THE NAUTILUS

QL

HOI

Volume 131, Number 3

September 29, 201 7

ISSN 0028-1344

A quarterly devoted

to malacology.

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|ose H. Leal

The Bailey- Matthews National

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National Museum of

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Smithsonian Institution

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Department of Invertebrates

Field Museum of

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Arthur E. Bogan

North Carolina State Museum ol

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Philippe Bouehet

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Center for Conservation Research

and Training

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Steffen Kiel

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Swedish Museum of Natural History

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Geerat j. Vermeij

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THE0NAUTILUS

Volume 131, Number 3

September 29, 2017

ISSN 0028-1344

CONTENTS

Carole S. Hickman

G. Thomas Watters

Paul Larson

Nemocataegis, a new genus and two new species of relictual seguenzioid

gastropods (Vetigastropoda: Cataegidae) and a geobiologieal framework for

integrating patterns in Deep Marine Wallacea . 151

A revision of the Hispaniolan genera Chondropomella , Chondropomium ,

and Clydonopoma (Gastropoda: Annulariidae), with the recognition of

a new genus, Superbiponur. phylogenetic, radular, and

conchological evidence . 163

Note

Arthur E. Bogan Designation of a lectotvpe for Paroaspina collina (Conrad, 1836)

(Bivalvia: Unionidae) . 202

Notice

205

THE NAUTILUS 131(3):151-162, 2017

Page 151

Nemocataegis , a new genus and two new species of relietnal

seguenzioid gastropods (Vetigastropoda: Cataegidae) and

a geobiological framework for integrating patterns in Deep

M arine Wallacea

Carole S. Hickman

Department ol Integrative Biology and Museum of Paleontology,

University of California,

Berkeley, CA 94720-3140 USA

[email protected]

ABSTRACT

Nemocataegis new genus, type species N. nmeani, is proposed for

two elaborately sculptured, small-shelled cataegid gastropods from

separate deep-water basins in the Indonesian biogeographic realm

of Wallacea. The Ope species is from 885 m in the Gulf of Bone,

between die southern and southeasteam anus of Sulawesi where

the narrow gulf opens into die Flores Sea. The second species,

N. quinni, is from 503 m in the Molucca Sea adjacent to 1 lalmaliera.

Both species have the oblique, broadly expanding aperture, fluted

outer lip and strong spiral cords characteristic oi Cat aegis McLean

and Quinn, 1987, but shells are smaller ( <20 nun) and the spiral

cords are densely nodose, with fine axial ribs connecting the nodes

and a unique microsculpture of fine threads and intritaealx in the

channels between node rows. A previously described true Cataegls

is endemic to the Makassar Strait in Wallacea, occurring at depths in

excess of 1000 m. Cataegidae is re-diagnosed to include two genera,

six nominal living species and two nominal fossil species. The family

is assigned to Seguenzioidea based on a combination of morpho¬

logical and molecular data. The new genus is endemic to Deep

Marine Wallacea, and the new species occur in geographically

isolated basins. Along with previously described deep-water gas¬

tropods in geologically ancient clades, they contribute to evidence

of relictual biotas. The biogeographic patterns are correlated with

factors that include the tracks of collecting expeditions, submarine

topography, biogeographic lines, patterns of oceanic circulation,

distribution of volcanic arcs and subduction zones, and major

structural features resulting from tectonic collision, fragmentation,

suturing, deformation, and extension over a period of more than

50 million years.

Additional ’Keywords: Sulawesi, Hahnahera, Molucca Sea,

Makassar Strait, Banda Arc, biogeography, hydrocarbon seep,

deep sea, endemism, biogeography, systematics.

INTRODUCTION

This is the third in a series of papers describing unusual and

relatively large-shelled (>20 mm) deep-water (200-2000 m)

gastropods from three basal vetigastropod families.

Remarkably, the specimens are from expeditions and

museum collections in which they may have been so

unusual that they were incorrectly filed, misidentified, or

simply waiting for resolution of the confused state of the

taxonomy of fossil and living basal gastropods that have

been variously arranged under many names at the

“archaeogastropod” grade of evolution.

