713 1 30 51 47-55 43 25-61

WP64 1-642 650 96 115 105-125 12] 95-47

WP267-496 294 57 126 112-140 163 49-277

B. Campagnari and D.L. Geiger, 2015

Page 87

180

160

120

Richness

40 a

0 100 200 300

Specimens

Figure 3.

400 500 600 700

Species richness estimates of samples over number of specimens in sample. Statistically insignificant positive correlation

(r < 0.38, p >> 0.1). Diamonds: ACE estimate. Squares: Chaol estimate. Solid line: Chaol regression. Dashed line: ACE regression.

same volume and were collected within kilometers of each

other. Depth (88-141 m) as a factor explaining those

differences could be ruled out (1? OS, jo» SS Ol,

Figure 4). The observed differences appear to reflect true

environmental heterogeneity.

The estimated number of species for each sampling site

was 62-88% lower than that for the whole area, which

shows high distribution heterogeneity. The sum of sample

species aighmness is much greater than the total species

richness due to many common species among samples.

Estimating species richness is known to be more difficult

for highly “ay erse locations (Chao, 2016), such as Maui.

Comparison of the data presented here is challenging,

because of the absence of comparable studies at other

localities. An environmental impact report (Russo et al.,

2018) from a waste water treatment outfall site at 33 m off

Oahu listed 165 species of mollusks of all sizes from

700

600

S00

400

300

Specimens

200 @ ¢

100 °

80 390 100 110

Figure 4.

Linear regression of mean depth vs number of specimens found (

approximately 4,500 individuals. Samples were collected

by SCUBA diver and small cores, hence, included the full

size-range of organisms. Diversity estimators calculated

from Ihe raw ate were highly consistent (this study).

Chaol estimates a species idhness of 193+12SE, 95%

confidence interval 169-217, and ACE estimated 197+

TSE, 95% confidence interval 183-211 (this study). The

different size compositions of the two studies make direct

comparison difficult. Based on the species list, we esti-

mate that less than half the species in Russo et al. (2018)

are of comparable small size as in our samples from Maui;

that proportion is conservative and in rough agreement

with other studies (Aravind et al., 2008; Middlefart et al.,

2016: see also below). Accordingly, the contrast of of

species richness of micromollusks alone is even more

remarkable: Oahu ~80 found, ~100 estimated; Maui

250 found, ~350 estimated. These differences suggest

120 130 140 150

Depth (m)

r = 0.06, p >> 0.1).

Page 88

THE NAUTILUS, Vol. 132, No. 3-4

Figures 5-14.

teloconch. 7-12. Paratypes SBMNH 45472

Murdochella dicate s new Me cies. 5-6. Holotype SBMNH 266361. 5. Shell. 6. Detail of protoconch and early

7. Shell of small s specimen with low sculpture. 8. Protoconch of 7. 9. Largest specimen

known (4.2 min). 10. SEM image. 11. oe mab of 10. 12. Enlargement of lamellae with irregular distal horizontal extensions towards

anterior. 13-14. Paratype SBMNH 452826. 13. Dark-colored specimen with strong axial lamellae. 14. Lighter colored specimen with

lower axial lamellae. Scale bars shells = 1 mm. Scale bars details

environmental heterogeneity and that the sites sampled in

this study are more tage rse than the site sampled in

Oahu. Differences in de »pth may account for the differ-

ence in diversity, as the Oahu samples were taken at 33 m

and the Maui samples were taken from 85-141 m. An-

thropogenic effects were discounted by Russo et al. (e018)

as similar species composition and numbers of species

were found in control sites.

Bouchet (2002) recorded 864 species smaller than

4.1 mm off Koumac, New

Caledonia. The higher

100 pm.

number of species found in New Caledonia is probably

due to the fact that the sampling area was larger

(295 km? vs 18 km?) with greater habitat diversity, and

the fact that New Cale Henin is less isolated than Maui.

The absolute latitudes of both areas are similar (Koumac

20.5°S, Maui: 20.7°N).

Many micromollusks are missed in habitat assessments

because they are smaller than the holes in the mesh used

to sort shells out of a sand sample. Additionally, some

micromollusks require the use of a scanning electron

B. Campagnari and D.L. Geiger, 2015

ace BC

Page 89

Figures 15-19. Pelycidion habei (Kay, 1979). SBMNH 454742. 15-17. Shells. 18. Protoconch of 15. 19. Enlargement of body

sculpture from 15. Scale bar shells = 1 mm. Scale bars details =

microscope to observe distinguishing characteristics

(Middlefart et al., 2016). Despite relatively few studies on

micromollusks, they appear to be very common. Of the

269 recorded gastropod species of the Western Ghats,

40% were micromollusks (Aravind et al. 2008), and 33% of

recovered species of mollusks from a study in New

Caledonia were micromollusks (Middlefart et al., 2016).

The present study shows that even a rather well-studied

area such as Hawai'i still harbors a significant percentage

of undescribed species. Given Blawentt s high rate of ma-

rine endemism, it highlights the need for strong con-

servation measures to protect the yet to be recognized

biodiversity.

ACKNOWLEDGMENTS

BC was supported in part by a generous donation from

Laura Francis, Education and Omirendh coordinator at

NOAA Channel Islands National Marine Sanctuary, and the

Sea Forward Fund. Jenna Rolle (SBMNH) kindly provided

insights into diversity estimates. Marta deMaintenon and

Cynthia Hunter helped with constructive comments to

improve the manuscript and provided reports on samples

from Oahu. José H. Leal provided editorial guidance.

Regina Kawamoto (BPBM) provided the registration

number for one paratype lot.

LITERATURE CITED

Aravind, N.A., R.K. Patil, and N.A. Madhyastha. 2008. Micro-

molluscs of the Western Ghats, India: diversity, distribution

and threats. Zoosymposia 1: 281-294.

Beechey, D. 2008. Murdochella macrina. http://seashellsofnsw.org.

awEpitoniidae/Pages/Murdoche Ha_macrina.htm [accessed 8/

28/2018]

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

Assessing the magnitude of species richness in tropical

100 um.

marine environments: exceptionally high numbers of

molluscs at a New Caledonia site. Biological Journal of the

Linnean Society 75: 421—436.

Brown, L.G. 2018. New species of Nystiellidae and Epitoniidae

(Mollusca: Gastropoda) from the northeastem Pacific, Mol-

luscan Research. DOI: 10.1080/13235818.2018.1433956

Chao, A., R.K. Colwell, C. Lin, and N.J. Gotelli. 2009. Sufficient

sampling for asymptotic minimum species richness esti-

mators. Ecology Society of America 90: 1123-1133.

Chao, A., T.C. Hsieh, and K.H. Ma. 2016. iNEXT: an R package for

rarefaction and extrapolation of species diversity (Hill num-

bers). Methods in Ecology and Evolution 7: 1451-1456.

Chao, A. 1984. Nonparametric estimation of the number of classes

in a population. Scandinavian Journal of Statistics 11: 265-270.

Chazdon, R.L., R.K. Colwell, J.S. Denslow, and M.R. Guariguata.

1998. Gitisticd methods for estimating species aries of

woody regeneration in primary and secondary rainforests of

aortieostermn Costa Rica. In: Forest Biodiversity Research,

Monitoring and Modeling. F. Dallmeier and J.A. Comiskey

(eds.). Parthenon Publishing, W ashington D.C., pp. 285-309.

Contrafatto, G. and A. Minelli 2011. Biological Science Fun-

damentals and Systematics. EOLSS Publishers, Oxford,

331 pp.

Dell, R.K. 1990. Antarctic Mollusca. The Royal Society of New

Zealand Bulletin 27: 1-311.

Engl, W. 2012. Shells of Antarctica. ConchBooks, Hackenheim.

402 pp.

Geiger, D.L. 2016. Severnsia strombeulima n. gen. & sp. from

Hawai'i (Mollusca: Gastropoda: Caenogastrpoda: Eulimi-

dae). Zootaxa 4084: 587-589.

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

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

preparing, storing and examining small molluscan speci-

mens. Molluscan Resear 97: 1-50.

Gotelli, N.J. and R.K. Colwell. 2011. Estimating species richness.

In: Biological Diversity. A.E. Magurran and B.J. McGill

(eds.). Oxford Biology, Oxford, pp. 39-54.

Hsieh, T.C., K.H. Maand A, Chao. 2016. iNEXT: iNterpolation and

EXT rapolation for species diversity. R package version 2.0.12.

Kay, A. 1979. Hawaiian Marine She lls. Bishop Museum Press,

Honolulu. 653 pp.

Page 90

Kilbum, R.N. 1985. The family Epitoniidae (Mollusca: Gas-

tropdoda) in southern Africa and Mozambique. Annals fo

the Natal Museum 27: 239-337.

Middlefart, P.U., L.A. Kirkendale and N.G. Wilson. 2016.

Australian tropical marine micromolluses: an overwhelming

bias. Molecular Diversity Preservation International 8: 17.

O'Hara, R.B. 2005. Species richness estimators: how many

species can dance on the head of a pin? Journal of Animal

Ecology 74: 375-386.

Oksanen, J., F.G. Blanchet, M. Friendly, R. Kindt, P. Legendre,

D. McGlinn, P.R. Minchin, R. B. O'Hara, G.L. Simpson, 2.

Solymos, M. Henry, H. Stevens, E. Szoecs and H. W agner.

2017. vegan: Community Ecology Package. R package

version 2.4-4. https://C RAN.R- -project. org/package =vegan

Okutani, T. 2000. Marine Mollusks in Japan. Takai University

Press, Tokyo. 1173 pp.

THE NAUTILUS, Vol. 132, No. 3-4

Okutani, T. 2017. Marine Mollusks in Japan. Second edition.

Takai University Press, Kanagawa. 1375 pp.

Rosenberg, G. 2014. A new critical estimate of named species-

level diversity of the recent Mollusca. American Malaco-

logical Bulletin 32: 308-32

Rivaso, AIR. J.H. Bailey- ee W. J. Cooke, C.L. Hunter, and

R.K. Kawamoto. 2018. Benthic Sampling in the Vicinity of

the Mokapu Ocean Outfall, O'ahu, Hawai'i, March 2018.

Water Resources Research Center. Univ ersity of Hawaii at

Manao, Honolulu. 182 pp.

Severns, M. 2011. Shells of the Hawaiian Islands. ConchBooks,

Hackenheim. 564 pp.

Zmihorski, M, P. Sienkiewicz, and P. Tryjanowski. 2013. Never

ending story: a lesson in using eae efficiency methods

with ground beetles. Journal of Insect Conservation 17:

3335337

THE NAUTILUS 132(3-4):91—100, 2018

ace

Page 91

Catalog of the taxonomic updates of northeastern Pacific Late

Cretaceous shallow-marine bivalves and gastropods named from

1874 to 1966

Richard L. Squires

Department of Geological Sciences

California State University

Northridge, CA 91330-8266, USA

and

Invertebrate Pale ontology!

Natural History Museum of Los Angeles C Jounty

Los Angeles, CA 90007 USA

richard.sq [email protected]

ABSTRACT

Published taxonomic updates of northeastern Pacific (southem

Alaska to Baja Califomia, Mexico) shallow-marine Late Creta-

ceous bivalves and gastropods named between 1874 and 1960,

are catalogued for the first time. Of the 143 species (including

“varieties,”) 68 are bivalves and 75 are gastropods. These species

were named prior to the usage of more effective paleontologic

procedures initiated by Willis P. Popenoe in 1957. The junior

homonym Inoceramus pacificus Anderson and Hanna, 1935 is

renamed as Inoceramus georgeedavisi new name Squires.

Additional Keywords: Vancouver Island, California, Inoceramus

INTRODUCTION

The purpose of this paper is to catalog, for the first time, the

taxonomic name changes of shallow-marine species of Late

Cretaceous bivalves and gastropods, which were named by

early molluscan paleontologists (between 1874 and 1966) in

the region extending from southern Alaska to northern Baja

Calligoveatn Mexico. This region corresponds to the “Northeast

Pacific Subprovince (NEP)” of Kauffman (1973: fig. 1).

Squires (2018) provided a discussion of the geologic setting

and paleoclimate conditions of this paleo-biotic subprovince.

Stemming from his field work in California in the 1860s,

William M. Cabb was the first paleontologist to collect and

name Cretaceous shallow-marine mollusks from the NEP

region. His monumental contributions were catalogued

hee Groves and Squires (2018). Subsequent NEP pale-

ontologists during the late 1880s to the late 1950s were, in

alphabetical order, Frank M. Anderson, Ralph Arnold,

James G. Cooper, Donald H. Dailey, E. B. Hall and

Arthur Ambrose, Roy D. McLellan, Charles W.

] .

Research Associate

Merriam, Michael A. Murphy, Earl L. Packard, Willis P.

Popenoe, Peter U. Rodda, Clarence A. Waring, Charles A.

White, and Joseph F. Whiteaves. Although Anderson’s

Late Cretaceous monograph was published in 1958, his

research was actually done in the 1940s. He died in 1945,

and the volume was published posthumously 13 years

after his death.

Starting in the late 1950s, there was a fundamental

transformation in how NEP Late Cretaceous bivalves and

gastropods were studied. Popenoe (1957: 430) imple-

mented a method that relied mainly on his own large and

well-curated collection of fossils: a collection now re ssicling

at the Natural History Museum of Los Angeles County

Museum of Natural History. “His method of careful

collecting, the recording of precise geographic locality and

stratigraphic position, ‘and accurate identifications has

resulted in so much more orderly knowledge of the Late

Cretaceous molluscan faunas that it now seems impossible

that these faunas could have been in such confusing

disarray as when he began his studies” (Saul et al., 1989:

117). Popenoe selected a representative number of speci-

mens from each major formation in which they occur and

then carefully cleaned the hinges of bivalves and the ap-

ertures of gastropods. C Careful “deamon of these critically

important morphologic areas allowed fora much better basis

for the systematic placement of taxa. He used detailed

biostratigraphy to more accurately show lineages of species

and their evolutionary trends. Initially, he waltts d only on

ammonite data for biostratigraphic control but later

(Popenoe et al., 1960) added inoceramid-bivalve zones. Saul,

who was mentored by Popenoe, perfected the method

begun by him and gr eatly helped to introduce it to current

NEP molluscan paleontologists. The method relies on new

biologic and stratigraphic data in order to keep everything

updated. In the last 20 years or so, there have been. many

updated systematic and biostratigraphic studies of NEP

Dace 92

Page 92

THE NAUTILUS, Vol. 132, No. 3-4

bivalves and gastropods, and these studies are the basis of

this present paper. It is relevant to mention that species

named after the onset of Popenoe’s transformation are much

less prone to being changed, than those named prior to it.

There are, nevertheless, many other bivalves and gastropods,

named prior to 1960, which are in need re-investigation.

More name updates will be made in the future.

MATERIALS AND METHODS

The information was largely extracted from the literature.

Study area papers concerning Late Cretaceous bivalves

and gastropods but without any taxonomic updates were

not included. Cold-seep taxa are not included. The au-

thors are listed in alphabetical order. Bivalves are listed

before gastropods, but a few of the listed species concern

only gastropods. The species names, which are also in

alphabetical order, include the parenthesized genus name

in which the author originally placed the species, along

with its date, page, figure number(s), and holotype

number, unless otherwise noted. The museum. type-

specimens numbers were mostly gleaned from the

cited literature. The CASG numbers were derived from

its in-house records. The USNM numbers were derived

from the USNM online database [natural history.si.edu/

rc]. All of the “original” information is followed by the

updated name (in bold) and the currently known ge ologic

age, geographic range (listed from north to south), and

FER oReOS (s) concerning the updates of the species. For

the geologic age, standard (European) geologic stages of

the Late Cretaceous are used, with the sequence from

older to younger being Cenomanian, Turonian, Con-

iacian, Santonian, Campanian, and Maastrichtian.

Abbreviations used in conjunction with type-specimen

numbers are: CASG: California Academy of Sciences, San

Francisco; GSC: Ge ological Survey of Canada, Ottawa,

Ontario; LACMIP: Neveu History Museum of Los

Angeles County, Invertebrate Pale ontology Department,

Los Angeles: LSJU, Leland Stanford Junior University,

Sramiiorl California [collection now at CASG]; UCMP:

University of California, Berkeley, Museum of Paleon-

tology,; UO, University of Oregon, Eugene, Oregon;

USNM, National Museum of Natural History, Smithso-

nian Institution. Abbreviations used for geographic areas

are: BX (Baja California, Mexico), CA (California), QC

(Queen Charlotte Islands, British Columbia, VI (Van-

couver Island area, British Columbia; including adjacent

islands along eastern coast), WA (Sucia Island, Washington).

ANDERSON

Bivalves

anaana (Pholadomya) Anderson, 1902: 73, pl. 7, fig. 151;

plastohypotypes UCMP 30251, 30252. Liopistha anaana

(Anderson, 1902), Turonian, CA. Popenoe (1937: 384),

Anderson (1958: 119).

banosensis (Glyc Yes) Anderson, 1958: 98-99, pl. 73,

figs. 1-3; CASG 28310.01. Glycymerita banosensis

(Anderson, 1958), late Campanian to latest Maas-

trichtian, CA, BX. Squires (2010: 908).

bowersiana (Trigonia) Anderson, 1958: 116, pl. 26, fig. 8

CASG 252.01. Musagonia bowersiana (Anderson,

1958), Turonian, CA. Cooper (2015: 27); no age update

found.

branneri (Trigonia) Anderson, 1958: 112, pl. 17, fig. 5; UO

26859. Louella fitchi (Packard, 1921). Cenomanian/

Turonian boundary, OR, CA. Saul (1978: 53), Cooper

and Leanza (2017: 330).

buttensis (“Trigonocallista”) Anderson, 1958: 140, pl. 59,

figs. 1, la; CASG 27838.15. Calva (Egelicalva) but-

tensis (Anderson, 1958), early Campanian, AK, VI,

CA. Saul and Popenoe (1992: 36).

churchi (Trigonia) Anderson, 1958: 115, pl. 17, figs. 6, 7

CASG 1788.01. Notoscabrotrigona evansana (Meek,

1858), early to late Campanian, VI, CA. Jones (1960:

436), Cooper (: Ke BS).

colusaensis (Trigonia) Anderson, 1958: 110, pl. 1, fig. 6;

lectotype UCMP 12171. Yaadia leana Gabb, 1876,

early Turonian to Coniacian, OR, CA. Saul (1978: 33).

contracostae (Inoceramus) Anderson (1958: 103, pl. 18,

figs. 3, 4), CASG 29084.01, two forms: a a Coniacian

species of Sphenoceramus? and a Coniacian? to

Santonian species of Mytiloides?, CA. Kauffman

(1977: 189).

duplicostatus (Inoceramus) Anderson, 1958: 100, pl. 17,

figs. 3, 4, CASG 61724.02. Mytiloides duplicostatus

(Anderson, 1958), Turonian, CA. Kauffman (1977: 189).

eolobatus (Inoceramus) Anderson, 1958: 99, pl. 18, fig. 13;

CASG 1291.01. Inoceramus crippsi? Mantell, 1822,

early to mid Cenomanian, CA. Kauffman (1977: 188).

gabbiana (Mactra) Anderson, 1902: 74, pl. 7, fig. 156;

CASG 1. Cymbophora gabbiana (Anderson, 1902),

Turonian, CA. Saul (1974: 1084).

glennensis (Inoceramus) Anderson, 1958: 99, pl. 17, figs.

1, 2; CASG 61629.01 Mytiloides sp., most likely M.

opalensis elongata (Seitz, 1934), early Turonian, CA.

(Kauffman, 1977: 188).

hemphilli (Trigonia) Anderson, 1958: 115, pl. 52, figs. 9,

Ya, 9b. CASG 994. Popenoella hemphilli (Anderson,

1958), early Maastrichtian, CA. Saul (1978: 50), Cooper

and Leanza (2017: 329).

jacksonensis (Inoceramus) Anderson, 1958: 100, pl. 43,

figs. 1, 2; not in CASG database. Inoceramus crippsi?

Mantell, 1822, early to mid Cenomanian, OR.

Kauffman (1977: 188).

klamathensis (Inoceramus) Anderson, 1958: 104, pl. 18,

figs. 1, 2; syntypes CASG 61914.01, 61914.02. Sphe-

noceramus lingua? (Goldfuss, 1836), Coniacian? to

Campanian, CA. Kauffman (1977: 186).

klamathonia (Trigonia) Anderson, 1958: 112, pl. 30, fig.

4., CASG 61873. Notoscabrotrigona klamathonia

(Anderson, 1958), Turonian. Cooper (2015: 25), CA.

meekianus (Inoceramus) Anderson, 1958: 101, pl. DO). figs.

5, 6; CASG 228.01. Mytiloides meekianus (Anderson,

1958), Coniacian?, WA. Kauffman (1977: 180).

pentzana (Glycymeris) Anderson, 1958: 98, pl. 74, figs. 9.

2a; CASG 61874.01. Glycymeris veatchii (Gabb,

R.L. Squires, 2018

Page 93

1864), Turonian to late

Squires (2010: 902).

pacifica (Opisoma) Anderson, 1958: 122, pl. 26, figs. 5-6;

CASG 29118.03. Opis (Hesperopis) triangulata

(Cooper, 1894), late Campanian through early late

Maastrichtian, CA, BX. Squires and Saul (2009: 1329).

pacificus (Inoceramus ) Anderson and Hanna (1935: 29, pl.

10, fig. 4); CASG 1430.01, early Maastrichtian, BX

(Miller and Abbott, 1989: 63, fig. 1). Anderson and

Hanna’s name is a junior homonym of Inoceramus

pacificus Woods (1917: 28, pl. 14, figs. 1, 2) from Upper

Cretaceous (lower Santonian) strata on South Island of

New Zealand (Crampton et. al, 2000: 322, fig. 12).

Anderson and Hanna’s junior homonym is renamed

herein as Inoceramus georgeedavisi new species

Squires, in honor of the late George Edward Davis,

geologist and molluscan paleontologist.

pac ificus ( (Pectunculus) Anderson, 1902: 74, pl. 7, fig. 159;

voucher GASG 70504. Glycymeris pacifica ( Usdin,

1902), late Cenomanian to Turonian, OR, CA, BX.

Squires (2010: 898).

peninsularis (Corbis) Anderson and Hanna, 1935: 31, pl.

10, fig. 1; UCMP 36119. Calva (Calva) peninsularis

(Anderson and Hanna, 1935), latest Campanian and

early Maastrichtian, CA, BX. Saul and Popenoe

(1992: 17).

roguensis (Avicula) Anderson, 1958: 106, pl. 38, fig. 7;

CASG 445.17. Pteria pellucida (Gabb, 1864),

Turonian to Santonian, OR, CA. Squires (2014: Dil),

rosarioensis (Opis) Anderson and Hanna, 1935: 31, pl. 10,

figs. 2, 3; UCMP 36120. Opis (Hesperopis) rosar-

ioensis (Anderson and Hanna, 1935), early middle to

middle late Campanian, CA, BX. Squires and Saul

(2009: 1322).

shastaensis (Glycymeris) Anderson, 1958: 97, pl. 19, figs.

8-10; CASG 27830.01. Glycymeris veatchii (Gabb,

1864), Turonian to late Campanian, VI, CA, BX.

Squires (2010: 902).

taff (“Trigonocallista”) Anderson, 1958: 139; CASG

Base OL, Calva (Egelicalva) taffi (Anderson, 1958),

Ganiheian to Santonian, CA. Saul and Popenoe

(1992: 34).

wheelerensis (Trigonia) Anderson, 1958: 116, (unfig.):

CASG 10632. Yaadia leana (Gabb, 1877), Coniacian

to early Turonian, OR, CA. Saul (1978: 33).

Campanian; VI, CA, BX.

Gastropods

argonautica (Cypraea) Anderson, 1958:177, pl. 21, figs. 4,

4a; CASG 61856.05. Protocypraea argonautica

(Anderson, 1958), Turonian, OR, CA. Lorenz

(2017: 205).

bellavistana (Acteonina) Anderson, 1958: 157, pl. 29, fig.

2; CASG 1293.01. Paosia californica (Gabb, 1864).

late Cenomanian to Turonian, VI, OR, CA, BX. Squires

and Saul (2004a: 492).

berryessensis (Acteonina) Anderson, 1958: 157, pl. 29, fig.

3; CASG 3192.01. Paosia californica (Gabb, 1864),

late Cenomanian to Turonian, VI, OR, CA, BX. Squires

and Saul (2004a: 492).

eed S ypraea) Anderson, 1958: 176, pl. 63, figs. 2,

2b; CASG 31918.02. Protocypraea ERAT

eae 1958), Turonian, CA. Lorenz (2017: 205).

biconica (Gosavia) Anderson, 1958: 175, pl. 75, figs. 3, 3a;

CASG 61935.01. Konistra biconica (Anderson,

1958), Turonian, CA. Saul and Popenoe (1993: 381).

colusaensis (Acteonina) Anderson, 1958: 158, pl. 21, fig.

14: CASG 1291.05. Paosia colusaensis (Anderson,

1958), late Albian to early Cenomanian, CA. Squires

and Saul (2004a: 491).

condoniana (Anchura) Anderson, 1902: 76, pl. 8, fig. 179:

CASG 445.30. Anchura (Helicaulax) condoniana

Anderson, 1902, Turonian, OR, CA. Saul and

Popenoe (1993: 354).

crossi (Volutoderma) Anderson, 1958: 174, pl. 16, figs. 3,

3a; CASG 61934.02. Cydas crossi (Anderson, 1958),

Turonian, CA. Saul and Popenoe (1993: 362).

frazierensis Acteonella Anderson, 1958: 161, pl. 29, figs. 7,

7a; CASG 28103. Trochactaeon (Trochactaeon)

frazierensis (Anderson, 1958), Turonian, CA. Sohl

and Kollmann (1985: 79).

garzana (Phasianella) Anderson, 1958: 162, pl. 73, fig. 5;

CASG 28311.02. Tylostoma? garzana (Anderson,

1958), late early to early late Maastrichtian, CA.

Squires and Saul (2004b: 27).

gualalaensis (Cypraea) Anderson, 1958: 176, pl. 62, figs. 8,

8a; CASG 61918.01. Protocypraea gualalaensis (Anderson,

1958), early Maastrichtian, CA. Lorenz (2017: 205).

jacksonensis Volutoderma Anderson, 1958: 174, pl. 21, fig.

1; CASG 445.16. Drilluta jacksonensis (Anderson,

1958), Turonian, OR, CA. Saul and Popenoe (1993: 368).

nortonensis Bullina Anderson, 1958: 178, pl. 21, figs. 13,

13a; CASG 61850.03. Ellipsoscapha nortonensis

(Anderson, 1958), Campanian, CA. Stecheson (2004: 102).

packardi Acteonella Anderson, 1958: 160, pl. 29, figs. 4,

4a, 4b: not in CASG database. Trochactaeon (Tro-

chactaeon) packardi (Anderson, 1958), Turonian.

Soh] and Kollmann (1985: 83).

robertiana (Nerinea) Anderson, 1958: 155, pl. 66, fig. 3;

UCMP 33954. Turritella chaneyi Merriam, 1941,

early Maastrichtian, CA. Saul (1983: 81), Saul and

Squires (1998: 465).

roguensis (Acteonina) Anderson, 1958: 158, pl. 30, figs. 5,

5a; CAS 61906.01. Paosia californica (Gabb, 1864),

late Cenomanian to Turonian, VI, OR, CA, BX. Squires

and Saul (2004a: 492).

rustica (Acteonella) Anderson, 1958: 161, pl. 29, fig 5;

CASG_ 33721.01. Trochactaeon (Trochactaeon)

frazierensis (Anderson, 1958), Turonian, CA. (Sohl

and Kollmann, 1985: 79, 85).

stewarti (Nerinea) Anderson, 1958: 155, pl. 30, figs. 2, 3;

CASG 61842.02. A cold-seep epitoniid, ?middle

Turonian, CA. Saul and Squires (1998: 465).

ursula (Acteonina) Anderson, 1958: 158, pl. 63, fig. 4

CASG 31210.01. Paosia ursula (Anderson, 1958),

Coniacian, CA. Squires and Saul (2004a: 494).

ursulagorda (Acteonina) Anderson, 1958: 159, pl. 63, fig.

5: CASG 31210.02. Paosia ursula (Anderson, 1958).

Coniacian, CA. Squires and Saul (2004a: 494).

> > C

Page 94

THE NAUTILUS, Vol. 132, No. 3-4

siskiyouensis (Gyrodes) Anderson, 1902: 76, pl. 8, figs.

167, 168: CASG 41. Natica conradiana (Gabb,

1864), Turonian, OR, CA. Popenoe et al. (1987: 92).

yrekensis (Acteonina) Anderson, 1958: 157, pl. 29, fig. 2;

CASG 61847.01. Paosia californica (Gabb, 1864),

late Cenomanian to Turonian, VI, OR, CA, BX. Squires

and Saul (2004a: 492).

ARNOLD

Bivalve

stantoni (Mactra) Arnold, 1908: 357, pl. 31, fig. 3; USNM

31001. Cymbophora_ stantoni (Arnold, 1908),

Campanian, CA. Saul (1974: 1076).

Gastropod

pescaderoensis (Turritella) Arnold, 1908: 358, pl. 31, fig.

7; CASG 69777. Turritella chicoensis pescaderoensis

Arnold, 1908, late Campanian, CA. Saul (1983: 64).

COOPER

Bivalves

triangulata (Corbula) Cooper, 1894: 49, pl. 2 [not 4], fig.

42; lectotype CASG 624. Opis tana! trian-

gulata (Cooper, 1894), late Campanian to early

Maastrichtian, CA, BX. Squires and Saul (2009: 1329).

bowersiana (Cucullaea) Cooper, 1894:48, pl. 5, figs. 61,

62; CASG 66028.0. Calva (Egelicalva) bowersiana

(Cooper, 1894), late early through late Campanian,

VI, CA. Saul and Popenoe (1992: 39).

Senta ety

capuloides ( Sieanania) Cooper, 1894: 47, pl. 2, figs. 38

39; CASG 612. Vasculum? capuloides (C Jooper,

1894), late Campanian to possibly early Maas-

trichtian, CA. Coan (1981: 160).

cretaceum (Sistrum (Ricinula?)) Cooper, 1896: 330, pl. 47,

figs. 1, 2; neotype LACMIP 9997. Perissitys cretacea

(Cooper, 1896), Coniacian, CA. Popenoe and Saul (1987: 12).

fairbanksi (Cerithium) Cooper, 1894: 44, pl. 1, fig. 12; not

in GASG database. Possibly Anchura, late C: ampanian

to possibly early Maastrichtian, CA. Coan (1981: 163).

kempiana ( (Calliostoma) C Jooper, 1894: 46 [pl. 2 [not 3],

figs. 33, 34; CASG 610. Calliostoma kempianum

Cooper, 1894, late Campanian to possibly early

Maastrichtian, CA. Coan (1981: 165).

normalis (Tornatella) Cooper, 1894: 46, pl. 2 [not 3], figs.

36, 37; CASG 625. Eoacteon normalis (Cooper,

1894), late Campanian to possibly early Maas-

trichtian, CA. Coan (1981: 168).

DAILEY and POPENOE

Bivalves

apletos (Glycymeris) Daily and Popenoe, 1966: 8, pl. 1

figs. 1, 5, 6; LACMIP 8891. Glycymerita veatchii

(Gabb, 1864), middle Turonian to late Campanian, VI,

CA. Squires (2010: 902).

pozo (Corbula) Daily and Popenoe, 1966:19, pl. 5, figs. 6-10;

LACMIP 8916. Panzacorbula pozo (Daily and Pope-

noe, 1966), early Campanian to early late Maastrichtian,

CA. Squires and Saul (2004¢: 117).

Gastropod

ainiktos (Pseudoglauconia?) Daily and Popenoe, 1966: 21,

pl. 6, figs. 3, 5, 6; LACMIP 8291. Bullamirifica

ainiktos (Dailey and Popenoe, 1966), middle through

late Campanian, CA, BX. Squires and Saul (2005: 141).

HALL and AMBROSE

Gastropod

branneri (Cerithium) Hall and Ambrose, 1916: 70, unfigured;

CASG 69804. Cerithium? teslaensis Hanna, 1924-162,

Late Cretaceous, CA. See Wiedey (1929: 25, pl. 1, fig. 6).

McLELLAN

Bivalves

suciensis (Cucullaea) McLellan, 1927: 132, pl. 17, figs.

4-6; UWBM 15010, Glycymerita veatchii (Gabb,

1984), middle Turonian to late Campanian, VI, WA,

CA, BX. Squires (2010: 902).

suciensis (Glycymeris ) McLellan, 1927: 131, pl. 17, figs. 7,

8; UWBM_ 15008, Glycymerita veatchii (Gabb,

1864), middle Turonian to late Campanian, VI, WA,

CA, BX. Squires (2010: 902).

MERRIAM

Gastropod

tolenasensis (Turritella) Merriam, 1941: 62, pl. 1, figs. 14,

15: UCMP 15328. Turritella hearni Merriam, 1941,

Turonian and probably early Coniacian, CA. Squires

and Saul (2006a: 54).

MURPHY AND RODDA

Gastropods

allisoni (Gyrodes) Murphy and Rodda, 1960:842, pl. 101,

figs. 18-20; LACMIP 9828. Natica? allisoni (Murphy

and Rodda, 1960), Cenomanian, OR, CA. Popenoe

et al. (1987: 92).

greeni (Gyrodes) Murphy and Rodda, 1960:543, pl. 101,

figs. 27-29; LACMIP 9830. Gyrodes (Sohlella?)

greeni Murphy and Rodda, 1960, Cenomanian, CA.

Popenoe et al. (1987: 79).

stewarti (Sollariella) [sic] Murphy and Rodda, 1960:839,

pl. 103, figs. 4, 5; LACMIP 9821. Igonoia stewarti

(Murphy and Rodda, 1960), late Cenomanian, CA.

Squires (201la: 142).

PACKARD

Bivalves

alisoensis (Tellina) Packard (1922: 426, pl. 33, fig. 3);

UCMP. 12309. Laternula? alisoensis (Packard,

R.L. Squires, 2018

Page 95

1922), middle Campanian, CA. Elder and Saul (1993:

pole 2, ress, IL)

angulata (Meretrix) Packard (1922: 425, pl. 33, fig.

UCMP 12307. Etea angulata (Packard, 1

Campanian, CA. Saul (1982: 73).

californica (Exogyra) Packard, 1922: 421, pl. 27, fig. 5;

UCMP. 12320. Costagyra californica (Packard,

1922), Turonian, CA. Squires (2017: 39).

californiana (Trigonia) Packard, 1921: 17, pl. 2, fig. 2; UO

6 (lost, but CASG 5658 is a plaster cast of it). Popenoella

californiana (Packard, 1921), Turonian, CA. Saul

(1978: 39), Cooper and Leanza (2017: 329).

chicoensis (Spisula) Packard, 1916: 300, pl. 27, figs. 6, 7;

UCMP 12205. Willimactra (Petromactra) truncata

(Gabb, 1864), early to late Campanian, VI, CA. Saul

(G72 LS).

cordiformis (Cucullaea?) Packard, 1922: 417, pl. 24, fig. 1;

UCMP 12311. Cucullaea (Idonearca) Condiformis

Packard, 1922, late Campanian, CA. Saul (1982: 68).

coronaensis (Cardium) Packard, 1922: 424, pl. 30, fig. 2:

UCMP 12281. Pachycardium coronaense (Packard,

1922), Turonian, CA. Popenoe (1937: 388).

crescentica (Ostrea) Packard, 1922: 420, pl. 26, fig. 3 [not

fig. 4]; UCMP 12318. Curvostrea crescentica (Packard,

1922), Turonian, CA. Squires (2017: 29).

demessa haidana var. (Acila (Truncacila)) Packard in

Schenck, 1936: 50, pl. 2, figs. 3, 4, 6, 10; CASG 69081.

Acila (Truncacila) haidana Packard in Schenck,

latest Albian (probably) to early Turonian, QC, CA.

(Squires and Saul, 2006b: 92).

deschutesensis (Trigonia) Packard, 1921: 24, pl. 10, fig. 3;

UO 9. Notoscabrotrigonia oregana (Packard).

middle Albian?, perhaps middle or late Cenomanian,

OR. Jones (1960: 437), Cooper (2015: 25).

evansana var. oregona (Trigonia) Packard, 1921: 26, pl. 9,

fig. 7; UO 4. Notoscarbrotrigona oregano (Packard),

middle Albian to perhaps middle or late Cenomanian,

OR. Jones (1960: 437), Cooper (2015: 25).

fitchi (Trigonia) Packard, 1921: 20, pl. 6, fig. 3; pl. 7, fig. 2;

UO 26859. Louella fitchi ( (Packard, i991), early ie

middle Turonian, OR, CA. Saul (1978: 53), Cooper and

Leanza (2017: 330).

hardingensis (Homomya) Packard, 1922: 423, pl. 32, figs.

la, Lb: UCMP 12291. Liopistha (Psilomya) hardingensis

(Packard, 1922), Turonian, CA. Popenoe (1937: 383).

inezana (Trigonia) Packard, 1921: 27, pl. §, figs. la, b; pl. 9,

fig. 1, pl. 10, fig. 1; UCMP 31464. Notoscabrotrigona

evansana (Meek, 1858), Coniacian through Campa-

nian, VI, OR, CA. Jones (1960: 436), Cooper (2015: 25).

inornata (Exogyra) Packard, 1922: 420, al, Bi, img, lly

UCMP 12284. Phygraea inornata (Packard, 1922),

Turonian, CA. Squires (2017: 46).

lapidis (Astarte) Packard, 1922: 423, pl. 30, figs. 4a, 4b;

UCMP. 12285. Eriphyla lapidis (Packard, 1922),

Campanian? CA. Popenoe (1937: 387).

nitida (Meretrix) var. major Packard, 1922: 425, pl. 33, fig.

2; UCMP 12279. Calva (Egelicalva) Doemana

(Cooper, 1894), late early through late Campanian,

WA, CA. Saul and Popenoe (1992: 39).

5);

922).

ovoides (Astarte) Packard, 1922, 424: pl. 30, fig. 1; UCMP

12280. Eriphyla ovoides (Packard, 1922), late

Turonian to Coniacian’, CA. Popenoe (1937: 386), Saul

(1982: 72).

striatus (Spondylus) Packard, 1922: 422, pl. 29; UCMP

12276, [a junior homonym renamed as Spondylus

fucatus Hanna, 1924: 181], late C Jampanian to early

Maastrichtian, CA. Elder (1991: E10).

subnodosa (Lima) Packard, 1922:421, pl. 28; UCMP

12275. Spondylus subnodosus (Packard, 1922), late

Campanian to early Maastrichtian, CA, BX? Elder

(1991: E10).

sulcata (Astarte?) Packard, 1922: 424, pl. 33, fig. 6; UCMP

12305, [a junior homonym renamed as Astarte? acerba

Hanna, 1924: 157 and as Astarte? earllergyi Anderson,

1958: 121: now Alleinacin acerba (Hanna, 1924)].

Turonian, CA. Squires and Ritterbush (1981: 896).

taxidonta (Ostrea) Packard, 1922: 420, pl. 26, fig. 2:

UCMP. 12317. Acutostrea taxidonta (Packard,

1922), Turonian, CA. Squires (2017: 32).

Gastropods

californica (Gyrodes) Packard, 1922: 429, pl. 35, figs. 2a,

2b; UCMP 12300. Euspira Shuntardiana (Gabb,

1864), late Campanian, CA. Pope noe (1937: 398).

californiensis (Lysis) Packard, 1922: 431, pl. 37, figs. 2

UCMP 12287. Lysis suciensis (Whiteaves, eieza)

late early Campanian to early Maastrichtian, VI, WA,

CA, BX. Saul and Squires (2008b: 128).

dubius (Siphonalia) Packard, 1922: 431, pl. 35, fig. 5;

UCMP. 12304. Saturnus dubius (Packard, 1922),

Turonian, CA. Saul and Popenoe (1993: 367).

nodosa (Alaria) Packard, 1922: 430, pl. 26, figs. 5a, 5b;

UCMP. 12297. Latiala nodosa (Packard, 1922),

Turonian, CA. Saul (1998: 132).

pseudoalveata (Amauropsis) Packard, 1922: 429, not pl.

35, figs. la, 1b, 3; UCMP 12301. Ampullina packardi

Popenoe, 1937, early to middle Campanian, CA.

Popenoe (1937: 399), Stecheson (2004).

suciaensis (Cerithium?) Packard, 1922: 430, pl. 35, fig. 4

UCMP. 12303. Zebalia suciaensis (Packard, 1922),

middle Campanian, WA, CA. Squires and Saul (2003a: 433).

vetus (Aporrhais) Packard, 1922: 431, pl. 36, fig. 1; UCMP

12298. Alarimella veta (Packard, 1922), Turonian,

CA. Saul (1998: 134).

tumida (Bullaria) Packard, 1922: 433, pl. 37, fig. 4, UCMP

12289, [a junior homonym renamed as Bullaria

obtenta Hanna, 1924: 159], Turonian, CA. Saul

(1982: 72).

WARING

Bivalves

chicoensis (Isocardia) Waring, 1917: 62, pl. 8, fig. 3; LSJU 399.

pace Clisocolus cordatus Whiteaves, 1879: 157, middle

Campanian, CA. Popenoe (1937: 390); Smith, 1978: 332).

cordata (Macrocallista) Waring, 1917: 62, pl. 8, fig. 1; not

in CASG database. Paraesa? lens (Gabb, 1864),

Campanian, VI, WA, CA. Saul (1993: 976).

Page 96

THE NAUTILUS, Vol. 132, No. 3-4

cowpert ( (Pecten (Propeamusium)) [sic] Waring, 1917: 63,

pl. 7, figs. 1, 2; ;: syntypes CASG 61644. Propeamussium

(Parpanrussiiin) cowperi (Waring, 1917), late

middle Campanian, CA. Sundberg (1989: 61-62). A

“variety” of cowperi occurs in a Paleocene wood-fall

(chemosynthetic) deposit in northern Japan (Amano

et al., 2018: 636).

triangulatus (Crassatellites) Waring, 1917: 59, pl. 9, fig. 1;

syntype CASG 397. Cymbophora _ triangulata

(Waring, 1917), late Campanian, CA. Saul (1974: 1083).

Gastropods

compressus (Gyrodes) Waring, 1917: 67, pl. 9, fig. 6;

CASG 61958.01. Euspira compressa (Waring,

1917), middle Campanian, CA. Stecheson (2004: 64).

crassa (Cancellaria) Waring, 1917: 66, pl. 9, fig. 5; CASG

61644.06. [a junior orasranaa renamed as Gincellann

simiana Hanna, 1924: 160], probably middle Cam-

panian, CA. No age update found.

plicata (Turris) Waring, 1917: 68, pl. 9, fig. 3; not in CASG

database, [a junior homonym renamed as Surcula

binotata Hanna, 1924: 181], Late Cretaceous un-

differentiated, CA. No age update found.

rotundus (Pugnellus ) Waring, 1917: 67, pl. 9, fig. 10; LS]U

402. Lispodesthes rotundus (Waring, 1917), middle

Campanian to early Maastrichtian, CA. Dailey and

Popenoe (1966: 22), Stecheson (2004: 43).

templetoni (Solariaxis) Waring, 1917: 68, pl. 9, fig. 22; not

in CASG database. Atira ornatissima (Gabb, 1864),

latest Santonian to late Campanian, VI, CA, BX. Squires

(2010: 1025).

WHITE

Gastropods

californiensis (Nerita) Orcutt, 1887 (based on White’s,

1885: 12, pl. 5, figs. 7, 8 of Nerita sp.); lectotype USNM

1341 la. Nerita (Bajanerita) californiensis (Orcutt,

1887), late Campanian to early Maastrichtian, BX.

Squires (1993: 1085).

condoni (Trophon) White, 1889: 21, pl. 3, figs. 4, 5;

syntypes USNM = 20122. Praesargana condoni

(White, 1889), Turonian, CA. Saul and Popenoe

(1993: 360).

dilleri (Scobinella) White, 1889: 25, pl. 4, figs. 1-3; syn-

types USNM 20123. Carota dilleri (White, 1889).

Turonian, VI, OR, CA, BX. Saul and Popenoe

(1993: 374).

dowelli (Gyrodes) White, 1889: 19, pl. 3, figs. 8, 9;

USNM 20126. Gyrodes (Gyrodes) dowelli White,

1889, Turonian, VI, OR, CA. Popenoe et al.