Tin1 first paper in the series (Hickman, 2012) described

a new genus and two new species in the family Gazidae

Hickman and McLean, 1990. It included anatomical and

radular features as well as new' shell characters. The

second paper (Hickman, 2016) described eight species in

the family Calliotropidae Hickman and McLean, 1990.

Both papers are focused on the Indo-West Pacific Region

and explore the complex geologic history of tectonic plate

interactions, arc volcanism, subduction, accretion, and

hydrography that identify a marine geobiogeographic

realm designated here as “Deep Wallacea."

Deep Wallacea is distinct from the original terrestrial

biogeographic concept that originated with Alfred Russel

Wallace when he drew his famous line. Ironically, the

region between Wallace’s Line and Weber’s Line was

named “Wallacea” bv geologist and paleontologist Roy

Ernest Dickerson in an overlooked paper (1924: 3, fig. 1)

designating “(1) the Asian (Sunda) Shelf, (2) Wallacea, and

(3) the Australian Sahul Shelf (as) the three well-defined

divisions of the Australasian Mediterranean Sea."

The primary objective of this paper is taxonomic: to de¬

scribe a new genus and two new species of extant deep-water

cataegid gastropods and to revise the diagnosis of the family.

A secondary objective is to provide a set of maps and dis¬

cussion of factors correlated with the disjunct and relictual

distribution patterns of basal marine gastropods in Deep

Wallacea. These include the tracks of collecting expeditions,

the path of the Indonesian Throughflow and patterns of

oceanic circulation, the division of the region into many

separate basins and sub-basins, the distribution of active

volcanoes and volcanic arcs associated with crustal collision

Page 152

THE NAUTILUS, Vol. 131, No. 3

and subduetion, and the distribution of major structural

features and terranes tliat record a long geologic history of

dramatic change in the distribution of land and sea.

A brief background of the ecology of living cataegids,

their paleontological record, and their previous classifica¬

tion follows as an introduction to the systematic treatment.

Ecological Background

Living cataegid gastropods have been reported previously

from bathval depths (>200 m) in two widely separated

regions of the world: (1) the combined basins of the

Caribbean Sea (McLean and Quinn; 1987; Waren and

Bouehet, 1993, 2001; Gracia et ah, 2001) and Gulf of

Mexico (McLean and Quinn; 1987; Waren and Bouehet,

1993, 2001; Carney, 1994; Cordes et ah, 2010) in the

Americas and (2) basins in the Southwestern Pacific that

include the South China Sea (Fu and Sun, 2006), Wallacean

Indonesia (McLean and Quinn, 1987), and Vanuatu

(Kano, 2007; Waren, 2001). Complex active teetonism in

these regions has produced some of the same geologic

settings and features (subduetion zones, thick accretionary

prisms and melange, fault zones, mud volcanoes) that in

turn give rise to unusual ecological settings. While geolo¬

gists have focused on fluid expulsion in these settings

(sulfides, petroleum, and other hydrocarbons, especially

methane), biologists have focused on the ecological com¬

munities exploiting chemical energy via chemosynthesis.

Living cataegid gastropods eonsistentlv are associated

with cold seeps where conduits for sulfide- and methane-

rich fluids are expelled. In these environments microbial

chemosynthesis creates both a rich nutritional source and

a challenging oxygen-depleted setting for development of an

extremophile macrobiota. In the Gulf of Mexico cataegids are

linked to specific sulfide and hydrocarbon seeps off Louisiana

(Carney, 2004; Cordes et ah, 2010) and in the Caribbean in

the thick accretionary prism off Venezuela, Trinidad, Barbados

and Colombia (Gracia et ;il., 2001: Gill et ah, 2005).

Fluid expulsion may also include barium-rich water that

precipitates at oxic/anoxic boundaries as barium sulfate

and barite sheets, mounds, cones and chimneys where

a cataegid is part of a unique seep community (Cordes

et ah, 2010). The cataegid collected in the volcanic arc of

Vanuatu was associated not only with cold seeps and

hydrothermal vents, but also was reported as common on

sunken wood (Waren, 201 1).

Paleontological Background

Fossil specimens all occur in cold-seep carbonates and

represent a broader geographic range that includes Lower

Cretaceous (Valanginian) rocks in California (Kaim et ah,

2014), and Upper Cretaceous (Campanian) rocks in Japan

(Kaim et ah, 2009; Kiel, 2010). Kiel and Campbell (2005)

note that “trochomorph” gastropods that are abundant in

some Lower Cretaceous cold-seep limestones in California

are difficult to classify without original shell material.

At the boundary between the South American and

Caribbean plates, collection records of Cataegis rneroghjpta

are more numerous from Paleogene and Neogene seep

carbonates than from modem seeps in the region (Gill et ah,

2005), although this is potentially biased by greater sam¬

pling effort in the fossil record. The record of Cenozoic

seep faunas in the Caribbean has been extended to sites

in the Dominican Republic and Cuba (Kiel and Hansen,

2015).

Tl le number of species is likely to increase with rec¬

ognition of fossil shells described under other names in

early literature. Kaim et al. (2014) note the morphological

similarity of a species described as Phasianema tauro-

crassum Sacco, 1895 from the Miocene “Calcare a Lu-

cina ," classic seep limestone ehemoherms (Moroni, 1966;

Taviani, 1994) in the Appenine chain of the Italian pen¬

insula. Of special note is a species from Trinidad described

as Solariella godineauensis bv Katherine Van Winkle (1919),

who noted (p. 26) both a flaring aperture and crenulated

carinae. Kiel and Hansen (2015, fig. 14F) provide an ex¬

cellent image that clearly shows seguenzioid axial threads in

the interspaces between spiral cords.

Nomenclature and Classification of Cataegid

Gastropods

The genus Cataegis , along with the family Cataegidae, is

a relatively late addition to the large global inventory of

vetigastropods. The family group was originally described

and treated as a trochid subfamily of uncertain affinity

(McLean and Quinn, 1987; Hickman and McLean, 1990;

Waren and Bouehet, 1993). In a transformative, new working

gastropod classification (Bouehet and Roeroi, 2005), the

family was tentatively reassigned to Seguenzioidea along

with Calliotropidae and Chilodontidae. This new view of

Catageidae is supported by molecular data (Kano, 2007;

Kano et ah, 2009; Aktipis and Giribet, 2012), although

the cataegid sequence used in all three analyses ( Cataegis

sp.) is, unfortunately, not tied to a shell voucher specimen.

Prior to this report there were four nominal species of

cataegid gastropods, all described under Cataegis : two from

the Caribbean (McLean and Quinn, 1987), one from

Indonesia (McLean and Quinn, 1987) and one from the

South China Sea (Fu and Sun, 2006).

Paleontologists have recognized fossil Cataegis specimens

as seguenzioid without further classification (e.g. Kiel and

Hansen, 2015), or with assignment to Cataeginae as a sub-

familv 'of Chilodontidae Wenz, 1938 (Kaim etah, 2009; 2014).

SYSTFMATICS

Subclass Vetigastropoda Salvini-Plawen, 19S0

Seguenzioidea Verrill, 1884 (unranked)

Fucycloidea Koken, 1897 (unranked)

Family Cataegidae McLean and Quinn, 1987

Genus Cataegis McLean and Quinn, 1987

Type Species: Hoinalopomafinkli Petueh, 1987 (senior

synonym of Cataegis toreuta McLean and Quinn, 1987).

C.S. Hickman, 2017

Page 153

Recent, Gulf of Mexico, Caribbean and Western Atlantic;

at continental slope depths, 337-1283 m.

Remarks: McLean and Quinn (1987) based the family

group and genus primarily on a highly unusual radula in

the t\pe species. The raehidian was missing, the in¬

nermost laterals lacked cusps and were elaborately fused,

and remaining three laterals had small cusps but com¬

plexly interlocking shafts and bases. Hickman and

McLean (1990) suggested that the radula represented

“an evolutionary excursion in the direction of disorder." It

was therefore surprising when Waren and Bouehet (1993:

fig 14C) figured a radula from C. ineroglypta McLean and

Quinn, 1987 with a robust raehidian tooth with a well-

developed and sharply-edged cusp. Habitat data help

make sense of the radular disparity, because C.

ineroglypta occurs on hard substrates at methane seeps

and feeds on bacterial films, whereas the type species is

reported to feed on sunken and decaying seagrass (Waren

and Bouehet, 1993). Although radular morphology is

usually the best indicator of suprageneric placement,

the cataegid shell is unusually rich in characters that

are used here to supplement the original description of

the family group and to accommodate a second genus,

Nemocataegis , for which the radula and anatomy are

unknown.