(1987: 75).

euryostomous (Trochus (Oxystele)) White, 1885: 12, pl. 5,

figs. 9-11; not in USNM online database. Homalo-

poma euryostoma (White, 1885), early Campanian to

early Maastrichtian, BX. Kiel and Aranda-Manteca

(2002: 29).

gabbi (Fulguraria) White, 1889: 23, pl. 3, fig. 1; lec-

totype USNM 20112. Volutoderma averillii (Gabb,

1864), early Campanian, VI, CA. Saul and Squires

(2008a: 226).

hilgardi (Fulgur) White, 1889: 22, pl. 3, figs. 2, 3; USNM

20117. Pentzia hilgardi (White, 1899). early Cam-

panian to early Maastrichtian, WA, CA, BX. Squires and

Saul (2003b: 57): Stecheson (2004: 83).

marcidulus (Faunus) White, 1889: 20, pl. 4, figs. 12, 13;

USNM 20127. Liocium marcidulum (White,

1889), early Campanian, CA. Squires and Saul

(2003c: 149).

nexilia (Ceratia) White, 1889: 21, pl. 3, figs. 13, 14; lec-

totype USNM 20119a. Acirsa nexilia (White, 1889),

early to late middle Campanian, WA, CA. Squires and

Saul (2003a: 39).

obstricta (Stomatia) White, 1889: 18, pl. 4, figs. 10, 11;

USNM 20124. Ariadnaria obstricta (White, 1889),

late Coniacian? to Santonian, CA. Saul and Squires

(2008b: 122).

obtusa (Mesalia) White, 1889: 20, pl. 4, figs. 6, 7]; lec-

totype USNM 20116a. Acirsa obtusa (White, 1889).

late Santonian to early late Campanian, CA. Squires and

Saul (2003a: 36).

oppansus (Lysis) White, 1889: 17, pl. 4, figs. 14, 15;

USNM 20115. Lysis duplicosta Gabb, 1864, Cam-

panian, WA, CA. Saul and Squires (2008b: 125).

pillingi (Cerithium) White, 1885: 13, pl. 5, figs. 3-6;

USNM 13408. Echinoaxis pillingi (White, 1885),

early Campanian to early Maastrichtian, BX. Kiel and

Aranda-Manteca (2002: 33).

totium-sanctorum (Cerithium) White, 1885: 13, pl. 5, figs.

12, 13; USNM 13409. Tympanotonos (Exechocirsus)

totiumsanctorus (White, 1885), early Campanian to

early Maastrichtian, BX. Kiel and Aranda-Manteca

(2002: 34).

wallaense (Solarium) White, 1885: 14, pl. 5, figs. 1, 2;

USNM_ 13412 Trochacanthus wallalense (White,

1885), early Campanian to early Maastrichtian, CA,

BX. Squires (201 1b: 3).

WHITEAVES

Bivalves

cretacea (Conchocele) Whiteaves, 1874: 266, plate of

fossils (figs. 2, 2a); vouchers CASG 61850.04, 61850.5.

Thyasira cretacea (Whiteaves, 1874), exact age

unknown, VI. Whiteaves (1903: 383).

cumshewaensis (Arca (Nemodon)), Whiteaves (1900: 294:

illustrated by Whiteaves 1884: 235, pl. 31, figs. 8, 8a,

8b): lectotype GSC 4915 (see Smith, 1978: 335).

Navonavis cusmshewaensis (Whiteaves, 1900), late

Santonian, QC. Haggart and Higgs (1989: 61).

meekana (Te Mina) Whiteaves, 1874: 268, plate of fossils

a6 6); GSC 5730. Paraesa? lens (Gabb, 1864),

Campanian, V 7 WA, CA. Saul (1993: 976).

nanaimoensis (Tellina) Whiteaves, 1903: 376, pl. 46, fig. 3;

GSC 5729. Willimactra (Petromactra) truncata

(Gabb, 1864), early to late Campanian, VI, CA. Saul

(1973: 23).

R.L. Squires, 2018

Page 97

suciense (Laevicardium) Whiteaves, 1879: 154, pl. 18, fig.

2; GSC 5713. Cymbophora suciensis (Whiteaves,

1879), Campanian, WA, CA. Saul (1974: 1079) used an

unnecessary emendation of Whiteaves’ name "suciense,"

instead of using the name "suciensis."

suciensis (Linearia) Whiteaves, 1879: 146, pl. 17, fig. 12;

GSC 5824 (Bolten, 1992: 235). No new name found,

hinge characters are unknown, and the only known

ena n has been lost (Squires and Goede srt, 1994:

262); thus, this bivalve is a nomen dubium, early middle

Campanian, WA. Squires and Graham (2014: fig. 2).

suciensis (Teredo) Whiteaves, 1879: 135, pl. 17, figs. 1, la;

syntypes GSC 5752, 5752 a-d (Bolton, 1992: 239). No new

name found, but this fossil cannot be assigned to “Teredo,”

whose certain identification requires soft-part morphology,

early late Campanian, VI. Squires and Graham (2014: fig. 2).

vancouverensis (Opis) Whiteaves, 1879: 158, pl. 18, figs. 4,

4a; GSC 5691. Opis (Hesperopis) vancouverensis

Whiteaves, 1879, middle to late Campanian, VI.

Squires and Saul (2009: 1328).

Gastropods

canadensis (Gyrodes) conradiana? var. Whiteaves, 1903:

365, unfigured; GSC 5777. Gyrodes (Sohlella) can-

adensis Whiteaves, 1903, Santonian to earliest Campanian,

VI. Popenoe et al. (1987: 85).

dakotensis vancouverensis var. (Serrifusus) Whiteaves

1879: 119, pl. 15, fig. 6. GSC 5794 (Bolton: 1965:

74). Serrifusus vancouverensis Whiteaves, 1879.

Boeebly early Maastrichtian, VI. Anderson (1958: 171).

harveyi (Cerithium) Whiteaves, 1903: 362, pl. 43, fig. 7;

syntypes 5933, 5933a (Bolton, 1965: 11). Alamirifica?

oe (Whiteaves, 1903), Coniacian to early

Campanian, VI. Saul and Squires (2003: 448).

intermedium (Mesotoma?) Whiteaves, 1903:360, pl. 43,

fig. 4; syntypes 5956, 5956 a-d (Bolton, 1965: 44).

Acirsa nexilia (White, 1889), early Campanian to late

middle Campanian, WA. Squires and Saul (2003a: 39).

lallierianum suciense var. (Cerithium) Whiteaves, 1879:

122, pl. 15, figs. 10, 10a; lectotype GSC 5764b. Bel-

liscala suciense (Whiteaves, 1903), middle to early

late Campanian, WA. Squires and Saul (2003a: 33).

newcombii (Mesostoma?) Whiteaves, 1903: 361, pl. 43, fig.

5; GSC 5298 (Bolton, 1965: 45). Confusiscala new-

combii (Whiteaves, 1903), latest Santonian to middle

Campanian, WA. Squires and Saul (2003a: 40).

nodulosa (Fasciolaria) Whiteaves, 1874: 268, figs. 7, 7a

(not 7b); lectotype GSC 5766. Forsia popenoei Saul,

1988. Late early to middle Campanian, VI, CA. Saul

(1988: 10), Squires and Graham (2014: fig. 2).

occidentalis radiatula? var. (Solariella) Whiteaves, 1903:

368, pl. 45, figs. 5, 5a; GSC 5918. Igonoia occidentalis

(Whiteaves, 1903), Santonian. Squires (201 1a: 144).

suciense (Mesostoma) Whiteaves, 1903: 359, pl. 44, fig. q

lectotype GS 5764b. Belliscala suciense (Whiteaves,

1879), middle Campanian to early late Campanian,

WA. Squires and Saul (2003a: 33).

suciensis (Cypraea) Whiteaves, 1895: 127, pl. 3, fig. 5; GSC

5937. Palaeocypraea suciensis (Whiteaves, 1895),

early middle

Squires and Graham (

Sepa WA. Groves (1990: 275):

2014: fig. 2); Lorenz, 2017: 210).

suciensis (Surcula) Whiteaves, 1879: 115, pl. 15, figs. 1, la

syntypes GSC 5784, 5784 a—b (Bolton, 1965: 79).

Amuletum? (Lutema)? suciensis (Whiteaves, 1879),

early middle Campanian, WA. Erickson (1974: 223),

Squires and Graham (2014: fig. 2).

suciensis carinifera var. (Stomatia) Whiteaves, 1879: 128,

pl. 16, fig. 4; lectotype GSC 5771. Lysis suciensis

(Whiteaves, 1879), late early Campanian to early

Maastrichtian, VI, WA, CA, BX (including Baja Sur).

Saul and Squires (2008b: 128).

suciensis carinifera var. (Stomatia) Whiteaves, 1879: 128,

pl. 16, fig. 5; lectotype UCMP 11975. Lysis duplicosta

Gabb, 1864, early to late Campanian, WA, no. CA.

Saul and Squires (2008b: 125).

tenuis nanaimoensis var. (Potamides) Whiteaves, 1879:

121, pl. 15, figs. 9, 9a; lectotype GSC 5763a. Anchura

nanaimoensis (Whiteaves, 1879), middle to late

Campanian, VI. Elder and Saul (1996: 390).

ACKNOWLEDGMENTS

Peter Roopnarine and Christine Garcia provided CASG

type-specimen numbers. Lindsey T. Groves (LACM,

Malacology Department) provided detailed information.

Lorenz (201 7), provided some type-specimen numbers,

critically reviewed the manuscript, and gave valuable

comments about it.

LITERATURE CITED

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

Mesozoic-relict mollusks from Paleocene wood-fall com-

munities in Urahoro Town, eastern Hokkaido, northerm

Japan. Journal of Paleontology 92: 634-637.

Anderson, F.M. 1902. Cretaceous deposits on the Pacific coast.

Proceedings of the California Academy of Sciences, 3rd

series, 2: 1-154.

Anderson, F.M. 1958. Upper Cretaceous of the Pacific coast.

The Geological Society of America Memoir 17, 378 pp.

Anderson, F.M. and G.D. Hanna. 1935. Cretaceous geology of

Lower California. Proceedings of the California Academy of

Sciences, 4th series, 23: 1-34.

Amold, R. 1908. Descriptions of new Cretaceous and Tertiary

fossils from the Santa Cruz Mountains, California. Pro-

ceedings of the United States National Museum 34:

345-390.

Bolton, T.E. 1965. Catalogue of type invertebrate fossils of the

Geological Survey of Canada. Geological Survey of Canada.

Volume 2, 344 pp.

Bolton, T.E. 1992. Catalogue of type invertebrate fossils of the

Geological Survey of Canada. Geological Survey of Canada.

Volume 8, 424 pp.

Coan, E. 1981. James Graham Cooper, pioneer western natu-

ralist. Northwest Naturalist Books, The University Press of

Idaho, Moscow, Idaho, 255 pp.

Cooper, J.G. 1894. Catalogue of Californian fossils (parts 2—5).

California State Mining Bureau, Bulletin 4: 1-65.

Page 96

THE NAUTILUS, Vol. 132; No. 3=4

Cooper, ].G. 1896. On some new Cretaceous (and Eocene?)

Mollusca of California. Proceedings of the California

Academy of Sciences 6: 330-337.

Cooper, M. R. 2015. On the Pte rotrigonia (Bivalvia, Trigoniida):

their biogeography, evolution, *“deasttention and relation-

ships. Neues Jahrbuch fiir Geologie und Palaontologie

Abhandlungen 277: 11-42.

Cooper, M.R. and H.A. Leanza. 2017. On the Steinmanellidae

(Bivalvia: Myophorelloidea); their palaeobiogeography,

evolution and classification. Neus Jahrbuch fiir Geologie

und Palaontologie Abhandlungen 285: 313-335.

Crampton, J., T. Mumme, I. Raine, L. Roncaglia, P. Schigler, P.

Strong, G. Turner, and G. Wilson. 2000. Revision of the

Piripauan and Haumurian local stages and correlation of the

Santonian-Maastricthian (Late Cretaceous) in New Zea-

land. New Zealand Journal of Geology and Geophysics 43:

309-333.

Dailey, D.H. and W.P. Popenoe. 1966. Mollusca from the Upper

Cretaceous Jalama Formation, Santa Barbara County,

California. University of California Publications in Geo-

logical Sciences 65: 1-27.

Elder, W.P. 1991. An unusual Late Cretaceous fauna form an

oyster-rich interval in the Santa Cruz Mountains of Cal-

ifornia. U.S. Geological Survey Bulletin 1934, Chapter E,

E1-E15.

Elder, W.P. and L.R. Saul. 1993. Paleogeographic implications

of molluscan assemblages in the Upper Cretaceous

(Campanian) Pigeon Point Formation, California. In:

Dunne, G., and K. McDougall (eds.). Mesozoic Paleoge-

ography of the Western United States-2. Pacific Section

SEPM, Book 71: 171-186.

Elder, W. and L.R. Saul. 1996. Taxonomy and biostratigraphy of

Coniacian through Maastrichtian Anchura (Gastropoda:

Aporrhaiidae) of the North American Pacific slope. Journal

of Paleontology 70: 381-399.

Erickson, J.M. 1974. Revision of the € Gastropoda of the Fox

Hills Formation, Upper Cretaceous (Maestrichtian) of

North Dakota. Bulletins of American Paleontology 66:

ISIE Zass

Gabb, W.M. 1864. Description of the Cretaceous fossils.

Geological Survey of California, Palaeontology 1(4):

57-217.

Gabb, W.M. 1876. Notes on American Cretaceous fossils with

descriptions of some new species. Proceedings of the

Academy of Natural Sciences of Philadelphia for 1876, pp.

276-324.

Gabb, W.M. 1877. Notes on American Cretaceous fossils, with

descriptions of some new species. Proceedings of the

Academy of Natural Sciences of Philadelphia 28: 276-324.

Goldfuss, G.A. 1836. Petrefacia Germaniae. Arnaz and Co.

Diisseldorf, 312 pp.

Groves, L.T. 1990. New species of Late Cretaceous Cypraeaca

(Mollusca: Gastropoda) from California and Mississippi,

and a review of Cretaceous cypraeaceans of North America.

The Meee 33: 272-285.

Groves, L.1T

the fossil invertebrates described by William More Gabb

(1839-1878). Zootaxa, 149 pp.

Haggart, J.W. and R. Higgs. 1989. A new Late Cretaceous

mollusc fauna from the Queen Charlotte Islands, British

Columbia. Geological Survey of Canada, Paper 89-1H:

59-64.

Hall, E.B. and A.W. Ambrose. 1916. Descriptions of new species

from the Cretaceous and Tertiary of the Tesla, Pleasanton,

_ and R.L. Squires. In press. Annotated Catalog of

San Jose, and Mt. Hamilton quadrangles, California. The

Nautilus 30: 68-71, 77-82.

Hanna, G.D. 1924. Rectifications of nomenclature. Proceedings

of the California Academy of Sciences, 4'” series, 13:

151-186.

Jones, D.L. 1960. Pelecypods of the genus Pterotrigonia from

the west coast of North America. Journal of Paleontology

34: 433-439.

Kauffman, E.G. 1973. Cretaceous Bivalvia. In, A. Hallam (ed.).

Atlas of palaeobiogeography. Elsevier Scientific Publishing

Company, Amsterdam, The Netherlands: 353-383.

Kauffman, E.G. 1977. Systematic, biostratigraphic, and_bio-

geographic relationships between middle Cretaceous

Buramerican’ and) north Pacific Inoceramidae, | Palae:

ontological Society of Japan Specie Papers 21: 169-212.

Kiel, S. andl Del). Aranda-Manteca. 2002. The g gastropod at the

Coralliochama beds on Punta Banda (Late Cretaceous,

northwestern Mexico) and a reconstruction of their pale-

oenvironment. Mitteilungen aus dem Geologisch-

Palaontologisches Institut Universitét Hamburg 86: 25-36.

Lorenz, F. 2017. Cowries a guide to the gastropod family

Cypraeidae. Volume 1. Biology and Systematics. Con-

chBooks, Harxheim, Germany. 644 pp.

Mantell, G. 1822. Fossils of the South Downs, or illustrations of

the geology of Sussex. London. 320 pp.

Mcl Alen, R.D. 1927. The geology of the San Juan Islands.

University of W ashington Phikeatons’ in Geology 2: 1-185.

Meek, F.B. 1858. Descriptions of new organic remains of fossils

from Vancouver's Island. Tenancies of the Albany In-

stitute 4: 37-49.

Merriam, C.W. 1941. Fossil turritellas from the Pacific coast

region of North America. University of California Publi-

cations Bulletin of the Department of Geological Sciences

26: 1-214.

Miller, V.V. and P.L. Abbott. 1989. In: Abbott, P.L. (ed.).

eae of the Upper Cretaceous Rosario Forma-

tion near Calonct and Camalu, Baja California. Geologic

studies in Baja California. Pacific Section, Society of

Economic Paleontologists and Mineralogists. Book 63. Los

Angeles, California: 63-74.

Murphy, M.A. and P.U. Rodda. 1960. Mollusca of the Creta-

ceous Bald Hills Formation of Califormia. Journal of Pa-

leontology 34: 835-858.

Orcutt, C.R. 1887. New Cretaceous fossils. The West American

Scientist 3: 28-31.

Packard, E.L. 1916. Mesozoic and Cenozoic Mactrinae of the

Pacific Coast of North America. University of California

Publications, Bulletin Department of Geology 9: 261-360.

Packard, E.L. 1921. The Trigoniae from the Bacife coast of

North America. University of Oregon Publications 1: 1-58.

Packard, E.L. 1922. New species from the Cretaceous of the

Santa Ana Mountains, California. University of California

Publications Bulletin of the Department of Geological

Sciences 13: 413-462.

Popenoe, W.P. 1937. Upper Cretaceous Mollusca of southem

California. leu of Paleontology 11: 370-402.

Popenoe, W.P. 1957. The Cretaceous gastropod genus Biplica

its evolution an biostratigraphic significance. University of

California Publications in Geological Sciences 30:425-454.

Popenoe, W.P. and L.R. Saul. 1987. Evolution and classification

of the Late Cretaceous-early Tertiary gastropod Perissitys.

Contributions in Science 380: 1—37.

Popenoe, W.P., R.W. Imlay, and M.A. Murphy. 1960. Corre-

lation of the Cretaceous formations of the Pacific coast

R.L. Squires, 2018

Page 99

(United States and northwestern Mexico). Bulletin of the

Geological ak ty of Ame rica 71: 1491-1540.

Popenoe, W.P., L.R. Saul, and T. Susuki. 1987. ¢

gastropods a the Pacific eons Cretaceous and Paleo-

cene. Journal of Paleontology 61: 70-100.

Saul, L.R. 1973. Evidence for the origin of the Mactridae

(Bivalvia) in the Cretaceous. University of California

Publications in Geological Sciences 97: 1-51.

Gyrodiform

Saul, L.R. 1974. Described or figured west coast species of

Cymbophora. Journal of Paleontology 48: 1068-1095.

Saul, L.R. 1978. The north Pacific Gremecons trioniid genus

Yaadia. University of California Publications in Ge ological

Sciences 119: 1—65.

Saul, L.R. 1982. Water depth indications from Late Cretaceous

mollusks, Santa Ana Mountains, California. In, D.J. Bottjer,

I.P. Colburn, and J.D. Cooper (eds.). Late Cretaceous

depositional environments and paleogeography, Santa Ana

Mountains, southern California. Pacific Section, SEPM,

Field Trip Volume and Guidebook: 69-76.

Saul, L.R. 1983. Turritella zonation across the Cretaceous-

Tertiary boundary, California. University of California

Publications ¢ Geological Sciences 125: 1-165.

Saul, L.R. 1988. New Late Cretaceous and early Tertiary Per-

issityidae (Gastropoda) from the Pacific slope of North

America. Contributions in Science 400: 1—25.

Saul, L.R. 1993. Pacific slope Cretaceous bivalves: eight venerid

species. Journal of Paleontology 67: 965-979.

Saul, L.R. 1998. Eight aporrhaid gastropod species from the

Cretaceous of the Pacific slope of North America and

clarification of the type species of Perissoptera. The Nau-

tilus 111: 119-142.

Saul, L.R. and W.P. Popenoe. 1992. Pacific slope Cretaceous

bivalves of the genus Calva. Contributions in Science 433,

68 pp.

Saul, L.R. and W.P. 1993. Additions to Pacific slope Turonian

Gastropoda. The Veliger 36: 351-388.

Saul, L.R. and R.L. Squires. 1998. New Sree Gastropoda

from California. Palaeontology 41(3): 461-488.

Saul, L.R. and R.L. Squires. 2003. New é retaceous cerithiform

gastropods from the Pacific slope of North America. Journal

of Paleontology 77: 442-453.

Saul, L.R. and R.L. Sguires. 2008a. Volutoderminae (Gastro-

poda): Volutidae) of Coniacian through Maastrichtian age

from the North American Pacific slope. Journal of Pale-

ontology 82: 213-237.

Saul, L.R. and R.L. Squires. 2008b. Cretaceous trichotropid

gastropods from the Pacific slope of North America: Pos-

sible pathways to calyptraeid morphology. The Nautilus

122: 115-142.

Saul, L.R, T. Susuki, and C.A. Nelson. 1989. Willis Parkison

Popenoe IIT, geology: Los Angeles. [http://texts.cdlib.org/view?

docld=hb4p30063r, NAAN = 13030&doc.view= frames&chunk.

id= div044&toc.depth = 1 &toc.id= &brand=calisphere |

Schenck, H.G. 1936. Nuculid bivalves of the genus Acila.

Geological Society of America Special Papers 4, 149 pp.

Seitz, O. 1934. Die Variabilitaét des Inoceramus labiatus v.

Schloth. Jahrbuch der Preussischen geologischen Land-

essanstalt zu Berlin. 55: 429-474.

Smith, J.T. 1978. Primary types in the Stanford paleontological

type collection. Bulletins of American P.: aleontology 72:

313-552.

Sohl, N.F. and H.A. Kollmann. 1985. Cretaceous actaeonellid

gastropods from the Western Hemisphere. U. S. Geological

Survey Professional Paper 1304: 1-95.

Squires, R.L. 1993. A new subgenus of neritid gastropod from

the Upper Cretaceous of Baja California, Mexico. Journal of

Paleontology 67: 1085-1088.