Three unique characters, in combination, were speci¬

fied as diagnostic of the family group and genus (McLean

and Quinn, 1987: 111-112): strong spiral cords, lack of

columellar plications, and the oblique, broadly expanding

aperture. Additional characteristic features include the

deep, concave spiral interspaces lined with extremely fine

prosocline or slightly sinuate axial threads and a dense

microsculpture of discontinuous, reticulate or anastomos¬

ing, ridges and grooves that are infilled or oveiprinted by

a chalky or crumbly brownish intritacalx (sensu D’Attilio

and Radwin, 1971). The intritacalx may be so extensively

developed as to obscure the microsculpture, but it clearly is

not a continuous periostracal sheet. Intritacalx is also an

important microstructural feature in two other seguenzioid

families: Eucyclidae (Herbert, 2012) and Calliotropidae

(see Hickman, 2016, for a discussion).

Macroscopic axial sculpture in Cataegis is restricted to

elongate, fine nodules on the spiral cords. In all cataegids

the columella is thickened, arcuate, lacking plications or

teeth and covered with a thin callus that extends to cover

the umbilicus completely or to leave a narrow umbilical

chink. Interior nacre is not covered with a translucent

inclined prismatic layer as in some seguenzioids, and it

does not extend all the way to the growing margin of the

shell, even in adults with descending sutures and final

apertures. Spiral ornament is visible through the interior

nacre as ridges and grooves. Fluting of the outer lip at the

termination of spiral ridges and grooves is also charac¬

teristic of the family.

Genus Nemocataegis new genus

Type Species: Nemocataegis mcleani new species.

Description: Shell small for family (height <15 mm),

turbiniform, low spired, with rapidly expanding bodv

whorl and large, oblique aperture; spiral ribs increasing by

intercalation to >10 on body whorl and crossed by nu¬

merous thin, continuous, prosocline axial threads that

form elongate nodes or sluup scales on spiral ribs; spiral

groves with microscopic sculpture and brownish intri¬

tacalx; columellar lip arcuate, smooth, and thickened by

opaque callus; callus almost completely covering umbi¬

licus as well as extending into interior margins of basal and

apertural lips and covering nacre; adult suture and ap¬

erture descending; interior nacre with weak ridges and

grooves reflecting exterior spiral sculpture.

Remarks: The new genus is clearly distinguished from

Cataegis by its smaller shell size ( < 1 5 mm), more nu¬

merous, narrower, and prominently-beaded spiral cords,

narrower spiral interspaces, lack of continuous periostracum,

and well -developed intritacalx in microscopic grooves

between crowded axial microsculptural threads.

It is unfortunate that the protoconchs are worn and that

there are no data on the anatomy or radula. It is re¬

markable that the empty shells are so exquisitely pre¬

served, and it is possible that the well-developed

intritacalx has served a protective function against dis¬

solution. Shells show no sign of encrustation, but there is

evidence of repaired breakages. The known specimens

come from gray mud at slope depths and are most likely

deposit feeders.

Etymology: Nemo (Cr. thread) + Cataegis, in refer¬

ence to the fine axial sculpture that distinguishes the

threaded cataegids.

Nemocataegis mcleani new species

(Figures 1-5)

Diagnosis: Enlarged bodv whorl evenly rounded,

without peripheral or basal demarcation; 6 primary spiral

cords visible in apical view, 5 additional spiral cords visible

in umbilical view; secondary spiral cords intercalated

between primary adapical spiral cords; axial ribs forming

short, sharp, anteriorly directed spines on primary and

secondary spiral ribs; thin callus deposit covering interior

nacre on outer and basal lips overlapping columellar callus

at base and top.

Description: The type species of Nemocataegis is

clearly distinguished from N. quinni new species (de¬

scribed below) by more numerous primary spiral cords

and more prominent, sharp, spinose projections where

they are crossed by axial threads. The short spines are

incompletely closed and project anteriorly (Figures 2, 3).

In the deep spiral grooves the depressions between axial

threads are filled with intritacalx that either obscures

underlying structure or shows faint axial lineation

(Figure 5).

Holotype: USN M 239464, height 14.4 mm, maximum

width 14.0 mm.

Page 154

THE NAUTILUS, Vol. 131, No. 3

Figures 1-5. Nevwcataegis mcleani new species 1 . Semiapertural, 2. Apical, 3. Basal, and 4. Abapertural views ol holotype, USNM

239464, height = 14.4 mm. 5. Detail from Figure 4 of nodose axial ribs and closely-spaced threads filled with brownish intritacalx.

Txpe Locality: 03°17'40" S, 120°36'45" E, Gulf of

Boni (Bone), Celebese (Sulawesi), SE of Olang Point, 484

fathoms ( = 885 m), U.S. Fish Commission, RA’

Albatross, Station 5656, 19 December 1909, gray mud.

Distribution: Known only from the type locality.

Etymology: Named for the late James 11. McLean in

recognition of his many important contributions to un¬

derstanding basal marine gastropods.

Remarks: The holotype appears to be an adult speci¬

men on the basis of terminal growth features that include

a descending suture and terminal translucent callus

covering, but not totally obscuring, nacre inside the outer

lip (Figure 3). Height and width are effectively equal. A

major breakage in the basal portion of the outer lip

(Figure 4) is repaired by a partially disjunct continuation

of sculpture. The shell is remarkably fresh in appearance

and was not encrusted by epizoans in the manner common

to many calliotropids, especially those living on hard

substrates.

Nemocataegis quinni new species

(Figures 6-10)

Description: Enlarged body whorl with slightly con¬

cave shoulder slope; final aperture strongly oblique, with

descending suture; spiral ribs of unequal strength and

spacing, interspaces broader than ribs; numerous fine axial

threads producing small blunt nodes at intersection with

spiral ribs; brownish intritacalx well developed between

axial threads in spiral interspaces; columellar callus re¬

flected and covering umbilical region; columellar lip

strongly arcuate, callus extending into aperture, partially

obscuring interior nacre; interior nacre with weak spiral

ridges.

Holotype: USNM 239279. Height 14.0 nun, maximum

width 13.4 mm.

Type Locality: 00°19'20" N, 127°28'30" E, Molucca

Sea, Halmahera, off Makian Island, 275 fathoms (=503 m).

U.S. Fish Commission, RA7 Albatross, Station 5622,

29 November 1909, gray mud.

Distribution: Known only from the type locality.

C.S. Hickman, 2017

Page 155

Figures 6-10. Nemocataegis quinni new species. 6. Semiapertural, 7. Apical, 8. Basal, and 9. Abapertural views of holotvpe, USNM

239279, height = 14.0 mm. 10. Detail from Figure 4 of nodose axial ribs and closely-spaced threads filled with brownish intritacalx.

Etymology: Named for James F. Quinn, Jr. in recog¬

nition of his insightful contributions to the systematics of

deep-sea gastropods.

Remarks: This species is distinguished from the type

species by its more numerous, shorter and bluntly

rounded nodes on tin1 spiral ribs, a completely obscured

umbilicus, slightly convex shoulder slope on the body

whorl, more strongly descending final aperture, and less

visible traces of spiral sculpture in the interior nacre. The

terminal growth features on the shell of the holotvpe

suggest that it is an adult. The brownish intritacalx

(Figure 10) is well developed in axial interspaces over the

entire shell except at the apex. The protoconch small, but

sufficiently worn that the boundary with the worn early

teleoconeh whorls cannot be discerned.

DISCUSSION

In light of pronounced under-sampling of the deep-water

fauna of Wallacean Indonesia, it is remarkable that the

region contains so many species and higher taxa that have

never been recorded in relatively well-sampled regions of

the Indo-Paeifie. The endemic genus and two new species

of Cataegidae described above join a diverse assemblage

of basal gastropods that includes an endemic genus of

Gazidae (Hickman, 2012) and six recently-described

large-shelled species of Calliotropidae (Hickman, 2016).

Adequate sampling of the deep Wallacean fauna re¬

quires a protocol that recognizes factors that appear to be

contributing the accumulation of relictual taxa. The pri¬

mary purpose of this discussion is to extend previous

characterization of deep Wallaeea (Hickman, 2009a;

2009b; 2009c; 2012; 2016) and to provide a set of graphic

illustrations (Figures 11-16) of the interacting compo¬

nents of deep Wallacean complexity.