Squires, R.L. 9010. Northeast Pacific Upper Cretaceous and

Paleocene glycymeridid bivalves. Journal of Paleontology

84: 895-917.

Squires, R.L. 201 la. A new genus of Cretaceous margaritine

gastropod (Turbinidae) from the northeastern Pacific

Ocean. The Nautilus 125: 137-149.

Squires, R.L. 2011b. New Cretaceous turbiniform vetigastro-

pods (Gastropoda) from the Pacific slope of North America.

The Nautilus 125: 1-14.

Squires, R.L. 2014. Cretaceous and Paleogene Pteria bivalves

from the Pacific slope of North America. The Veliger 51:

216-23.

Squires, R.L. 2017. Late Cretaceous oysters from the Pacific

slope of North America: revision of named species and

discove Ty of new species. Contributions in Science 525:

25S"

Squires, R.L. 2018. Late Cretaceous endemic shallow-marine

gastropod genera of the northeast Pacific: biodiversity and

faunal changes. PaleoBios 35: 1-16.

Squires, R.L. and J.L. Goedert. 1994. New species of early

Eocene small to minute mollusks from the Crescent For-

mation, Black Hills, southwestern Washington. The Veliger

37: 253-266.

Squires, R.L. and R. Graham. 2014. Additions and refinements

to Sycodes (Shumard, 1858), a poorly known Late Creta-

ceous (Campanian) marine gastropod from the northeast

Pacific: taxonomic and biostratigray yhic implications. Ca-

nadian Journal of Earth Sciences 51: 775.782

Squires, R.L. and L. Ritterbush. 1981. A new astartid bivalve

genus from the Upper Cretaceous of southern California.

jonmel of Paleontology 55: 896-897.

Squires, R.L. and L.R. Saul. 2003a. New Late Cretraceous

epitoniid and zygopleurid gastropods from the Pacific slope

of North America. The Veliger 46: 20-49.

Squires, R.L. and L.R. Saul. 2003b. New Late Cretaceous

(Campanian and Maastrichtian) marine gastropods from

California. Journal of Paleontology 77: 50-63.

Squires, R.L. and L.R. Saul. 2003¢. editions to Late Cretaceous

shallow-marine gastropods from California. The Veliger 46:

145-161.

Squires, R.L. and L.R. Saul. 2004a. The pseudomelaniid gas-

tropod Paosia from the marine Cretaceous of the Pacific

slope of North America and a review of the age and

paleobiogeography of the genus. Journal of Paleontology

78: 484—500.

Squires, R.L. and L.R. Saul. 2004b. Uncommon Cretaceous

naticiform gastropods from the Pacific slope of North

America. The Veliger 47: 21-37.

Squires, R.L. and L.R. Saul. 2004c. Cretaceous corbulid bivalves

of the Pacific slope of North America. The Veliger 47:

103-129.

Squires, R.L. and L.R. Saul. 2005. New Late Cretaceous

(Santonian and Campanian) gastropods from California and

Baja Califomia, Mexico. The Nautilus 119: 133-148.

Squires, R.L. and L.R. Saul. 2006a. Additions and refinements to

Aptian to Santonian (Cretaceous) Turritella (Mollusca:

Gastropoda) from the Pacific slope of North America. The

Veliger 48: 46-60.

Squires, R.L. and L.R. Saul. 2006b. Cretaceous Acila (Trun-

cacila) (Bivalvia: Nuculidae) from the Pacific slope of North

America. The Veliger 48: 83-104.

Page 100

THE NAUTILUS, Vol. 132, No. 3-4

Sguires, R.L. and L.R. Saul. 2009. Cretaceous opine bivalves

from the Pacific slope of North America and_palae-

obiogeography of subfamily Opinae Chavan, 1969. Palae-

ontology 52(6), pp. 131 Ay,

Stecheson, M.S. 2004. Systematic paleontology of marine gas-

tropods from the Upper Cretaceous Chatsworth Forma-

tion, Simi Hills, southern California. California State

University Northridge, unpub. MLS. thesis. 142 pp. [http://

hdl handle.net/10211.3/164139]

Sundberg, F.A. 1989. Propeamussium species (Bivalvia: Pro-

peamussiidae) from the Upper Cretaceous of southern

California. Journal of Paleontology 63: 53-63.

Waring, C.A. 1917. Stratigraphic antl faunal relations of the

Martinez to the Chico and Tej jon of southern California.

Proceedings of the California Academy of Sciences, qth

eS 7: 41-124.

White, C.A. 1885. On new Cretaceous fossils from California.

U. e ‘Gosleste al Survey Bulletin 22: 352-372.

White, C.A. 1889. On invertebrate fossils from the Pacific coast.

U. S. Geological Survey Bulletin 51: 1-102.

Whiteaves, ].F. 1874. Notes on the Cretaceous fossils collected

by Mr. James Richardson at Vancouver and the adjacent

islands. Canada Geological Survey, Report of Progress for

1872-1873, pp. 260-268.

Whiteaves, J.F. 1879. On the fossils of the Cretaceous rocks of

Vancouver and adjacent islands in the Strait of Georgia.

Geological Survey of Canada, Mesozoic Fossils, Part. 2.

Volume 1: 263-307.

Whiteaves, ].F. 1884. On the fossils of the coal-bearing deposits

of the Queen Charlotte Islands collected by Dr. G.M.

Davson in 1878. Geological Survey of Canada, Mesozoic

Fossils, Part 3. Volume 1: 191-262.

Whiteaves, J.F. 1895. On some fossils from the Nanaimo Group

of the Vancouver Cretaceous. Transactions of the Royal

Society of Canada, 2 94 series, 1: 119-133.

Whiteaves, ].F. 1900. On some additional or impertectly un-

derstood fossils from the Cretaceous rocks of the Queen

Charlotte Islands, with a revised list of the species from

these rocks. Mesozoic Fossils, Part 4. Volume 1: 263-307.

Whiteaves, J.F. 1903. On some additional fossils from the

Vancouver Cretaceous, with a revised list of the species

therefrom. Geological Survey of Canada, Mesozoic Fossils,

Part 5. Volume 1: 309-415.

Wiedey, L.W. 1929. Some previously unpublished figures of

type mollusks from California. The Nautilus 48: 21-26.

Woods, H. 1917. The Cretaceous faunas of the northeastern part

of the South Island of New Zealand. New Zealand Geo-

logical Survey Paleontological Bulletin 4: 1-41.

THE NAUTILUS 132(3-4):101-112, 2018

Page 10]

Anatomy of Engoniophos unicinctus from Isla Margarita, Venezuela

(Gastropoda: Caenogastropoda: Nassariidae), with a discussion on

the buccinid-nassariid relationship

Daniel Abbate

Luiz Ricardo L. Simone

Museu de Zoologia da Universidade de Sao Paulo

Sao Paulo, (4299- 970 BRAZIL

[email protected]

Daniel C. Cavallari

Centro para Documentagao da Biodiversidade

Faculdade de Filosofia, Ciéncias e Letras de Ribeirao Preto

Universidade de Sao Paulo

Ribeirado Preto, BRAZIL

[email protected]

ABSTRACT

The taxonomic position of Engoniophos unicinctus (Say, 1826) in

the family Nassariidae is comfirmed based on morphological

results. Our detailed anatomical analysis reveals some disparities

with typical nassariids, e.g., having a non-serrated operculum,

a simpler and proportionally smaller head, and the absence of

epipodial tentacles. However, the similarities in shell and radula

are present and are easily noticeable, which corroborates the

taxonomic data and the positioning of E. unicinctus within the

Nassariidae.

Additional Keywords: Anatomy, Buccinidae, Neogastropoda,

Taxonomy

INTRODUCTION

The families Nassariidae and Buccinidae are among the

most diverse within the Caenogastropoda. They occur

from the poles to the equator, inhabiting a wide varie ty

of environments, from shallow water to abyssal ocean

depths (Harasewych, 1998;

the families are small- to medium-sized (20-250 mm),

with ovate to fusiform, weakly shouldered shells, an open

siphonal canal, columella lacking plicae, and axial sculpture

usually consisting of broad aie limited to early whorls.

These families ewe a rich fossil record dating from the

lower Cretaceous (Taylor and Morris, 1980), together with

Fasciolariidae, Melongenidae, and Cancellariidae, while

other neogastropod families appeared between the late

Cretaceous and early Paleocene (Tracey et al., 1993).

The systematics of both families are poorly understood,

and there is no consensus as to their limits and their

relationships (Simone, 2011). Ponder (1974) considered

the Buccinidae closely related to the Nassariidae, Fas-

ciolariidae, and Melongenidae, based on the absence of

accessory salivary lands and an anal gland. This close

affinity was emphasized by Kantor (1996). based on

Hayashi, 2004). Members of

shared characteristics such as a long or very long pro-

boscis, the loss of glandular dorsal galas. and a te ages ncy

toward the reduction of the gland of Leiblein. The mo-

lecular systematics work by Galindle et al. (2016) provided

a hypothesis for the sy stematic relationships within the

Nassariidae.

Buccinids are usually distinguished from the nassariids

by having a radula with three or more cusps on the central

teeth, hie ral teeth with two large flanking cusps (Golikov,

1980; Kantor, 1990a), and zero to five intermediate cusps

(Harasewych, 1998). These radular distinctions, however,

have not been observed in Engoniophos unicinctus (Say,

1825), which shows shell and radula greatly similar to

those of the Nassariidae (Bandel, 1984). This led some

authors to suggest its relocation to Nassariidae, and not in

Buccinidae as it is traditionally allocated (Bandel, 1984;

Faber, 2004).

The present paper is part of a larger project aiming at an

improvement of the taxonomic knowledge on the Bucci-

noidea. Focusing in a problematic species with a clear duality

of familiar attribution, we intend to discuss its morpho-

anatomical attributes which has been published on nas-

sariids and buccinids, and provide new data on the species.

MATERIALS AND METHODS

Specimens examined in this study are deposited in the

collection of the Museu de Zoologia da Universidade de

Sao Paulo (MZSP). A list of examined specimens is given

after the species description. Shells were broken prior to

soft-tissue extraction. Specimens were immerse din 70%

ethanol and dissected by standard techniques under

a stereomicroscope. The terminologies of Amold (1965)

and Simone (2011) were employed for shell cone TS,

and Diver's (1931) whorl-counting method was applied.

Details of the radula were examined under the scanning

electron microscope (SEM) at the MZSP. The synonymic

list presented herein is restricted to taxonomic and oc-

currence papers.

Page 102

Abbreviations used in anatomical drawings: aa, anterior

aorta; ab, albumen gland; ae, anterior oesophagus; ag,

albumen gland; an, anus; au, auricle; bg, buccal ganglia;

br, subradular membrane; ce, cerebral ganglion; eg,

cement gland; em, columellar muscle; co, cement gland

orifice; ev, ctenidial vein; df, dorsal fold of buccal mass:

dg, digestive gland; dl, gland of Leiblein duct; ep, pos-

terior oesophagus; es, oesophagus; fp, female pore; fs,

stomach folds; ft, foot: ge, suboesophageal ganglion; gi,

gill; hg, hypobranchial gland; kd, kidney, ml-ml11.

odontophore muscles: il, mantle border; me, medium

oesophagus; mj, julgal muscle; mo, mouth; mt, mantle;

ne, nephrostome; ng, nephridial gland; nr, nerve ring; oe,

odontophoral cartilage; od, odontophore; og, osphradium

ganglion; os, osphradium; ot, oral tube; ov, pallial oviduct;

oy, Ovary; pa, posterior aorta; pb, proboscis; pd, penial

duct; pe, penis; pf, propodial expansion; pg, pedal gland;

po, palial oviduct; pp, penial papilla; pr, propodium: pu,

pleural ganglia; ra, radula; rm, retractor muscle of pro-

boscis; rm, radial ar nucleus; rs, radular sac; rt, rectum; sd,

salivary duct; sf, siphon; sg, salivary gland; so, salivary duct

orifice; su, suboesophageal ganglia: te, tentacle; va,

vaginal atrium; vd, vas de ses -rens: ve, ventricle: vo, visceral

oviduct; ye, eye.

RESULTS

Family Nassariidae Iredale, 1916

Subfamily Photinae Gray, 1857

Genus Engoniophos Woodring, 1928

1873 by original

Type Species: Phos erectus Guppy,

designation. Miocene, Jamaica.

Engoniophos unicinctus (Say, 1826)

(Figures 1—25)

Nassa unicincta Say, 1826: 211-212 (pl. 57, figs 1, la

Tryon, 1873: 35 (fig. 55), 1882: 245.

Nassa pallida Powys, 1835: 96.

Nassa guadelupensis Petit de la Saussaye, 1852: 171 (pl. 2

figs & 4).

Strongylocera textilina Morch, 1852: 80 (Lister, pl. 965,

fig. 20).

Phos guadalupensis |sic]: Arango, 1880: 201.

Phos guadeloupensis [sic]: Tryon, 1881: 219 (pl. 83, figs

512, 520).

Nassa textilina: Tryon, 1882: 243.

Phos unicinctus: Dall, 1889: 178; Gardner, 1926: 460.

Strongylocera unicincta: Dall and Simpson, 1901: 402.

Engoniophos guadelupensis: Warmke and Abbott, 1962:

115 (pl. 21, fig. F); Humfrey, 1975: 150; Bandel, 1976:

99 (fig. 3a—b).

Engoniophos unicinctus: Rehder, 1962: 131; Warmke and

Abbott, 1962: 116 (pl. 21, fig. J); Work, 1969: 672; Abbott,

1974: 220 (fig. 2428): Wolves andl Volzes, 1983: 26 (pl. 14,

fig. 21); Bande 1, 1984: 142 (fig. 235; pl. 16, fig. 6); Jong and

THE NAUTILUS, Vol. 132, No. 3-4

Coomans, 1988: 83 (pl. 38, fig. 455); Diaz and Puyana,

1994: 187; Ramos and Role. 1994: 102: Faber, 2004: 8:

Buitrago et al., 2006: 639; Cruz and Gandara, 2006: 132;

Riemivindle z and Jiménez, 2007: 5; Reyes et al., 2007: 384;

Rosenberg et al., 2009: 650.

Pallacera unicincta: Macsotay and Campos, 2001: 88.

Type Locality: Coast of South Carolina (Say, 1826 in

error; see Woodring, 1964: 269).

Distribution: Lower Caribbean to Venezuela (Macsotay

and Campos, 2001).

Description: SHELL (Figures 1, 3, 4). Fusiform, twice as

long as wide, with 7-8 convex whorls, light gray to cream.

Protoconch wide, smooth, dome- shaped, white, with

three whorls; transition indistinct. Teleoconch sculpture

consisting of 5—S thin spiral cords along entire surface of

all whorls (twice as many on body volharal)s space between

cords equals three to four times their width, becoming

more closely spaced near suture; spiral cords crossed Ib

wide axial ribs (width ~1/10 of whorl width) bearing

rounded nodules at middle level on earlier whorls, and on

upper portion of body whorl; axial ribs becoming less

prominent toward suture and basal portion of body whorl;

depression between ribs as wide as ribs. Spire angle Oo.

Aperture elliptical, twice as long as wide, white, glossy;

length ~1/2 of shell length. Siphonal canal short, broad,

dorsally recurved. Armall notch present, but not well-

marked. Outer lip thick, lirate, with thickened outer

edge. Columella straight, without folds.

Heap-roor (Figures 8, 9, 11, 13). Head protruded, ten-

tacles elongate d and narrow, twice as long as head; eyes

located on “oral protuberances at mid level of tentacle,

clearly separating basal broad and distal narrow portions.

Rhynchostome as transverse slit, located in middle region

of ventral surface of head (Figure 9, 15). Foot large,

occupying whole body whorl (retracted), bearing pair Oe

smal] propodeal expansions; ene gland located in central

region of foot (Figure 11), forming groove, extending

from dorsum of foot (propodium) to sole. Cement gland

of females rounded, located in anterior region of meso-

podium, inner space wide, duct long (Figure 11). Cement

gland orifice located on median line of anterior sole region

(Figure 13). Opercular pad elliptical, ~80% as wide as

dorsal surface of foot; attachment with operculum oc-

cupying ~70% of foot area. Penis originated on right

lateral region, posterior to cephalic base, at level of mantle

edge. Columellar muscle wide and broad, 1% whorl long.

Haemocoel long and thin, extending dorsally along center

of foot and columellar muscle (Figure Qi),

OpeRcULUM (Figure 2). Small, oval, corneous, pale brown,

located close to edge of foot; occupying 1/2 of apertural

area. Nucleus terminal, inferior. Outer surface with

normal concentric oe lines, forming undulations.

Scar oval, occupying ~7/3 of inner aries,

D. Abbate et al., 2018 Page 103

007

Figures 1-7. Engoniophos unicinctus, MZSP 77798 8, shell, operculum and radula. 1. Shell, apertural and dorsal views (L = 18.7

mim). 2. Operculum, inner and outer views. Scale bar = 2 mm. 3-4. Protoconch and first teleoconch whorls. 3. Lateral view. Scale bar =

0.5 mm. 4. Apical view. Scale bar = Imm. 5-7. SEMs of radula. 5. Middle portion of radula. Scale bar = 30 xm. 6. Detail of central

teeth. Scale bar = 10 wm. 7. Detail of lateral teeth. Scale bar = 10 pm.

Page 104 THE NAUTILUS, Vol. 132, No. 3-4

po rt

Figures 8-14. Engoniophos unicinctus, detailed anatomy. 8. Complete specimen extracted from shell (operculum removed), an-

terior-right view. 9. Detail of head, dorsal view. 10. Reno-pericardial region, ventral view, some adjacent structures also shown. 11. Foot

of female, sagital section. 12. Pallial cavity roof, ventral view, and coiled visceral mass. 13. Sole of foot, female, showing propodial

expansions and cement gland opening. 14. Pallial cavity roof, transverse section at middle level of osphradium. Scale bars = 2 mm.

D. Abbate et al., 2018

Page 105

MANTLE Cavity Orcans (Figures 10, 12, 14). Mantle cavity

covering ~1 whorl. Siphon width ~1/2 of mantle cavity

width, i ngth ~2/3 of mantle cavity length. Right base of

siphon high, width ~twice of mantle e dge Satie left base

low. Osphradium long and narrow, elliptical, length ~1/3 of

pallial cavity length, seagthh ~1/6 of its width. Osphradial

filaments short, w aks similar to mantle edge. Ctenidial vein

(efferent branchial vessel) with uniformly narrow along its

length. Gill elliptic, occupying ~80% of le ngth and wlVa of

width of pallial cavity. Anterior end of gill aun d, ending

gradually, inserted directly into pallial cavity. Gill filaments

relatively uniform in size along its entire le ngth. Posterior

end of gill rounded, located in posterior region of mantle

cavity Rinse to pericardium. Gill filaments height ~1/3 of

pallial cavity height, apex rounded, at middle portion, tilted

to right; right and left ec dges of filament straight; space

between gill and right pallial organs about twice of gill

width. Hypobranchial gland inconspicuous. Right side of

pallial cavity almost entirely filled by gonoducts. Rectum

long and thin, with ~1/2 of p: allial cavity length. Anus sessile,

distanes between anus and mantle border 1/3 of total pallial

cavity length. Anal gland absent.

ViscERAL Mass (Figures 8, 10, 12). Extending ~3 whorls

posteriorly to pallial cavity. Digestive gland Genk beige,

occupying ~S0% of visceral mass Palme! encircling

stomach. Gonad orange with small black spots, located on

columellar surface, posterior to stomach. Seminal vesicle

of males located in anterior portion of gonad, ~% of its

size. Kidney occupying ~1/3 of visceral mass volume,

located on right side of anterior visceral end. Stomach

not seen in ale ail.

CIRCULATORY AND EXCRETORY SysTEMS (Figure 10, 12).

Reno-pericardial region occupying 1/3 whorl, oval, on right

margin of last soho of visceral mass, adjacent to mantle

cavity. Pericardium occupying 1/3 of reno-pericardial

region, posterior to gill; situated on left anterior margin

of wisearall mass. Aunties reniform, anterior to wreralvatelles

with three connections: on upper right side with kidney, on

anterior right side with gill, and on posterior left side with

ventricle. Vemaaalle four times size of auricle, with common

aorta on left posterior margin. Aortas wide, located along

left-posterior region of ee ele Kidney occupying 1/5 of

pallial cavity volume, renal lobe single, solid, with glandular

transverse folds along its ventral Moree etrerent renal

vessel located at its right portion. Nephridial gland not seen

in detail. Nephrostome a small, transverse slit, located in

anterior region of membrane between kidney and pallial

cavity.

DicEsTIVE SysTeM (Figures 9, 15-21). Mouth longitudinal,

narrow, located on center proboscis tip. Proboscis long,

straight and thin, occupying ~80% of hemocoel aeeltinnnes

not ‘completely retractable. Rhinchodeal wall thin, in-

volving ~1/3 of proboscis. Retractor muscles covering ~ 1/3

of proboscis; several thin retractors muscles originated on

dorsal surface of foot, inserted into posterior end surface of

proboscis. Odontophore and buccal mass muscles: mj, thin

pairs of perioral muscles connected on both sides, dorsal

and ventral, surrounding odontophore cartilages; m1, jugal

muscles, several small muscle fibers connecting buccal

mass to adjacent inner surface of proboscis; m2, pair of

strong retractor muscles of buccal mass, originating on

inner surface of proboscis, running along entire odontophore,

inserting into anterior region of odontophore cartilages:

m2a, pair of retractor muscles of buccal mass, originating

on dorsal surface of haemocoel, inserting at end of

posterior margin of odontophore cartilages; m2b, ventral

single, thin Paeele auxiliary of m2, originating on ventral

medial fibers of m2, detaching from it in region just

posterior to m6; m3, long, ~80 % of odontophore le ngth,

cylindrical muscle forming outer wall of odontophore, eatlh

transverse fibers: m4, pairs of strong radular dorsal tensor

muscles cove ring almost entire rane of poste nor por-

tion of odontophore cartilages, inserting into subradular

membrane; m5, pair of sxovaltierey dorsal tensor muscles of

radula, originating inside edges of cartilage, adjacent to

rasertion of m4: m6, horizontal muscle, thick, connecting

ventral edges of cartilages, running almost along its entire

length; m8, pair of small elliptical muscles, length ~1/3 of

odontophore cartilage length, originating at anterior end

of odontophore cartilages, running along ventral surface

of odontophore, inserting on anterior ventral surface of

cartilages; m11, pair of ventral tensor muscles of radula,

elongated, about 1/2 of total odontophore length,

originating at ventral-posterior end of cartilages, crossing

ventrally entire odontophore, inserting into ventral posterior

surface of radula. Additional odontophore structures: br,

subradular membrane, thin, translucent, along entire length

of radular ribbon, covering inner surface of odontophore

cartilages; oc, odontophore cartilages, about 3 times as long

as wae inner ventral surface concave, ~1/4 of anterior end

fused with each other, ~1/3 of anterior region, concave,

involving radular ribbon; rs, radular sac thin-walled,

cylindrical, located at posterior end of radula. Radular

teeth (Figures 5, 6, 7): rachidian tooth wide, comb-like,

occupying about half of radular width; base curved, width

~3% its length; ~9 triangular, sharp pointed cusps of similar

size, except for some lateral reduction; lateral tooth hook-

like, bicuspid, base broad (equivalent to rachidian base

width), obliquely disposed; external lateral cusp widely

curved inwards, about as long as base; inner cusp

approximately half size of main lateral cusp. Salivary

glands (Figure 15) small, located at anterior portion of

aeroeoel occupying ~1/8 of haemocoel volume, entirely

involving nerve ring, middle esophagus and anterior

portion mat proboscis. Salivary ducts very narrow, except

for short proximal region running completely attached to

anterior esophagus wal] and, more anteriorly, inside dorsal

folds of buccal cavity (Figure 16); opening very small

(Figure 16: so), in anterior-middle region ae dorsal folds

of buccal cavity. Valve of Leiblein (Figures 15, 18: vl) large,

about 1/8 of odontophore volume, located in medium

esophagus anterior to nervous ring, antenor region w ith

transverse white band bearing long cilia, middle and

posterior regions white, corresponding to inner gland

Page 106 THE NAUTILUS, Vol. 132, No. 3-4

Figures 15-19. Engoniophos unicinctus. Detailed anatomy. 15. Head and haemocoel, ventral view, foot and columellar muscle

removed. 16. Proboscis and anterior esophagus opened longitudinally, showing salivary ducts and their apertures. 17. Extended proboscis

opened longitudinally, ventral view, odontophore as in situ. 18. Mid and anterior region of posterior esophagus and associated structures,

showing valve and gland of Leiblein. 19. Odontophore, dorsal view, superficial layer of membrane removed. Scale bars = 2 mm.

D. Abbate et al., 2018 Page 107

Figures 20-25. Engoniophos unicinctus. Detailed of anatomy. 20. Odontophore, dorsal view, superficial muscles dissected. 21.

Odontophore cartilages, dorsal view, some adjacent muscles shown. 22. Penis, ventral view, penis duct shown by translucency. 23-24.

Nerve ring, ventral and dorsal views. 25. Pallial oviduct, ventral view, transversely sectioned at its middle level, some adjacent structures

also shown. Scale bars = 2 mm.

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THE NAUTILUS, Vol. 132, No. 3=4

occupying most of inner surface; bypass absent. Gland of

Leiblein narrow and elongated (Figures 15, 18: gl), ~twice

as long as middle esophagus, becoming gradually narrower

posteriorly; duct of gland of Leiblein straight, long and very

narrow, length ~1/2 of medium esophagus length, width

~1/6 of medium esophagus width. Accessory salivary glands

absent. Anterior esophagus broader; walls muscular, with

several dorsal internal longitudinal folds, occupying entire

length of proboscis. Middle esophagus slender, short,

diameter and length approximately half those of anterior

esophagus. Posterior esophagus narrow, straight, about

as long as anterior esophagus; anterior region broad,

differentiation between middle and posterior esophagus

clear with a thin duct. Stomach not seen in detail.

REPRODUCTIVE SysTEM, MALE (Figure 22). Vas deferens

narrow, simple, straight, running along ventral surface

of kidney up to pallial cavity. Prostate totally closed

Figures 26-28. Hard structures of some nassarids for comparison. 26. Nassarius albus MZSP: 109759 shell, apertural and dorsal

views. 27. Operculum of Nassarius arcularia plicatus MZSP: 99863 outer view, scale bar = 2 mm. 28. Radula of Nassarius arcularia

plicatus, SEM, MZSP: 99863, detail of central and lateral teeth. Scale bar = 20 pm.

D. Abbate et al., 2018

Page 109

(tubular), running through right mantle edge, at ~2/3

of total pallial cavity length. Vas deferens anterior to

yrostate, straight, running immersed into integume nt of

dorsum, next to mantle arias sr and penis base. Penis large,

length ~1/2 of total head-foot length, dorso-ve ori

flattene -d; base curved, apical region pointed, Penial duct

29 cm

Figures 29-31.

straight, running through center of penis, closed (tubular).

Penial papilla long, slender, subterminal, located at anterior

left region of penis. Penial aperture apical, far from papilla.

REPRODUCTIVE SYSTEM, FEMALE (Figures 10, 12, 25). Visceral

oviduct narrow, straight, running along ventral surface of

Anatomy of Nassarius vibex. 29. Complete specimen extracted from shell ( (operculum removed), right-anterior view.

30. Penis, ventral view, transverse section at indicated levels also shown. 31. Head-foot, dorsal view, showing met :podis al tentacles. Scale

bars = 2 mm.

Page 110

THE NAUTILUS, Vol. 132, No. 3-4

kidney in first whorl of visceral mass, located in front of

pallial cavity. Posterior region of pallial oviduct protruding

into kidney, occupying almost entire length of pallial cavity

and 1/3 of its width. Albumen gland posterior, whitish,

thick-walled, occupying ~1/5 of pallial oviduct volume.

Capsule gland ~4/5 of oviduct volume, elongated, orange,

thick- male d. Vaginal atrium anterior to capsule Bland

occupying ~1/7 of oviduct volume: sinuous, walls nck.

muscular, female genital pore narrow, protruded, papilla-

like, with thick edges. Bursa copulatrix absent.

CreNTRAL Nervous SysTEM (Figures 25, 26). Nerve ring lo-

cated in ventral basal proboscis region (Figure 15: nr),

occupying ~ 1/12 of haemocoel volume, highly concentrated.

Ganglia mostly fused, hard to distinguish from each other,

SOE asymmetrical. Pairs of pleural and cerebral

ganglia fully fused with each other. Sub- esophageal

ganglion about half of cerebro- pleural ganglion size, shortly

andl broadly connected to right cere beal ganglion. Pair of

buccal ganglia small, located close to posterior edge of

cerebral ganglia. Esophageal aperture small, occupying

about 1/9 of ventral surface of nerve ring. Statocysts not

seen.

Shell Measurements (length < width in mm): MZSP

THis YS LO. K 1O¢Ze 175 XK VOr GS IBD) X< Bil

18.7 X 9.5.

Habitat: Muddy and sandy bottoms, intertidal.

Material Examined: MZSP 77798: 26, 22, Venezuela:

Isla Margarita; Playa Bella Vista, 10°56’ N, 63°50 W,

3-4 m als »pth ( (Simone col., 02/ii/1995).

DISCUSSION

Engoniophos unicinctus has a clear conchological re-

semblance to some representatives of Nassariidae, e.g.,

Tritia alba (Say, 1826) (Figure 26), in having a small shell

with a poorly develope d parietal callus, a high spire, short

siphonal canal, weak anal notch, and uniform sculpture

throughout the- shell surface. Moreover, there is also

a resemblance in radular configuration. The radula of

E. unicinctus shows lateral reas bearing two similarly

sized cusps, lacking the smaller alert FEU teeth

present in Buccinum undatum Linnaeus, 1758, type

species of Buccinidae (Fretter and Graham, 1962: 171,

fig. 105E). On the other hand, the species in the nassariid

genus Buccinanops also bears intermediate cusps in lat-

eral teeth. It also shows rachidian teeth with a larger

number of cusps, as in Nassarius arcularia plicatus

(Roding, 1798), a subspecies of the type species of

Nassariidae (Figure 28). Such characteristics led Faber

(2004) to argue that E. unicinctus would be better placed

in Nassariidae, following Bandel’s (1984) suggestion, in-

stead of being allocated in Buccinidae, as proposed by

certain authors in more traditional classifications (e.g.,

Cernohorsky, 1984; Miloslavich, 1999; Reyes, 2007).

Engoniophos unicinctus presents some anatomical

differences from those of typical nassariids, for example,

the operculum (Figure 2) lacking ae serrations

commonly found in nassariines (Figure 2 27), but these are

also absent in dorsanine fiiceinanopanes The head of

nassariids is generally well developed (Figure 29), while in

E. unicinctus it is proportionally much smaller and sim-

pler, with the tentacles inserted directly into the dorsum

(Figure 8: te). The penis is short and broad, with a long

and slender papilla in E. unicinctus (Figure 22), while in

other nassariids it is normally long and slender, lacking

papillae (Figure 30). The epipodial tentacles (Figure 31),

either singly or in pairs, are a traditional character and

a demonic anatomical feature of Nassariidae (Simone

and Pastorino, 2014). This feature is not found in E.

unicinctus, which only bears a pair of propodeal (anterior)

expansions (Figure 13), also ee found in other

nassariids, but no tentacles. Last but not least, recent

molecular analyses, place E. unicinctus within the Nas-

sariidae (Galindo et al., 2016).

The morphological similarities above mentioned

common with nassariids and the aforementioned recent

findings in the literature so far indicate that E. wnicinctus,

in fact, belongs to Nassariidae. This conclusion adds an-

atomical argumentation to the already known conchological

and anole similarities, and further corroborates the fa-

miliar placement, and its positioning within Buccinoidea

proposed by Galindo et. al (2016).

ACKNOWLEDGMENTS

The authors are grateful to Lara Guimaraes (MZSP) for

helping with the SEM examination. This work was par-

tially supported by a doctoral grant by the Conselho

Nacional de Desenvolvimento Games e Tecnolégico

(CNPq, proc. No 159448/2012-3).

LITERATURE, CITED

Abbott, R.T. 1974. American Seashells: The Marine Mollusca of

the Atlantic and Pacific Coasts of North America, 2"¢ ed.

Van Nostrand Co., New York, 663 pp.

Arango, R.M. 1880. Contribucion a la fauna malacologica

ants yana. Havana: Imp. de G. Montiel y comp., 315 pp.

Arnold, W.H. 1965. A glossary of a thousand and one terms used

in conchology. The Veliger 7: 1-50.

Bandel, K. 1976. Morphologie der Gelege und 6kologische

Beobachtungen an Buccinaceen (Gastropoda) aus der

siidlichen Karibischen See. Bonner Zoologische Beitrige

27: 98-133.

Bandel, K. 1984. The radulae of Caribbean and other Meso-

gastropoda and Neogastropoda. Zoologische Verhandelin-

gen 214: 1-188.

Buitrago, J., J. Capelo, J. Gutiérrez, M. Rada, R. Hernandez,

and S. Grune. 2006. Living macromolluses from a paleo-

reef region on the northeastern Venezuelan continental

shelf. Estuarine, Coastal and Shelf Science 66 (3-4):

634-642.

Cernohorsky, W.O. 1984. Systematics of the family Nassariidae.

Bulletin of the Auckland Institute and Museum 14: 1-356.

D. Abbate et al., 2018

Page 11]

Cruz, F.V. and C.G. Gandara. 2006. Lista actualizada de los

gasterOpodos de la planicie del Arrecife Lobos, Veracruz,

México. Revista UDO Agricola 6 (1): 128-137

Dall, 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 2,

Gastropoda and Scaphopoda. Bulletin of the Museum of

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

10-40.

Dall, W.H. and C.T. Simpson. 1901. The Mollusca of Porto Rico.

Bulletin of the United States Fish and Fisheries Com-

mission 20: 351-524, pls. 53-58.

Diaz, J.M.M. and M.H. Puyana. 1994. Moluscos del Caribe

Colombiano: Un catélogo ilustrado. Colciencias Fundacién

Natura, Bogota, 291 pp.

Diver, C. 1931. A method of determining the number of the

whorls of a Shell and its application to Cepaea hortensis

Mill. and C. nemoralis L. Proceedings of the Malacological

Society of London 19: 234-239.

Faber, M.J. 2004. Marine gastropods from ABC-islands and

other localities. 2. The family Nassariidae (Gastropoda:

Buccinoidea). Miscellania Malacologica I (ie 71s.

Femandez, J.R. M. and M.M.P Jiménez. 007 Fauna mala-

colégica del litoral rocoso de la costa sur del Golfo de

Cariaco y costa norte del estado Sucre, Venezuela. Boletin

del Instituto Oceanografico de Venezuela 46 (1): 3-11.

Fretter, V. and A. Graham, A. 1962. British Prosobranch

Molluscs. Their Functional Anatomy and Ecology. The Ray

Society Institute, London, 775 pp.

Galindo, L.A., N. Puillandre, J. Utge, P. Lozouet, and P.

Bouchet. 2016. The phylogeny and systematics of the

Nassariidae revisited (Gastropoda, Buccinoidea). Molecular

Phylogenetics and Evolution 99: 337-353.

Gardner, J.A.

Group of Florida. Florida. U.S. Government Printing Of-

fice, 709 pp.

Golikov, A.N. 1980. Molluses Be of the World Ocean.

Fauna SSSR, Molluski 5 (2): 1466.

Guppy, R.J.L. 1873. On some new Tertiary fossils from Jamaics:

Raster sei ss of the Scientific Association of Trinidad 2

(2/10): 7 2-88.

Harasewych, "M.G. 1998. Infraorder Neogastropoda, pp.

819-845. In: Beesley, P.L. and G.].B. Ross (eds.). Mollusca

the Southern Synthe sis, Vol. 5, Part. B. Csiro Publishing,

669 pp.

Hayashi, S. 2004. The molecular phylogeny of the Buccinidae

(Caenogastropoda: Neogastropoda) as inferred from the

complete mitochondrial 16S rRNA gene sequences of

selected representatives. Molluscan Research 25 (2):

§5-958.

Humfrey, M. 1975. Sea Shells of the West Indies: A guide to

the marine molluscs of the Caribbean. London, Collins,

351 pp

Jong, K.M. and H.E Coomans. 1988. Marine Gastropods from

Curagao, Aruba and Bonaire. E.J. Brill, Leiden, 261 pp.

Kantor, Yu.I. 1990a. Anatomical basis for the origin and evo-

lution of the toxoglossan mode of feeding. Malacologia 32:

3-18.

Kantor, Yu.I. 1996. Phylogeny and relationships of Neo-

gastropoda, p. 221- 930 in: Taylor, J. (ed.) Origin and

evolutionary radiation of the Mollusca. Oxford Unive rsity

Press, Oxford.

1926. The Molluscan Fauna of the Alum Bluff

Lister, M. 1688. Historiae sive synopsis methodical con-

chyliorum. Liber IV. London, Published by the author, pl.

965, fig. 20.

Macsotay, O. and R.A. C Jampos. 2001. Moluscos Representativos

de la Plataforma de Margarita, Venezuela: Descripcién de

24 especies nuevas. Vale ncia, Venezuela, Privately pub-

lished. iii + 230 pp.

Miloslavich, P. 1999. Nutritional value of the intracapsular

liquid of Beet yee unicinctus Say, 1825 (Caenogas-

sae Buccinidae). Journal of Molluscan Studies 65

(4): 502-503.

Morch, O.A.L. 1852. Catalogus Conchyliorum quae ee liquit D.

Alphonso d’Aguirra & Gade a Comes de Yoldi.

Primus. Hafniae, Typis I mdovici Kleini, vi + 170 pp.

Rehder, H.A. 1962. Contribucién al conocimiento de_ los

moluscos marinos del Archipiélago de los Roques y la

Orchila. Memoria de la Sociedad de Ciencias Naturales La

Salle 22 (62): 116-1358.

Reyes, J.L., A.F Sanchez, J. Carruyo-Noguera, C.L. Casler, S.

Narciso, M. Nava, and A. Guerra-Gémez. 2007. Moluscos

Gasterépodos y Bivalvos de la Alta Guajira, Estado Zulia,

Venezuela. eeleen del Centro de Investigaciones Bio-

logicas 41 (3): 376-393.

Petit de la eaten S. 1852. De scription de coquilles nouvelles.

Journal de Conchyliologie Be Dil).

Ponder, W.F. 1973. The origin and evolution of the Neo-

Sere Malacologia 12: 295-338.

Powys, W.T. 1835. C Neencrers of new species of shells collected

by ve Cuming. Proceedings of the Zoological Society of

London 3: 94— 96.

Ramos, G.H. and G. Robaina. 1994. Contribucién al con-

ocimiento de los moluscos gasteropodos y pelecipodos de

la bahia de Mochima, Bale, Sucre, Venezuela. Memoria

de la Sociedad de Ciencias Naturales La Salle 54 (141):

95-106.

Say, T. 1826. Descriptions of marine shells recently dis-

covered on the coast of the United States. Journal of

the Academy of Natural Sciences of Philadelphia 5:

207-221.

Fasciculus

Simone, L.R.L. 2011. Phylogeny of the Caenogastropoda

(Mollusca), based on comparative morphology. Arquivos de

Zoologia 42 (2-4): 83-323.

Simone, L.R.L. and G. Pastorino. 2014. Comparative mor-

phology of Dorsanum miran and Bullia granulosa from

Morocco (Mollusca: Caenogastropoda: N Nassariidae). Afri-

can Invertebrates 55 (1): 125-142.

Rosenberg, G., F. Moretzsohn, and E.F. Garcia. 2009. Gas-

tropoda (Mollusca) of the Gulf of Mexico. pp. 579-699 in:

Felder, D.L. and D.K. Camp (eds.). Gulf of Mexico —

origins, Waters, and Biota. Biodiversity. Texas, College

Station, A&M University Press.

Taylor, J.D., N.J. Morris, and C.N. Taylor. 1980. Food spe-

cialization and the evolution of predatory prosobranch

gastropods. Palaentology 23 (2): 375-409.

Tracey, S., J.A. Todd, and D.H. Erwin. 1993. Mollusca: Gas-

tropoda. Pp. 131-167 in: Benton, M_J. (ed.) The Fossil

Record 2. London, Chapman and Hall.

Tryon, G.W. 1873. American marine conchology: or, de-

scriptions of the shells of the Atlantic coast of the United

States from Maine to Florida. Philadelphia, published by

the author, 224 pp.

Tryon, G.W. 1881. Manual of Conchology: Structural and

Systematic, with illustrations of the species. ser. 1, v. 3.

Tritonidae, Fusidae, Buccinidae. Philadelphia, published

by the author, 510 pp.

Page 112

THE NAUTILUS, Vol. 132, No. 3-4

Tryon, G.W. 1882. Manual of Conchology: Structural and

Systematic, with illustrations of the species. ser. 1, v. 4.

Nassidae, Turbinellidae, Volutidae, Mitridae. Philadelphia,

published by the author, 408 pp.

Vokes, H.E. and E.H. Vokes. 1983. Distribution of shallow-water

marine Mollusca, Yucatan Peninsula, Mexico. Mesoamerican

Ecology Institute Monograph 1. Middle American Re-

search Institute Publication 54. New Orleans, Middle

American Research Institute, 154 pp.

Warmke, G.L. and R.T. Abbott. 1962. Caribbean Seashells: A

Guide to the Marine Mollusks of Puerto Rico and Other

West Indian Islands, Bermuda and the Lower Florida Keys.

Livingston, Livingston Pub. Co., 348 pp.

Woodring, W.P. 1928. Miocene mollusks from Bowden,

Jamaica. Contributions to the Geology and Palae-

ontology of the West Indies. Carnegie Institution of

Washington publication, no. 385. Carnegie Institution of

Washington.

Woodring, W.P. 1964. Geology and paleontology of Canal Zone

and adjoining parts of Panama; Description of Tertiary

mollusks (gastropods; Columbellidae to Volutidae). United

States Geological Survey Professional Paper 306-C:

241-297.

Work, R.C. 1969. Systematics, ecology and distribution of the

mollusks of Los Roques, Venezuela. Bulletin of Marine

Science 19: 614-711.

THE NAUTILUS 132(3—4):113-116, 2018

Page 113

Mericella zhangsupingae, a new cancellariid species from the South

China Sea (Gastropoda: Cancellariidae)

Shugian Zhang’

Peng Wei

Laboratory of Marine Organism Taxonomy and Phylogeny

Institute of Oceanology, C hinese Academy of Sciences

Qingdao 266071, CHINA

And

Center for Ocean Mega-Science, Chinese Academy of Sciences

Qingdao, 266071, CHINA

ABSTRACT

A new western Pacific species belonging to the family Can-

cellariidae is described from off Dongsha Islands, South China

Sea, at bathyal depths of 500-600 m. Mericella shangsupingae

new species is most similar to Mericella bozzettii Petit and

Harasewych, 1993 in shell size and proportion. However, the

new species can be separated from that species in having a larger,

depressed protoconch, more convex teleoconch whorl, much

weaker sculpture, and more sinuous outer lip. The new species

represents the first member of genus Mericella discovered from

westerm Pacific.

Additional Keywords: Volutoidea, Dongsha Islands, China,

protoconch

INTRODUCTION

The genus Mericella was originally established by Thiele

(1 929), as a subgenus of Craccionio, to accommodate

a single species previously treated as Cancellaria (Merica)

jucunda Thiele, 1925. To date, three species have been

recognized in the genus, including Mericella jucunda

(Thiele, 1925) fost off Tanzania, Mericella paschalis

(Thiele, 1925) from off Tanzania and Mozambique, and

Mericella bozzettii Petit and Harasewych, 1993 from off

Somalia. Olsson and Bayer (1972) proposed the generic

name Gerdiella to include three cancellariid species from

bathyal depths (516-897 m) of the Florida Straits and the

Caribbean Sea. In that publication, they recognized that

Gerdiella was related to, and possibly congeneric with,

Mericella, but would differ by shell size and “geographical

distribution. Verhecken and Bozzetti (2006) compared

the two genera based on conchological characters, and

found no reason for further generic separation, although

recognizing some degree of difference (e.g. spire height,

suture form, relatively aperture height). However, a

l ; ~ ae

Author for correspondence: [email protected]

significnt distinction between the two genera was over-

looked by Verhecken and Bozzetti (2006). Mericella

species have a smooth protoconch, while Gerdiella species

have axial ribs on the protoconch. In addition, some

microscopic spiral threads were observed on the proto-

conch of Gerdiella alvesi Lima, Barros, and Petit, 2007.

Verhecken (2002) stated that protoconch characters are of

no diagnostic importance at the generic level. This opinion

was not followed by Lima et al. (2007), who regarded

Gerdiella as a distinct genus endemic to the western and

southern Atlantic Ocean.

In this paper we describe a western Pacific species

belonging to the Cancellariidae and assign it to the genus

Meric ila based on its smooth protoconch and geographic

proximity to other species of Mericella. The finding ex-

tends the distribution of Mericella from East Neen

waters to the western Pacific.

MATERIALS AND METHODS

Specimens were trawled by fishermen from bathyal

depths (500-600 m) off Dongsha Islands, South China

Sea (see Figure 1). Shells were observed using a light

microscope and protoconch characters by scanning

electron microscope (SEM). All specimens have been

deposited at the Marine Biology Museum of Chinese

Academy of Sciences (MBMCAS). The following ab-

breviations are used in the text: MBM, Marine Biological

Museum, Qingdao, China; USNM, National Museum of

National History, Smithsonian Institution, Washington, DC.

SYSTEMATICS

Family Cancellariidae Forbes and Hanley, 1851

Genus Mericella Thiele, 1929

Type Species: Cancellaria jucunda Thiele, 1925 (off

Dar es Salaam, Tanzania).

Page 114

THE NAUTILUS, Vol. 132, No. 3-4

Indian Ocean

@ Mericella jucunda (Thiele, 1925)

#F Mericella bozzettii Petit & Harasewych, 1993

¥ Mericella zhangsupingae new species

30 40 50 60

Geographic distribution of known Mericella species.

Figure 1.

Mericella zhangsupingae new species

(Figures 1—12)

Description: Shell (Figures 2-12) large for genus, up to

31.5 mm, solid, elongated, with a tall spire, conical,

protoconch rounded, aperture narrow. Protoconch

(Figures 11-12) with two smooth, glassy whorls, large,

maximum diameter nearly 2 mm, indicating plankto-

trophic larval development. Transition to teleoconch

distinct, marked by numerous weak axial incremental

processes. Protoconch forms smooth keel prior to tran-

sition to teleoconch. Teleoconch with up to five roundly

convex whorls. Suture constricted, narrow, deep. First

teleoconch whorl with four weak initial spiral cords, be-

coming five on second whorl, eight on penultimate whor!

and ca. 30 on body whorl, occasionally with intercalated

spiral threads. Axial sculpture of regularly spaced, sinuate,

opisthocline ribs, 17 on first teleoconch whorl, 22 on

second whorl, forming sharp nodules and cancellated

appearance at intersections with spiral cords. Axial ribs

and nodules becoming weak and smoother toward pen-

ultimate and body whorl. Incremental lines distinct,

dense, forming conspicuous axial ridges between spiral

interspaces. Varices up to seven in number, very weak on

spire whorls, distinct on body whorl. Aperture narrowly

sO) 10) 00) 1910) 420) WE

elongate, elliptical, deflected from coiling axis by 18° (in

holotype). Outer lip thickened, with about 20 obsolete

denticles along entire length of the flaring, strongly sin-

uate outer lip; columellar lip with two developed folds.

Radula and soft parts unknown.

Type Material: Holotype, MBM286508 (height 31.5 mm;

width 13.8 mm); Paratypes 1-2, MBM286509, all from the

type locality.

Type Locality: Off Dongsha Islands, South China Sea,

500-600 m.

Etymology: This new species is named after Prof. Suping

Zhang in recognition of her contribution to gastropod

taxonomy in China.

Comparative Remarks: Based on its elongate shell with

a smooth protoconch and a flaring, sinuate outer lip, we

assign this new cancellariid species to genus Mericella.

Mericella zhangsupingae new species is most similar to

Mericella bozzettii Petit and Harasewych, 1993 (Figures

13-15) in shell size and proportion, but differs in having

a larger, lower, more globose protoconch, more convex

teleoconch whorls, much weaker axial sculpture, and

S. Zhang and P. Wei, 2018 Page 115

Figures 2-15. Shells of Mericella species. 2-12. Mericella zhangsupingae new species. 2-4. Holotype, MBM286508, 31.5 mm. 5-7.

Paratype 1, MBM286509, 26.6 mm. 8-10. Paratype 2, MBM286509, 23.2 mm. 11-12. Protoconch of paratype 2. 13-15. Holotype

Mericella bozzettii Petit and Harasewych, 1993, USNM 860315, 28.7 mm.

Page 116

THE NAUTILUS, Vol. 132, No. 3-4

more sinuous outer lip. In addition, spiral sculpture is

dominant in Mericella zhangsupingae, whereas the axial

sculpture is dominant in Mericella bozzettii. Mericella

zhangsupingae new species can be clearly distinguished

from the other two congeners, Mericella jucunda

(Thiele, 1925) and Mericella “paschalis | (Thiele, 1925), by

its larger, thicker shell with roundly convex tele oconch

whorls. ana a white color. Previously, all members of

genus Mericella were only known from off East Africa

(see Figure 1). The finding of Mericella zhangsupingae

new species extends the dicen bution of the genus to the

western Pacific Ocean.

ACKNOWLEDGMENTS

We would like to express our sincere thanks to Drs. M.G.

Harasewych and José H. Leal for valuable comments and

meticulous editing. This research was financially sup-

ported by the N Jational Natural Science Foundation of China

(3] 750002, 41376167).

PIMP RARURE CED

Lima, S.F.B., R.E. Petit, and J. C.N. Barros. 2007. A new species

of Gerdiella (Gastropoda: Cancellariidae) from the South

Atlantic Ocean off Brazil with discussion of an undescribed

species. The Nautilus 121: 99-103.

Olsson, A.A. and F.M. Bayer. 1972. Gerdiella, a new genus of deep-

water cancellariids. Bulletin of Marine Science 22: 875-880.

Petit, R.E. and M.G. Harasewych. 1993. A new Mericella (Mollusca:

Gastropoda: Cancellariidae) from northeastem Africa. Pro-

ceedings of the Biological Society of Washington 106: 221-224.

Thiele, J. 1925. Gastropoda der Deutschen Tiefsee- -Expedition,

II. Deutsche Tiefsee-Expedition 17: 35-382.

Thiele, J. 1929. Handbuch der systematischen Weichtierkunde.

Gustav Fischer, Jenna 1: 7B.

Verhecken, A. 2002. Atlantic bathyal Cancellariidae (Neogastropoda:

Cancellarioidea): additional data, and description of a new

species. Journal of Conchology 37: 505-514.

Verhecken, A. and L. Bozzetti. 2006. New data on East-African

ee species, and description of a new species of

Scalptia (Neogastropoda: Cancellarioidea: Cancellariidae).

Gloria Maris 45: 14-25.

THE NAUTILUS 132(3-4):117-123, 2018

Page 117

A newly discovered Paleocene species of Boreocomitas

(Gastropoda: Pseudomelatomidae) from eastern Hokkaido, Japan,

with implications for the biogeography of the Paleocene

Bering Strait

Kazutaka Amano Krzysztof Hryniewicz

Department of Geoscience Institute of Paleobiology

Joetsu University of Education Polish Academy of Sciences

1Yamayashiki ul. Twarda 51/55

Joetsu 943-8512, JAPAN 00-818 Warszawa, POLAND

[email protected] [email protected]

Robert G. Jenkins

School of Natural System,

College of Science and Engineering

Kanazawa Unive rsity, Kanazawa City

Ishikawa 920-1151, JAPAN

[email protected]

ABSTRACT

Two species of the gastropod family Pseudomelatomidae, genus

Boreocomitas, ee the new species B. inouei, are described

from the Paleocene (upper Selandian to lowest Thanetian)

Katsuhira Formation in eastern Hokkaido, Japan. These species

represent the first Paleocene record of Boreocomitas. The paper

discusses also the new recognition of species of Boreocomitas

in Paleocene deposits of westerm Greenland and Denmark.

Occurrences of some taxa, including this genus, are suggestive of

faunal exchanges between the northern Pacific and the North

Atlantic/Artic regions from the middle Paleocene (Selandian) to

late Eocene. These exchanges could have resulted from direct

marine connections between both areas via the Bering Strait.

Additional Keywords: Paleocene, gastropod, Boreocomitas, new

species, palaeogeography

INTRODUCTION

Two species of a Paleocene bathyal gastropod genus

including a new species are reported from eastern

Hlclakeetidlo, Japan. These species are here assigned to the

pseudomelatomid gastropod genus Boece The

deep-sea gastropod genus Boreocomitas was proposed

by Hickman (1976) as an extinct subgenus of Comitas

Finlay, 1926, based on specimens from the Cowlitz and

Keasey formations in Oregon, western USA. Recent data

show that the age of ane. Cowlitz Formation is middle

to late Eocene Sand that of the Keasey Formation is

late Eocene to earliest Oligocene (Prothero, 2001).

Boreocomitas has also been recorded from the upper

Eocene Kovachinskaya Formation in western Kamchatka

(Gladenkov et al., 1991).

Many well-preserved molluscan fossils have been de-

scribed from the upper Selandian to the lowest Thanetian

Katsuhira Formation in eastern Hokkaido, northern Japan

(Figure 1; Amano and Jenkins, 2014, 2017; Amano and

Oleinik, 2014: Amano et al., 2015a, b, 2016, 2018). These

fossils occur mainly in small calcareous concretions (about

30 cm in diameter) with plant debris, considered to be

wood-fall communities (Amano et al. ee 2018). From

the locality at Katsuhira along the Urahoro River, two

species of Boreocomitas including a new one have been

recovered. In this paper, we describe these species and

discuss their paleobiogeographical significance.

MATERIALS AND METHODS

Three specimens of Boreocomitas inouei new species and

one specimen of Boreocomitas species were obtained

from carbonate concretions (20 em in diameter) with

many bored wood fragments included in dark gray mud-

stones of the ravigulbitie Formation. These concretions are

exposed along the Urahoro River, 44 m south from the

mouth of the Kokatsuhirazawa River, Urahoro Town,

eastern Hokkaido (Figure 1; 42°59'10" N, 143°37'38" E).

Rocks in the direct proximity to the locality contain

dinoflagellate fossils indicating a late Selandian to ear-

liest mene ‘tian age (Amano et al., 2018). These species

were associated with 22 species of mollusks (Table 1) and

one species of echinoid. Among them, Myrtea ezoensis

(Nagao, 1938), and Thyasira oliveri Amano and Jenkins,

2018, were proposed as chemosymbiotic bivalves. As

mentioned by Amano et al. (2018), the paleobathymetry

of the Katsuhira Formation can be estimated as 200 to

500 m in depth. From these occurrences of mollusks, it is

possible to consider the two species of Boreocomitas as

members of the deep-sea wood-fall community.

All specimens of Boreocomitas are catalogue d in the

University Museum of the University of Tokyo ( UMUT).

The associated fauna is stored at Joetsu University of

Education (JUE). Classification at family level follows

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THE NAUTILUS, Vol. 132, No. 3-4

Okhotsk Sea

\ Hokkaido Island _-

- Study area

—42N

Pacific Ocean

Figure |

the current taxonomy of Conoidea by Bouchet et al.

(2011, 2017).

SYSTEMATIC PALEONTOLOGY

Class Gastropoda Cuvier, 1797

Order Neogastropoda Wenz, 1938

Superfamily Conoidea Fleming, 1822

Family Pseudomelatomidae Morrison, 1966

Remarks: Boreocomitas was originally proposed as

a subgenus of the genus Comitas Finlay, 1926, and placed

in Turriculinae Boxee 1 1942, of Turridae H. Adams and

A. Adams, 1853 by Hickman (1976). Family Pseudome-

latomidae including the genus Comitas is characterized by

a smooth paucispiral protoconch.

Katsuhira

A SS .

ad so°

Sy A / M

Map showing the locality yielding the two species of Boreocomitas discussed.

Genus Boreocomitas Hickman, 1976

Type Species: Comitas (Boreocomitas) oregonensis

Hickman, 1976

Remarks: Boreocomitas is characterized by having

a medium to moderately large-sized fusiform shell srati

axial nodes on the keel and fine spiral threads, which are

less prominent on its wide shoulder than anterior ward. Its

anal sinus is broad and moderately deep on the shoulder.

According to Hickman (1976), “the protoconch is missing

or worn on all specimens examined, but it is appare sntly

paucispiral”, The genus Comitas Finlay, 1926, has a more

slender shell, a narrower shoulder, a longer anterior canal

and weaker nodes on the periphery than in Boreocomitas.

It can be judged that these differences are enough to

separate the two taxa as distinct genera. Although the

kK. Amano et al., 2018

Page 119

Table 1. Mollusks associated species with Boreocomitas inouei

new species.

Species

Leionucula yotsukurensis (Hirayama)

Acila (Truncacila) hokkaidoensis (Nagao)

Ezonuculana aff. obsoleta Tashiro

Malletia poronaica (Yokoyama)

Menneroctenia plena Kalishevich

Pseudoneilonella? sp.

Tindaria paleocenica Amano and Jenkins

Propeamussium yubarense (Yabe and Nagao)

Limaria sp.

Myrtea ezoensis (Nagao)*

Thyasira oliveri Amano and Jenkins*

Astarte sp.

Cidarina? sp.

Naticidae gen. et sp. indet.

Kangilioptera sp.

Urahorosphaera kanekoi Amano and Oleinik

Admete katsauhiraensis Amano, Oleonik and Jenkins

Acteocina sp.

Retusa sp.

Biplica paleocenica Amano and Jenkins

Striodentalium sp.

Laevidentalium sp.

* Chemosymbiotic species.

pseudomelatomid genus Nekewis Stewart, 1927, re-

sembles Boreocomitas, Nekewis can be distinguished from

Boreocomitas by having a long and straight anterior canal

with a fasciole and ahelltoree anal sinus. Wenz (1938)

mistakenly considered Nekewis to be a synonym of the

raphitomid genus Clinura Bellardi, 1875. Clinura is very

similar to Boreocomitas in having biconic general outline

with distinct axial nodes on its periphery. However, Cli-

nura has a protoconch that is narrowly conical and di-

agonally cancellate. Its diagonally cancellate protoconch

demonstfate »s that C Pease belongs in Raphitomidae

Bellardi, 1875. Clinura also has an anal sinus whose apex is

located near the suture on the shoulder slope.

Boreocomitas inouei new species

(Figures 2, 3, 4)

Diagnosis: Moderate-sized, short, biconic Boreocomitas

species consisting of five and half whorls and at least one

smooth paucispiral protoconch. Surface sculptured with

23 spiral cords on base, nine on keel and six above wide

shoulder, and fine sinuous growth lines on last whorl. Apex

of anal sinus located at midpoint of shoulder.

Description: Shell moderate-sized, attaining 20.4 mm in

height, fusiform, consisting of 5.5 teleoconch whorls and

one protoconch whorl. Apical angle ranging from 59° to

73°. Protoconch smooth, ee and with large di-

ameter (d = 1.6 mm in paratype UMUT CM 32942). Last

whorl very large, occupying about 74% of shell height in

holotype; spire low; subsutural band very weak; shoulder

slope broad and gently concave. Surface of penultimate

whorl sculptured with one fine spiral cord below keel, six

on keel, shoulder slope smooth, without growth ae sand

21 axial nodes on keel: keel of last einan with 22 axial

nodes, 23 spiral cords on base, nine on keel, and six very

weak fine cords just above keel and five very faint on

shoulder below subsutural band: among spiral cords on

last whorl, two cords below keel stronger ae other cords.

Anal sinus moderately deep, its apex ‘erat d at midpoint

of shoulder slope on last whorl and just below subsutural

band on penultimate whorl. Aperture pyriform; outer lip

very thin; inner lip covered by thin, narrow callus. Anterior

canal short, slightly broken on anteriormost part, but

having slight siphonal fasciole.

Type Material: Holotype, UMUT CM 32793 (shell height,

20.4 mm-+: diameter, 13.5 mm): Paratype, UMUT CM

32794 (shell height, 9.0 mm; diameter, 5.9 mm); Paratype,

UMUT CM 32942 (diameter, 7.5 mm-+).

Type Locality: Cliff along Urahoro River, 44 m south

from mouth of Kokatsuhirazawa River, Urahoro Town,

eastern Hokkaido.

Remarks: This is the first record of Boreocomitas in the

Paleocene and from the northwestern Pacific region.

Boreocomitas inouei new species is similar to Bor-

eocomitas biconica (Hickman, 1976) from the middle to

upper Eocene Cowlitz Formation in northwestern Ore-

gon in having a similar size (shell height of B. biconica,

20.0 mm), a malcvely low spire and a Sane number of

nodes on the last whorl (20 in B. biconica). However, the

present new species has higher ratio of diameter (D)/shell

height (H) than the B. biconica species (D/H = 0.66 for B.

inouei: 0.53 in B. biconica). Also, unlike B. biconica, the

present species has a spiral cord below the periphery and

no beaded subsutural cords.

Some previously identified species belonging to the

now invalid genus Pleurotoma Lamarck, 1799. are now

allocated to other genera. Among them, Pleurotoma

(Pseudotoma) brevior von Koenen, 1885 (p. 35-36, pl. 2,

figs. 5a—c), was described from the Selandian of Copen-

hagen, Denmark. Later, this species was re-described as

Genotia brevior by Ravn (1939, p. 93-94, pl. 4, fig. Lla—b).

Judging from the size, outline, sculpture antl smooth

protoconch, P. brevior can be confidently allocated to

Boreocomitas. Boreocomitas brevior new combination

differs from B. inouei new species in having a more

slender, larger shell (ca. 30 mm in height), with fewer

nodes on the axial keel (18 to 19 in B. brevior) and distinct

spiral cords on the shoulder slope, and with more nu-

merous protoconch whorls (3.5 in B. brevior).

The genus Clinura Bellardi, 1875, has a similar shell

outline to Boreocomitas, with similar, moderately deep

anal sinus and many nodes on its keel. Among the species

of Clinura, Clinura sp. 1 from the Sonja Lens of the

Selandian Agatdal Formation in the western part of

Greenland (Nagssuaq) was illustrated by Kollmann and

Peel (1983, p. 97-98, fig. 220). This species is very similar

to the present new species in having a short biconic

outline, apex of the anal sinus located at the midpoint of

the broad shoulder slope, and very fine spiral cords above

Page 120

THE NAUTILUS, Vol. 132, No. 3-4

Figures 2-5.

32794: Qa.

3a. Aol view; 3b. Abapertural view; :

the keel. This species also has a smooth paucispiral

protoconch, which warrants its allocation to Boreocomitas,

not in Clinura. This species can be distinguished from B.

inouei new species by its larger shell (ca. 40 mm in he ight),

less numerous axial nodes, and by h: wing only one spiral

cord below the keel on the last whorl.

Etymology: The new species is named for Kiyokazu

Inoue (Obihiro), who collected the holotype.

Distribution: Known only from the type locality, in the

upper Selandian to lowe st Thanetian Katsuhira Forma-

tion, Urahoro Town, eastern Hokkaido.

Boreocomitas species

(Figure 5)

Description: Shell small, 13.1 mm in height, fusiform,

consisting of 5 teleoconch whorls; protoconch missing.

Apical angle 47°. Last whorl large, occupying about 69%

of shell he ight; spire high; subsutural band very weak;

shoulder slope broad and gently concave. Surface of

penultimate whorl sculptured with four fine spiral cords

below keel, three on keel, shoulder slope smooth, and 21

axial nodes on keel; keel of last whorl with 20 axial nodes,

12 spiral cords on base, three on keel, and no cord on

shoulder below subsutural band. Anal sinus moderately

deep, its apex located at midpoint of shoulder slope on last

whorl and just below subsutural band on penultimate

Boreocomitas species from the Katsuhira Formation. 2-4. Boreocomitas inouei new species. 2, Paratype, UMUT CM

Apical view; 2b. Lateral view showing deep anal sinus of growth lines above shoulder slope. 3. Holotype, UMUT CM 32793;

3c. Apical view. 4. Paratype, UMUT CM 32942, Apicz al view. 5. Boreocomitas sp., UMUT CM

32943; 5a. Apertural view; 5b. Side ae Be Apical view.

whorl. Aperture pyriform; outer lip very thin; inner lip

covered by thin, narrow callus. Anterior canal short.

Remarks: This species is similar to Boreocomitas inouei

new species as above described. However, Boreocomitas

species differs from B. inouei by having a slender form,

a higher spire, no spiral cord on the shoulder. Bor-

eocomitas oregonensis (Hickman, 1976), the type species

of the genus, is similar to Boreocomitas species in having

a rather higher spire. However, the type species has some

distinct spiral threads on its shoulder.

Distribution: Known only from the type locality of

Boreocomitas inouei new speices, in the upper Se landian

to lowest Thanetian Katsuhira Formation, Urahoro Town,

eastern Hokkaido.

DISCUSSION

The described species possibly lived in a wood-fall

community of upper bathyal depths. When Hickman

(1976) proposed Boreocomitas, this genus appeared to

be confined mainly to the middle Eocene to lower Oli-

gocene deposits in Oregon. As noted above, Gladenkov

et al. (1991) described and illustrated Comitas (Bor-

eocomitas) sp. from the upper Eocene Kovachinskaya

Formation in western Kamchatka. However, that shell is

poorly preserved, with weak nodes on the keel and

kK. Amano et al., 2018

ave 119)

Page 12]

a narrow shoulder. It is uncertain whether this species

can be classified in Boreocomitas. Our study shows that

Boreocomitas dates back to the Selandian and it had

a broad distribution including Denmark, westem Greenland

and eastern Hokkaido.

The aporrhaid gastropod Kangilioptera Rosenkranz,

1970 shows the same geographical pattern to Bor-

eocomitas. Kangilioptera has been recorded from the

upper Danian Kangilia Formation in western Greenland,

the Selandian Kerteminde Marl in Denmark and the

upper Selandian to lowest Thanetian Katsuhira Formation

in eastern Hokkaido (Rosenkranz, 1970; Kollmann and

Peel, 1983; Amano and Jenkins, 2014; Schnetler and

Nielsen, 2018) (Table 2).

The astartid bivalve Astarte paleocenica Amano and

Jenkins in Amano et al., 2018 from the Katsuhira For-

mation is similar to A. parvula Kalishevich in Kalishevich

et al., 1981, from the Danian Krasnoy arskaya Formation

in southeastern Sakhalin and A. trigonula Koenen, 1885,

from the Selandian Lellinge Gre end (see also Amano

et al. 2018: Schnetler, 2001).

The bivalve genus Conchocele Gabb, 1866, has its

earliest confirmed occurrences in the latest Cretaceous

(Maastrichtian) of Antarctica, the Danian of western

Greenland, and the Thanetian of Spitsbergen

(Rosenkranz, 1970; Little et al. 2015; Hryniewicz et al.,

2016, 2017). In Spitsbergen, the genus survived to the late

Eocene-early Oligocene (Thiedig et al., 1980). There are

so far no woneiainel records of @onchucele from the

Cretaceous and Paleocene of the northern Pacific area.

However, some publications suggest it could have been

present in the Pacific during the Late Cretaceous. Con-

chocele cretacea has been recorded from the Upper

Cretaceous deposits from Vancouver Island area

(Whiteaves 1874, jo, Wi, figs. 2. Der). Thyasira cretacea

(= Conchocele cretacea) has also been recorded from the

Coniacian(?) Funks Formation of Sacramento Valley area

in California (Anderson, 1958). Therefore, the absence of

Conchocele in the Upper Cretaceous and Paleocene of

northern Pacific area might be just an artifact of the fossil

record. The genus became very common in the northern

Pacific region after the Paleocene (Hickman, 2015:

Hryniewicz et al., 2017). This distribution pattern shows

that Conchocele populations could have interchanged

between the northern Pacific and the North Atlantic/

Arctic regions at least by the late Eocene, or probably

earlier.

Moreover, the oldest record of the deep-sea arcid bi-

valve genus Bentharca Verrill and Bush, 1898, has been

found in the upper Selandian to lowest Thanetian Kat-

suhira Formation in eastern Hokkaido (Amano et al.,

2015). As pointed by Amano et al. (2015), Barbatia (Acar)

hennigi Heinberg, 1978, from the Maastrichtian to

Danian deposits in Denmark is possibly an ancestor of this

genus.

Adding to Boreocomitas, the four deep-sea mollusks

above mentioned occur in both northern Pacific and

North Atlantic/Arctic, indicating the sea connection be-

tween both areas during the Paleocene, although most

paleontologists believe that the Arctic was isolated from

the Pacific from the Albian to the latest Miocene

(Marincovich et al., 1990, 2002: Thiede et al., 1990:

Marincovich, 1993: Marincovich and Gladenkov, 1999:

Beard and Dawson, 1999). Deep-sea genera commonly

have broad geographic distribution (e.g. Amano et al.

2015a, b; Amano and Jenkins 2017), and similarities be-

tween the fauna of the Paleocene Katsuhira Formation

from Hokkaido and that of the North Atlantic/Arctic

Paleocene mollusk faunas could be partially caused by

that. However, an alternative explanation is that the

Bering Strait was to some extent open and connected

northern Pacific and Arctic oceans, allowing for faunal

interchange between both areas during the Selandian to

late Eocene time. When Brikiatis (2014) reviewed the

early Cenozoic paleogeography around the Arctic region

based on the land mammals and flora, he showed no land

bridge around the Arctic Ocean during the Selandian

in ie Figure 8. Gleason et al. (2009) also reconstructed

the early Eocene paleogeographic map without any

land bridge surrounding the Arctic Ocean, based on the

Table 2. Taxa of Paleocene faunas shared between the Northwestern Pacific, Arctic and Northem Atlantic regions.

Region NW. Pacific

E. Hokkaido — SE. Sakhalin Ellesmere Is.

Taxa Age L. Selandian— Danian Paleocene

E. Thanetian

Bentharca +

Small Astarte + +

Thyasira oF =P

Conchocele

Kangilioptera tr

Drepanocheilus + +

Aporrhaidae

Boreocomitas +

Reference This study Kalishevich Marincovich and

(1981) Zinsmeister

(1991)

Arctic/ N.Atlantic

Spitsbergen Is. W. Greenland

Denmark

Selandian Danian— Danian—

Selandian Selandian

ata

+ +

> +

Rosenkranz (1970), Ravn (1939), Heinberg

Kollmann and (1999), Schnetler

Peel (1983) vat Nielsen (2018)

Hryniewicz

et al. (2016)

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THE NAUTILUS, Vol. 132, No. 3-4

examination of Nd-Sr isotopes in fossil fish debris from

Lomonosov Ridge. Although these data support our hy-

pothesis on the Paleogene temporary opening of the

Bering Strait, more data are needed to eaten it.

ACKNOWLEDGMENTS

We acknowledge Alan Beu (GNS Science) and Geerat

I[- Vermeij (UC Davis) for providing comments to an

earlier version of the manuscript. We thank Carole S

Hickman (UC Berkeley) for her information on the type

specimens of Boreocomitas species. We thank Anton E.

Oleinik (Florida Atlantic University) and Kai I. Schnetler

(Denmark) for their information on fossil species from the

Paleogene in Kamchatka and Denmark. We also express

many thanks to Kiyokazu Inoue (Obihoro in eastern

Hokkaido) who found the holotype and kindly donated

Boreocomitas specimen. This study was supporte d_ by

a Grant-in-aid for Scientific Research from the Japan

Society for Promotion of Science (C, 17K05691, 2017—

2019) to KA and RG]. Financial support to KH was

provided by the Polish National Science Centre (NCN)

research grant 2014/15/B/ST10/04886 “The influence

of Paleocene/Eocene Thermal Maximum on oceanic

chemosynthesis-based ecosystems”.

LITERATURE CITED

Adams, H. and A. Adams. 1853. The genera of recent Mollusca.

John Van Voorst, London.

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

Aporrhaidae (Gastropoda) from eastern Hokkaido, Japan.

Paleontological Research 18: 33-39.

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

bivalves from Urahoro Town in eastern Hokkaido,

Northern Japan. Journal of Conchology 42: 283-297.

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

Mesozoic-relict mollusks from Paleocene wood-fall com-

munities in Urahoro Town, eastern Hokkaido, northern

ee Journal of Paleontology 22: 105-111.

Amano, k., R.G. Jenkins and K. Nishida. 2015a. A new Pa-

eee ne species of Bentharca (Bivalvia; Arcidae) from

eastern Hokkaido, with remarks on evolutionary adaptation

of suspension feeders to the deep Sea. Paleontological

Research 19: 128— 138.

Amano, K., C.T.S. Little, K.A. Campbell, R.G. Jenkins, and K.P.

Saether. 2015b. Paleocene and Miocene Thyasira sensu

stricto (Bivalvia: Thyasiridae) from chemosynthetic com-

munities from Japan and New Zealand. The Nautilus 129:

43-53.

Amano, K. and A. Oleinik. 2014. A new genus of Buccinoidea

(Gastropoda) from Paleocene deposits in eastern Hokkaido,

Japan. The Nautilus 128: 122-128.

Amano, K., A. Oleinik and R.G. Jenkins. 2016. A new species of

Admete (Gastropoda: Cancellariidae: Admetinae) from the

Paleocene of eastern Hokkaido, northern Japan. The

Nautilus 130: 116-121.

Anderson, F.M. 1958. Upper Cretaceous of the Pacific Coast.

Geological Society of America Memoir 71: 1-378.

Bellardi, L. 1875. Novae pleurotomidarum Pedimonti et Lig-

uriae onsthovine dispositionis prodromus. Bollettino de Ta

Societa Malacologica Italiana 1: 16-24.

Beard, K.C. and M.R. Dawson, 1999. Intercontinental dispersal

of Holarctic land mammals near the Paleocene-Eocene

boundary: Paleogeographic, Plaeoclimatic and biostratigraphic

implications. Bulletin de la Société Géologique de France 170:

697-706.

Bouchet, P., Yu. I. Kantor, A. Sysoev, and N. Puillandre. 2011. A

new operational classification of the Conoidea (Gastro-

poda). Journal of Molluscan Studies 77: 273-308.

Bouchet, P., J.-P. Rocroi, B. Hausdorf, A. Kaim, Y. Kano, A.

Niitzel, P. Parkhaev, M. Schrédl, and E.E. Strong. Sie

Revised classification, nomenclator and typification of ¢; gastropod

and monoplacophoran families. Malacologia 61: 1-526.

Brikiatis, L. 2014. The De Geer, Thulean and Beringia routes:

key concepts for understanding early Cenozoic bio-

geography. Journal of Biogeography 41: 1036-1054.

Finlay, H.J. 1926. New shells from New Zealand Tertiary Beds.

Transactions and Procee dings of the Royal Society of New

Zealand 56: 55-60.

Fleming, J. 1822. The philosophy of zoology: or a general view of

the structure, functions and _ classification of animals.

Edinburgh 1: I-LII 1-432. 2: 491.

Gabb, W.M. 1866. Pale ontology of California. Vol. 2. Cretaceous

and Tertiary fossils. Sect. 1 (Tertiary Invertebrate Fossils).

124 pp. Geological Survey of California, Philade Iphia.

Gladenkov, Yu.B., V.N. Sinelnikova, A.E. Shantser, A.I.,

Chelebaeva, A.E. Oleinik, L.V. Titova, G.M. Bratseva,,

N.A. Fregatova, E.V. Ziryanov, and K.G. Kazakov. 1991.

The Eocene of western Kamchatka. Transactions of the

Geological Institute, Academy of Sciences of the USSR

467: 1-182. (in Russian).

Gleason, ].D., D.J. Thomas, T.C. Moore Jr., J.D. Blum and R.M.

Owen. 2009. Early to middle Eocene history of the Arctic

Ocean from Nd-Sr isotopes in fossil fish de bris, Lomonosov

Ridge. Paleoceanography 24: PA2215.

Heinberg, C., 1978. Bivalves from the white chalk (Maas-

trichtian) of Denmark, II: Arcoida. Bulletin of the Geo-

logical Society of Denmark 27: 105-116.

Hickman, C.S. 1976. Bathyal gastropods of the Family Turridae

in the early Oligocene Keasey Formation in Ore gon, with

a review of some dee p-water genera in the Paleoge one of the

eastern Pacific. Bulletins of American Paleontology 70:

5-119.

Hryniewicz, K., M.A. Bitner, E. Durska, J. Hagstroém, H.R.

Hjalmarsdottir, R.G. Jenkins, C.T.S. Little, Y. Miyajima,

H.A. Nakrem and A. Kaim. 2016. Paleocene methane seep

and wood-fall marine environments from Spitsbergen,

Svalbard. Palaeogeography, Palaeoclimatology, Palaeoecology

462: 41-56.

Hryniewicz, K., K. Amano, R.G. Jenkins, and S. Kiel. 2017.

Thyasirid bivalves from Cretaceous and Paleogene cold

seeps. Acta Palaeontologica Polonica 62: 705-729.

Kalishevich, T.G., E. D. Zaklinskaya, and M.Ya. Serova. 1981.

Development of organic life of the Pacific belt on the

boundary between the Mesozoic and Cenozoic. Foramin-

ifers, mollusks and palynoflora of North-West Sector: 164 p.

Nauka, Moscow. (in Russian, title translated).

Keene n, A. von. 1885: Ueber eine Paleociine Fauna von

Kopenhagen. Abhandlungen der kéniglichen Gesellschaft

der Wissenschaften (Gottingen) 32: 1-128.

Kollmann, H.A. and J.S. Peel. 1983. Paleocene gastropods from

Niigssuaq, West Greenland. Bulletin Gronlands Geologiske

Undersggelse 146: 1-115.

Lamarck, J. B. P. A. de M. 1799. Prodrome dune nouvelle

Classification des Coquilles. Mémoires de la Sociéte

Naturelle de Paris, An. VIII: 63-91.

kK. Amano et al., 2018

Little, C.T.S., D. Birgel, A.J. Boyce, A.J. Crame, ].E. Francis, S.

Kiel, Ife Peclomanns D. Pirrie, G.K. Rollinson, and J.D. Witts.

2015. Late Cretaceous (Maastrichtian) shallow water hy-

drocarbon seeps from Snow Hill and Seymour Islands,

James Ross Basin, Antarctica. Palaeogeography, Palaeo-

SE Palaeoecology 418: 213-228.

Marincovich, Jr. 1993. Danian mollusks from the Prince

Creek Beas. Northem Alaska, and implications for

Arctic Ocean paleogeography. Paleontological Society

Memoir 35: 1—35.

Marincovich, L., Jr., K.B. Barinov, and A.E. Oleinik. 2002. The

Astarte (Bieler Astartidae) that document the earliest

opening of Bering Strait. Journal of Paleontology 76:

239-245.

Marincovich, L. Jr., E.M. Brouwers, D.M. Hopkins, and M.C.

Mckenna. 1990. Late Mesozoic and Cenozoic paleo-

geographic and paleoclimatic history of the Arctic Ocean

Basin, based on shallow-water marine faunas and ter-

restrial vertebrates. In: A. Grantz, L. Johnson, and J.F.

Sweeney (eds.), The Arctic Ocean Region. Geological

Society of America, The Geology of North America,

Volume L, pp. 403-426.

Marincovich, L., Jr. and A. Yu. Gladenkov. 1999. Evidence

for an early opening of the Bering Strait. Nature 397:

149-151.

Morrison, ].P.E. 1966. On the families of Turridae. The

American Malacological Union. Annual Reports, for 1965,

Powell, A.W.B. 1942. The New Zealand Recent and fossil

Mollusca of the family Turridae. With general notes on

turrid nomenclature and systematics. Bulletin of the

Auckland Institute and Museum 2: 1-192.

Prothero, D.R. 2001. Chronostratigraphic calibration of the

Pacific coast Cenozoic: a summary. In: D. R. Prothero (ed.),

Magnetic stratigraphy of the Pacific coast Cenozoic, Pacific

Section SEPM, “eal 91, pp. 377-394.

Ravn, J.P.J. 1939. Etudes sur les mollusques du Paléocéne de

Copenhague. Det Kongelige Danske Videnskabemes Sel-

skab. Biologiske Skrifter 1: 1-106.

Rosenkrantz, A., 1970. Marine upper Cretaceous and lowermost

Tertiary deposits in west Greenland. aus tin of the Geo-

logical oe ty of Denmark 19: 406-45:

Schne tle r, K.I. 2001. The Selandian (Paleoce ae molluse fauna

from Cope snhagen: the Poul Harder 1920 Collection. Geo-

logy of Denmi ma Survey Bulletin: 37: 1-85.

Samnatilor: K.I. and M. S. Nielsen. 2018. A Palaeocene (Selan-

dian) molluscan fauna from boulders of Kerteminde Marl in

the gravel-pit at Gundstrup, Fyn, Denmark.

Research 18: 3-81.

Stewart, R. B. 1927. Gabb’s California fossil type | gastropods.

Proceedings oi the Academy of Natural Sciences of Phil-

adelphia 78: 287-447.

Thiede, ]., D.L. Clark, and Y. Herman. 1990. Late Mesozoic and

Cenozoic paleoceanography of the northern polar oceans.

In: A. Grantz, L. Johnson and J. F. Sweeney (eds.), The

arctic Ocean Region. Geological Society of America, The

Geology of North America, Vallone L, pp. 427-457.

Thiedig, F., C.A.G. Pickton, U. Lehmann, W.B. Harland, and

H. J. Anderson. 1979. ie ee von Renardodden (éstlich

Kapp Lyell, Westspitzbergen, Svalbard). Mitteilungen der

Ge ologisch- Palaontologisches Institut der Universitiit

Hamburg 49: 135— 146,

Vernill, AE. and K.J. Bush. 1898. Revision of the deep-water

Mollusca of the Atlantic coast of North America, with

descriptions of new genera and species. Proceedings of the

United States National Museum 20: 775-932.

Wenz, W. 1938. Gastropoda. Handbuch der Palaéozoologie, Band

6. Gebriider Borntraeger, Berlin-Nikolassee, 1639 pp.

Whiteaves, G.F. 1874. Notes on the Cretaceous fossils collected

by Mr. James Richardson, at Vancouver and the adjacent

islands. Ge ological Survey of Canada Report of Progress for

1873-74: 5-18.

Cainozoic

THE NAUTILUS 132(3-4):124-130, 2018

Page 124

Three new species of the genus Humboldtiana (Gastropoda:

Pulmonata: Humboldtianidae) from Mexico

Omar Mejia’

Benjamin Lopez

Departamento de Zoologia, ENCB-Instituto

Politécnico Nacional

Prolongacion de Carpio y Plan de Ayala s/n

C.P. 11340 Ciudad de México, MEXICO

José Ma. Reyes-Gomez

Centro de Cultura Ambiental y Biodiversidad,

Bolanos, Jalisco, MEXICO

ABSTRACT

Three new species of the genus Humboldtiana are described.

The presence of a mantle mottled with dark spots allocates H

paquimei from Chihuahua to the subgenus Gymnopallax. On the

other hand, the embryonic whorls, a smooth and granular

sculpture distributed over the shell surface, allocate H. wixarika

from Jalisco and H. aurea from Chihuahua to the subgenus

Humboldtiana s.s. The newly described species are distinguishe d

from other species of the genus by the combination of die ll and

anatomical characters and by their geographic distribution.

Additional Keywords: Taxonomy, land snail, neotropical region

INTRODUCTION

The land snail genus Humboldtiana von thering 1892 (see

Pilsbry, 1927) is comprised by more than 50 species

patchily distributed from the southwestern United States

to central Mexico (Thompson, 2011). With the exception

of three “widely distributed species”, H. buffoniana, H.

nuevoleonis, and H. durangoensis, most of the species are

microendemic and known only from the type locality. The

classification followed herein was proposed by Thompson

(2011). It recognizes six subgenera based on shell and

reproductive anatomy: Polyomphala Thompson and

Brewer, 2000; Gymnopallax Thompson, 2006; Clydo-

nacme Thompson, 2006; Aglotrochus Thompson, 2006;

Oreades Thompson and Brewer, 2000; and Hum-

boldtiana Thompson and Brewer, 2000. The subgenus

Polyomphala includes species with depressed shells with

nodular tubercles and dart bulbs exposed; the subgenus

Gymnopallax is composed of species with mottled

mantle with dark gray spot, and shell apex usually de-

collated; the subge nus Clydonacme includes species

with an embryonic shell sculpture consisting of wavy

transverse striations and granular sculpture re stricted to

the first postembryonic whorl; the subgenus Aglotrochus

Author for correspondence: [email protected]

comprises species that completely lack the granular

sculpture; species in the subgenus Oreades do not have

the dart apparatus; and the subge snus Humboldtiana is

composed of species with e »mbryonic whorls present (not

decollated) and a smooth and granular sculpture dis-

tributed over the entire shell surface. Despite the

presence of a more or less conserved pattern that sug-

gests morphostatic radiation (Gittenberger, 1991), it is

difficult to find morphological synapomorphies for the

entire genus owing to the fact that the granular sculpture

is Angst nt in certain species (Thompson and Brewer,

2000; Thompson, 2006). Nevertheless, the molecular

phylogeny clearly supports the monophyly of the genera,

although not the monophyly of the subgenera (Mejfa and

Zuniga, 2007). Despite of this, we manere to the current

classification until a new review of the genus is available.

Authorities for the new species are as follows: Hum-

boldtiana wixarika new species Mejia, Lopez, and

Reyes-Gomez; Humboldtiana aurea new species Mejia

and Lé6pez; and Humboldtiana paquimei new species

Mejia and Lopez.

MATERIALS AND METHODS

De spite being medium- to large-sized land snails, the

species of ae genus Humboldtiana are ve ry elusive and

hard to collect. In fact, with selected exceptions noted,

most of the species previously named have been described

based on very few individuals. For this reason, the de-

scriptions of new species herein are based on the holotype

and one or two paratypes only. Shell description and

morphological measurements were performed according

to Thompson and Brewer (2000). In all cases, the first

measurement (or counts, for the number of whorls) is

from the holotype and the measurements in parentheses

are from paratypes | and 2, respectively. The types of the

newly described species are deposited at Coleccién

Nacional de Moluscos (CNMO), Instituto de Biologia,

UNAM, Ciudad de México. Color photographs of the

currently described species shells are available at https://

osf.io/3cswm.

O. Mejia et al., 2018

SYSTEMATICS

Family Humboldtianidae Pilsbry, 1939

Genus Humboldtiana von Ihering, 1892

Humboldtiana wixarika new species Mejia, Lopez,

and Reyes-Gomez

(Figures 1-4, 13)

Diagnosis: Medium-sized Humboldtiana with pale-brown

shell bearing three equal-sized dark brown bands clearly

visible on internal shell surface. Presence of short verge

composed by four lobes and penis with two thick longi-

tudinal fleshy columns distinguish this species from mihee

species in subgenus Humboldtiana, where dart glands are

separated from dart sacs by distance equal to or longer than

glands length.

Description: SHELL ( Figures 1—4) globose, outer lip not

reflected, pale brown, with three chestnut to dark-brown

bands of same size, first band often lighter in color toward

end of embryonic whorl, 4 whorls (4.1, 4.0). Embryonic

shell caramel in color, with 1.7 whorls (1.8, 1.7), first whorl

without sculpture, then with almost imperceptible growth

lines. Sculpture of rest of shell consisting of white growth

lines more evident toward shell aperture and randomly

distributed small oval granules absent toward aperture.

Umbilicus completely covered by basal portion of

10 mm

Figures 1-4. Humboldtiana wixarika new species. Holotype, CNMO 756. 1) Apertural view 2) Apical view 3) Oblique basal view 4)

Detail of embryonic whorls.

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THE NAUTILUS, Vol. 132, No. 3-4

peristome. Parietal callus thin, transparent to white. Shell

height 37 mm (36, 38); shell diameter 39 mm (43, 45);

aperture height 28 mm (33, 29); aperture diameter 26 mm

(25, 29).

REPRODUCTIVE ANATOMY (Figure 13): Penis short 8 mm

(6.5, 7.2), bulbous to globose, tapering at distal end, penis

lumen with two thick columns, short verge extending to

first third of penis and composed of four lobes. Penis

retractor muscle 14 mm long (14, 15). Epiphallus long,

cylindrical, measuring 38 mm (46, 34). Flagellum short,

51 mm inle ngth (44, 59), near 0.85 times (1.18, 0.67) the

combined length of penis plus epiphallus. Atrium short,

1.28 mm (2.27, 1.65). Lower vagina short and slightly

shorter than half penis size, 3.76 mm _ (3.76, 3.40),

extending to region of dart sacs; four dart sacs approxi-

mately of same size: ds] (2.41 mm), ds2 (2.33 mm), ds3

(2.70 mm), and ds4 (2.07 mm) (3.14, 3.02, 4.00, 3.38)

(2.53, 2.17, 3.40, 2.56). Median vagina short, bearing four

dart glands forming a ring, dart glands reaching a maxi-

mum height of 2.91 mm (3.24, 3.31) and separated from

dart sacs by distance of 2.55 mm (2.8, 2.11). Spermathecal

duct §1 mm length (128, 114), spermatheca with caecum

of 11 mm le meth ( (8, 8) adhering to the albumen gland,

elongated and sac-shaped, witty a length of 11 mm

(10, 7).

Type Material: Holotype: CNMO 7561; collected 29

August 2017 by José Maria Reyes Gomez; Paratypes:

CNMO 7562 (2): same data as the holotype. All from type

locality.

Type Locality: Jalisco, km 29 Carretera Bolanos, Tux-

pan, 11.1 km northwest of Bolaios, Jalisco, 2454 m al-

titude (21°54'04” N, 103°51/33” W). The individuals were

collected on the ground in a pine-oak forest.

Distribution: Known only from type locality.

Remarks: The subgenus Humboldtiana comprises a very

disparate group of species where the embryonic whorls

are smooth (Thompson, 2006). In this subge snus, three

groups of species have been recognized, the Hum-

boldtiana buffoniana species group in which the dart

glands are located just above the dart sacs, the Hum-

boldtiana bicincta species group where the dart sac ap-

paratus is reduced, and the Humboldtiana texana species

group where the dart glands are separated from the dart

sacs by a distance equal or higher than the length of the

dart sacs and the epiphallic chamber is absent (Thompson

and Brewer, 2000). Humboldtiana wixarika new species

belongs to the Humboldtiana texana species group. By

having a short verge with four digitiform lobes, this species

is similar to H. fasciata (Burch and Thompson, 1957) from

the state of Hidalgo, Mexico, but in contrast to H. fasciata,

H. wixarika prese Ss two thick columns in the interior of

the penis instead seven longitudinal folds. Besides, this

latter feature, the verge, and the median vagina in H.

wixarika are smaller than in H. fasciata. On “ihe ether

hand, the shape of the penis of H. wixarika is similar to

that of H. balanites from the state of Chihuahua, Mexico,

but in this last species, the verge is multilobed and its size

is almost the size of the penis, in opposition to H. wixarika,

where the verge is short. Additionally, although in both

species the atrium is short, it is quite evident in H. wix-

arika, while in H. balanites it is barely distinguishable.

Etymology: The specific epithe t, anoun in apposition, is

dedicated to the Wixarika people, better known by their

Spanish name, Huicholes, a brave people that reject

succumbing to the pressures of the modern world.

Humboldtiana aurea new species Mejia and Lopez

(Figures 5-8, 14)

Diagnosis: Medium-sized Humboldtiana with thin and

fragile shell. The new species is similar to other species in

tne. Humboldtiana texana species complex by having

protruding dart bulbs at the base of the dart sacs chile

differing from the rest of the species in the group by

possessing four sub-equal dart sacs and lacking a caecum

on the spermathecal duct. The description is based on the

holotype and two paratypes.

Description: SHELL (Figures 5-8) small, subglobose and

thin, outer lip slightly thickened, pale-brown to golden

with three clearly defined dark-brown bands, in certain

specimens bands fade toward the aperture, with first and

second band of same width, third band narrower, 4.1

whorls (3.2, 3.2). Embryonic shell with 1.2 whorls (1.1,

1.3), paler than rest of shell. First whorl without sculpture,

remainder of whorls with few, weak growth lines. Re-

mainder of shell sculptured with growth lines and small

ovate granules absent on umbilicus. Umbilicus partially

cove ted by basal portion of peristome. Callus thin,

transpare nt. Shell height 25 mm (25, 23); shell diameter

29 mm (31, 28); aperture height 19 mm (17, 16), aperture

diameter 19 mm (18, 18).

RepropuctivE ANaTomy (Figure 14): Penis short and

globose 7.61 mm (6.94, 7.98), penis with verge extending

els penis length, composed of three skirts of tissue, one of

them with three digitiform processes. Penis retractor

muscle 5 mm (5.77, 5.2). Epiphallus short and cylindrical

20 mm (19.2, 20.0). Flagellum short 36 mm (38, 32),

nearly 1.2 times (0.68, 0. 87 ) the combined length of penis

plus ‘epiphallus. Atrium abbreviated and ree ly distin-

guishable. Lower vagina 5.82 mm (4.29, 5.32), exte nding to

region of dart sacs; fa sub-equal dart sacs ds] (3.22 mm),

ds2 (3.73 mm), ds3 (2.81 mm) and ds4 (2.30 mm) (2.39,

2.47, 1.98, 1.68) (2.35, 2.95, 2.81, 2.56). Dart bulbs

embedded in vagina wall, forming conspicuous bulges.

Median vagina sibbrreutetedl, bearing ring of four aia

glands, dae glands reaching maximum height of 3.80 mm

(3. 05-3.39) arn separatec alarm dart sacs =by distance of

1.83 mm length (2.86-2.81). Spermathecal duct short,

47mm (37-50) and without a caecum. Spermatheca

elongated and sac-shaped, 9 mm length (7.54—9.81).

Type Material: Holotype: CNMO 7563; collected 9

August 2007 by Omar Mejia. Paratypes: CNMO 7564

(2): same data as the holotype. All from type locality.

O. Mejia et al., 2018

Page 127

10 mm

Figures 5-8. Humboldtiana aurea new species. Holotype, CNMO 7563. 5) Apertural view 6) Apical view 7) Oblique basal view 8)

Detail of embryonic whorls.

Type Locality: Chihuahua, E] Oro, municipality of Casas

Grandes, near the border with the State of Sonora, 67 km

southwest of Nuevo Casas Grandes, Chihuahua, 2054 m

altitude (30°09’ 35” N, 108°33'56” W). The specimens

were collected on the rocks next to a small waterfall,

where the predominant vegetation is an oak-pine forest.

Distribution: Only known from the type locality.

Remarks: Humboldtiana aurea new species belongs to

the H. texana species group based on its dart glands

separated from the dart sacs. Two other species of the

group had been described from the state of Chihuahua, H.

balanites (Thompson, 2006) and H. corruga (Thompson

and Mejia, 2006). Humboldtiana aurea is similar to H.

balanites as both have a thin shell, an abbreviated atrium,

and bulges on the dart sac caused by the underlying dart

bulbs; nevertheless, the light shell color with dark-brown

bands of H. aurea contrasts with the dark-brown ground

color with black bands of H. balanites. In addition, H.

balanites has a multilobed verge, a long flagellum two

times longer than the combined length of the penis plus

epiphallus, a spermathecal duct that is twice the length of

that present in H. aurea and, last but not least, H. balanites

presents a short caecum in the spermathecal duct that is

absent in H. aurea. On the other hand, H. aurea resembles

H. corruga in size, but the shells of H. corruga are darker

in color. The reproductive anatomies of H. corruga and H.

aurea are similar in that both species have ware that

protrude at the basis of the dart sacs, and both also have

a short flagellum that is of same size as the combined

length of the penis plus epiphallus; nevertheless, H.

corruga differs from H. aurea by having a slenderer

atrium, larger penis, dart sacs of the same size, a globular

spermatheca, and a spermathecal duct that with a caecum.

Page 128

THE NAUTILUS, Vol. 132, No. 3-4

Etymology: The specific epithet, aurea, alludes to the

name of the type locality, “El Oro” (“The Gold”) where

the species was collected.

Humboldtiana paquimei new species Mejia and Lépez

(Figures 9-12, 15)

Diagnosis: Medium-sized Humboldtiana. The new

species has a similar shell to H. regula (Thompson, 2006),

but differs in reproductive anatomy particularly on the

verge, the length of the epiphallus and the spermathecal

ain. The description is based on the holotype and one

paratype.

Description: SHELL (Figures 9-12) large, external lip not

reflected, background color pale- Prous becoming darker

toward aperture. Three well-defined chestnut- to dark-

brown bands, third one wider than first and second. 4.1

Whorls (4.1). sEmbsyonte shell lighter than remainder of

2 (1.4) whorls and with sculpture of weak

shell, with 1

wavy striations. Sculpture of second whorl with small

granules: remainder of shell with striations and white

eerie Umbilicus almost completely covered by basal

portion of ae Callus very thin, transparent. Shell

height 37 mm (31); shell diameter 38 mm (33); aperture

height 25 mm (21); aperture diameter 23 mm (20).

REPRODUCTIVE ANATOMY (Figure 15): Penis short 7.91

mm (7.24), slender at base and swollen in middle; short

and stout verge of approximately one-third penis length

and consisting of two tissue folds with digitiform

processes. bens retractor muscle 3.04 mm long (7.7).

Epiphalus long, cylindrical and stout, measuring 34 mm

(29), four times penis length. Flagellum short 60 mm in

length (62), nearly 1.4 times (1.7) eombined length of penis

plus epiphallus. Atrium short and bare ly perceptible. Lower

vagina of almost same length as penis, measuring 6.13 mm

(5.68) and extending to region of dart sacs; four dart sacs of

different sizes, ds] largest, remainder decreasing in size,

Figures 9-12.

view 12) Detail of embryonic whorls.

Humboldtiana paquimei new species. Holotype, CNMO 7565. 9) Apertural view 10) Apical view 11) Oblique basal

O. Mejia et al., 2018

Ds > IC

Page 129

gen atr

5mm

Figure 13. Reproductive anatomy of Humboldtiana wixarika

new species. Abbreviations. cae :spermathecal cae cum; dgla: dart

glands;ds; dart sacs; epi: epiphallus; fla: flagellum; gen atr; genital

atrium pen: penis; pr: penis retractor; spd: spermathecal duct:

spt: spermatheca; vag: vagina; vd: vas deferens.

spt

spd

pen

gen atr

Figure 14. Reproductive anatomy of Humboldtiana aurea

new species. Abbreviations. dgla: dart glands;ds; dart sacs; epi:

epiphallus; fla: flagellum; gen atr; genital atrium pen: penis; pr:

penis retractor; spd: spermathecal duct; spt: spermatheca; vag:

vagina; vd: vas deferens.

spt

fla

aan . i epi

ds"

vag

gen atr

‘18

Figure 15. Reproductive anatomy of Humboldtiana paquimei

new species. Abbreviations. cae :spe srmathecal caecum: dgla: dart

glands;ds; dart sacs; epi: epiphallus; fla: flagellum; gen atr; genital

atrium pen: penis; pr: penis retractor; spd: spermathecal duct; spt:

spermatheca; vag: vagina; vd: vas deferens.

dsl (4.12 mm), ds2 (3.51 mm), ds3 (3.02 mm) and ds4

(3.77 mm) (4.25, 4.08, 3.0, 3.5). Median vagina bearing four

dart glands that form ring just above Gime sacs arg] wal

maximum height of (3.07mm) (3.02). Spermathecal duct

long, measuring 112 mm (103), bearing short caecum of

8 mm length (6). Spermatheca sac-shaped, adhered to

albumen gland, measuring 7 mm (10).

Type Material: Holotype: CNMO 7565; collected 9

August 2007 by Omar Mejia; Paratype: CN MO 7566 (1):

same data as the holotype. All from type locality.

Type Locality: Chihuahua, bridge over Rio Piedras Verdes.

Approximately 1 km Southeast Bh El] Willy, municipality of

Casas Grandes, Chihuahua, 1829 m altitude (30°10'29" N,

108°18'40" W). Individuals were collected on the south wall

of the canyon that surround the Piedras Verdes river, the

predominant vegetation is an oak-pine forest.

Distribution: Only known from the type locality.

Remarks: The wavy sculpture of the embryonic whorl

and the granular sculpture restricted to the first post-

enilraworante assigns the new species to the subgenus

Clydonac me, a taxon so far limited to the State of Chi-

huahua that comprises six currently recognized species

(Thompson, 2006). The shell of the new species is similar

to H. regula (Thompson, 2006), but differs in three re-

productive traits. In H. paquimei, the verge is stout and

consisting of two tissue folds, in contrast to H. regula,

where the verge is composed of a thin skirt of tissue. In

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THE NAUTILUS, Vol. 132, No. 3=4

H. paquimei, the epiphallus is four times larger than the

penis, and in H. regula, the epiphallus is twice the length

of the penis. Last but not least, in H. paquimei, the

spermathecal duct plus the spermatheca is three times

longer than in H. regula. Two shells deposited at the Florida

Museum of Natural History (UF 185700 and UF103138)

may be H. paquimei based on geographical proximity. These

two specimens unfortunately alk reproductive anatomy and

were excluded from the de scription.

Etymology: Paquimé was an ancient culture that de-

veloped in northwestern Mexico between the years 700

and 1475. The maximum expression of this culture. is

found in the municipality of Casas Grandes, Chihuahua.

The name of the new species honors this culture and the

archaeological zone of Paquimé, Intangible Cultural

Heritage of Humanity since 1998.

ACKNOWLEDGMENTS

We are grateful to César Guzman for taking the photo-

graphs, to Alejandra Rocha- poe z for the illustrations of

the reproductive anatomy, to John Slapcinsky at Uni-

versity of Florida Museum a Zoology who provided

some photos of H. regula and two anonymous reviewers

for their comments. This work was pé arti: ly funded by

Conacyt project number 165990. This work is dedicated

to the me mory of Prof. Fred G. Thompson.

LITERATURE CITED

Burch, J.B. and F.G. Thompson. 1957. Three new Mexican land

snails of the genus Humboldtiana. Occasional Papers of the

Museum of Zoology, University of Michigan 590: 1-11: pls.

]—4.

Mejia, O. and G. Zuniga. 2007. Phylogeny of the three brown

banded land snail genus Humboldtiana (Pulmonata:

Humboldtianidae). Molecular Phylogenetics and Evolution

45: 587-595.

Pilsbry, H.A. 1927. The structure and affinities of Humboldtiana

and related helicid genera of Mexico and Texas. Pro-

ceedings of the Academy of Natural Sciences of Phila-

delphia 79: 165-192.

Pilsbry, H.A. 1948. Inland Mollusca of Northern Mexico. I. The

genera Humboldtiana, Sonorella, Oreohelix and Ashmu-

nella. Proceedings of the Academy of Natural Sciences of

Philadelphia 100: 185-203: pls. 12-14.

Thompson, F.G. 2006. A new land snail of the genus

Humboldtiana (Gastropoda: Pulmonata: Humboldtia-

nidae) from Nuevo Leén, Mexico. The Nautilus 120:

21-24.

Thompson, F.G. 2006. Some landsnails of the genus Hum-

boldtiana from Chihuahua and Wester Texas. Bulletin of

the Florida Museum of Natural History 46: 61-98.

Thompson, F.G. 2011. An annotated che cklist and bibliography

of the land and freshwater snails of Mexico and Central

America. Bulletin of the Florida Museum of Natural

History 50: 1-299.

Thompson, F.G. and G.P. Brewer. 2000. Landsnails of

the genus Humboldtiana from Northern Mexico

(Gastropoda, Pulmonata, Helicoidea, Humboldtiani-

dae). Bulletin of the Florida Museum of Natural History

43: 49-77.

Thompson, F.G. and O. Mejia. 2006. Two new land snails of the

genus Humboldtiana (Gastropoda: Pulmonata: Humbold-

tianidae) from Chihuahua, Mexico. The Nautilus 120:

95-99.

Von Ihering, H. 1892. Morphologie und Systematik des Geni-

talaparates von Helix. Zeitschrift fiir Wissenschaftliche

Zoologie 54: 425-520.

NAUTILUS

Volume 132

2018

AUTHOR INDEX

ANB BAU E at LD) Seetewe tee een OR ee ont oe aa tet ane ee Tee ae eran Ss neta 101 TER NAIN Eg ge retrer coments a Seth ee te SM Sars Src dav cesses aeteaavietaeeti- 51

/NIBGINUING), TRS -Secosenectbescnonedabssobucc coe ncceoDae eee Bee Cece ec eoer ne ees ROSE ane 19 EAGTETSSS) LITINUA, IMs “aoscecnconsanssesce soacde0 bodes SacBee A onecccl AE ooC AOA AEERACOOMESSRE 45

JAIVNINTOWR ISS oeconsnonceoroacedo testcase Bese nec enc aacane Recon c ee cece eee eeeenee 117 |EGISISZA® 18}.. coneaqposaanaescoosbocaceedoaadsoseLcesacen cits ape sess oSionneEe Bacco EOE 124

IBYOLGAINIE: Ns} eacdiadseodse dap tooncecaactoeticSentecibaSoeed ee ouceadaactodce nee eceaetnaees 45 NM Ne BUA eK O) otter cect onckdeciareceonee te acan eetnccr ocece at arese cio esse neers 124

IENORRIUATOWA, Ins -csesepsadticoadasockeesaae sondasshidececbodesaconsaouctenecuceeeese aarere 45 COOLING ACID Chat coed eaanccac cece eo poact ead tame sen OsaetE once Das REB CEE Pee 19

JBSIOTRINE, Ro, Seneasccsdanacaeeapncseonccesict ie aaaadicaodeaaneanoced Raaarasaaa eee meaner 33 IRUESWESS= (© @ NATE: FU easement ete ee 124

(GANSINAINO) (GANG [1S )o:. dchasaasstacatece eid oSecBee caEEe Bo ond eOCOS SOC CaS aaa REE 65 SAILILIBG. JNA CAN. Gk op sbansopaaescescos Boasecasscocuce ease Gack Coc eso Sa SeSCEECEEE 19

(CAIMIPANGIIATRT, 18%, accsondanodancsnsbocsecsadosuenactionnenc onuabdo nado OnE aT see EEE HEE §3 SUIMIOINTS: |] Lal RGILS cscoosdboaancteoccucebaecsosoaoer ase cBaenon ace ene ase onaeaaaceeee 101

CAWATITEATR Teg IB) Gara or essessacens Seas cons Ste seontveemeenrsdcsesscoyuesteaceeree 101] STARGINS Reh esses seortnss vis stare chasasareneemenes tet Soertantierovtotnre sain: |

IDYOTUIGILAGS Sis. sosodenssandtoasesc5szeces seed neaeaee ss86oe ASSOUSSE EE SEES Pe anEcERRee 30 SONUZZIN, SAIS ia cacacectauc a ecniG i omaeene a shae ber oo eee O ne OOD Oc ERO 65

FAR © 1 ya Ege octet Bale wns cena nae ede ea ss the bare oa SoS 45 S OUMRES RS tiga serene tee ere eee te aban eRe rare er ol, 91

(ETOGI AR, ID) Uns Sacccocsacese eS Ese Tce e ce Oe SSN SOR ees E CoE ee 83 AMEN ORI OMIM Se ccccsescvsces esters ses sce seat sees senses sor nesevsses os sess seaesoes: 71

| SUATRACTERIAKOIRL, IMIG... conaonesseconscnosonoooncascoobeondesossonoebecooasebed SD, Uf UTE WAINNGY | F cieetcecnarstcgne voces tevesecyaecdudarsé cin? savodeeesn scadeaseecocieaes 30

USS ANDIDIAHICZZ ISG" acooceensoseacrosoobacecnadoenbedonask onc sonatanse opeconOs Oe Sea 117 AVES iad De syeabe stich So, Santen oder ci osha ki aaeeateneencmneaesieeahLcdeessieeeet: 08, 113

IJIN SORE (G8 Se oe erence oot coe Sete so loco, ucts Sadacecuacle st 117 LITAIN Ga | [eager ce eects Ase deetea Rare ese ous tages so. shore sen acest eben: 58

IKOATRYATITAWIE AIA pee tetra teeter ee rca ene Serre Rae cee er eet ccas Pastore teeta: 45 TABUNNNGE SNE DN aceettoccrces ecco ee eee eee Rc eee ae 13, 113

ILHANUVILIIG: (Gs Reds badconesosconespessonsue aac ROO SSNSODE coS NCEE EOS Deore 30 CSEVIAIN (Ca SU incest eee Oe oes DS ee Rite ila re a 13, 58

NEW TAXA PROPOSED IN VOLUME 132

GASTROPODA

Boreocomitas inouei Amano, Hryniewicz, and Jenkins, 2018 (Pseudomelatomidae, fossil) .............00cccccceeeceeees settee eee eeeeeeerereeeeeeeees 119

G@alliostomassnesaaAnancmWeimandeAhnanes2 OlSosmewaspeciess Calliostomatidac) pes eee 62

Calliotiopissuapensise7nanveandeAnanees? 0ommewaspeciess (Calliotropid ac) pmseeeeeee eee eee 16

Humboldianaaurea Mejiarandeltonezs 2018 tmnewsspecies | (Elumboldtianid ac) esses sees esos eee ee ee 126

Humboldtiana paquimei Mejia and Lépez, 2018, new species (Humboldtianidae) ................c.cccececees ec eeeeeeseeeeeeeeeseseseseseeeesesenesenees 128

Humboldtiana wixarika Mejia, Lopez, and Reyes-Gémez, 2018, new species (Humboldtianidae)..........0..0.0.0:.0c cee eee 125

Memecilok-hanesupinzacw/nan teand Vel 0ommevvas pe cies (Gancel lari dae) pees eee nena tenn ee 114

Murdochella tricingulata Campagnari and Geiger, 2018, new species (Epitoniidae) ...........0..ccs cece eseeee ees ests esses esse eseseeeeesereseeeey 85

Reonidla Umanorum Sowines, AUS, messy GoSHES (ACBOMCS, HOSS) scccccosnce3e90005¢560580006003050000857505 65595345 0055650000 509093555050 OOCIGICEE 54

BIVALVIA

Cyclonaias necki Burlakova, Karatayev, Lopes-Lima, and Bogan, 2018 (new species) (Unionidae) ........0...0.00c0:cce cece eects 46

Beu, Alan

Bogan, Arthur E.

Carew, James

Cowie, Robert H.

Del Rio, Claudia Julia

deMaintenon, Marta

Eernisse, Douglas

Fedosov, Alexander

Ford, David

Garner, Jeff

Groves, Leslie

REVIEWERS FOR VOLUME I:

Harasewych, M.G.

Hayes, Kenneth A.

Hunter, Cynthia

Kabat, Alan

Kantor, Yuri

Kiel, Steffen

Lee, Taehwan

Nielsen, Sven

Padilla, Dianna

Pastorino, Guido

Pearce, Timothy A.