This narrative begins with the currently known oc¬

currences of species with deep (Paleozoic or Mesozoic)

evolutionary origins and endemic or strongly disjunct

representation in separate basins within Wallaeea. It

proceeds with components of explanation that are con¬

sistent with the objectives and tracks of previous collecting

expeditions, the basins and sub-basins of Deep Wallaeea,

shallow oceanic circulation and path of the Indonesia

Throughfiow, the distribution of active volcanoes and

THE NAUTILUS, Vol. 131, No. 3

Page 156

volcanic arcs, and the distribution of trenches, faults, and

major tectonic features that are essential to understanding

the geologic history of changing distribution of land

and sea.

The term “Indonesian Archipelago” (the Malay Archi¬

pelago of Wallace, I860; 1863; 1869) is a misnomer. A

modern map Indonesia at the scale typically used to depict

the islands of Oceania misses most of the more than 1 7000

islands and their arrangement. The region is actually

a composite of separate archipelagic seas, straits and gulfs

that have opened, expanded, contracted, and in some

instances disappeared over a 200 My geologic history that

began with the break-up of Gondwana (Metcalfe, 201 1).

The larger islands in these seas have likewise moved,

collided, fused, separated or disappeared during the

tectonic evolution of the region. Many islands have

emerged or submerged during its more recent eustatie

history. The biogeographic effects of sea level change

during the Pliocene and Pleistocene have received con¬

siderable attention from terrestrial and shallow marine

biogeographers. Regional biogeographv still lacks a deep

marine perspective. Appreciation of the paleobathymetrie

complexity of the Wallacean seaways is, however, of in¬

creasing interest to physical oceanographers and meteo¬

rologists investigating large-scale influences of ocean

circulation on global climate (e.g.. Cane and Molnar,

201 l).

Endemic and Relictual Taxa in Wallacea

Occurrences of cataegid gastropods in deep Wallacea are

illustrated in red in Figure 1 1, along with occurrences of

calliotropid (green) and gazid (blue) vetigastropods,

abyssochrvsid gastropods (yellow), and a living coelacanth

fish (star). Previous discussion (Hickman, 2016) of the

deep geologic origins (Paleozoic or Mesozoic) and pro¬

nounced global disjunctions in these groups are consistent

with the hypothesis that they are relictual taxa. The fossil

records of more widespread geographic occurrence and

greater taxonomic diversity are similarly indicative of

a concentration of relictual taxa in Deep Wallacea.

Expeditions and Benthic Sampling Coverage

Within Wallacea, the distributions shown in Figure 1 1 are

correlated with the tracks of the Dutch Siboga Expedition

of 1889-1900 and the R/V Albatross Philippines Expe¬

dition of 1907-1910 (Figure 12). Although both expedi¬

tions produced remarkable oceanographic data, dredge

samples, and new marine taxa from >200 meters, the

research objectives were different in intent and execution

as well as in their subsequent study and publication of

results.

The Siboga Expedition was conducted under the

leadership of Professor Max Weber, a distinguished zo¬

ologist at the University of Amsterdam, whose proposed

track (shown in orange in Figure 12) was a zig-zag ex¬

ploration of the margins of the region of deep marine

basins between the Sunda and Salmi continental shelves.

His proposed track was not followed precisely, but it

appears to reflect Weber’s curiosity about the terrestrial

biogeographic lines that recently had been drawn by

Wallace (1863) and Lyddeker (1896) and their potential

oceanographic correlates. The commander of the Siboga,

G.F. Tydeman, was expert at recording deep soundings

and oceanographic data (Tydeman, 1903) from the >300

sampling stations. Most remarkably, the taxonomic results

were published in a series of >100 monographs and in¬

cluded descriptions of the prosobranch gastropods

(Sehepman 1908; 1909). For a summary of the Siboga and

other Dutch Oceanographic research in Indonesia in

colonial times, see Van Aken (2005). The specimens were

from both shallow and deep stations on the periphery of

the region, and the Siboga did not sample in the gulfs,

basins, and sub-basins of Sulawesi and Halmahera.

Thi‘ Albatross Philippines Expedition was designed as

a focused survey of the aquatic resources of the Philippine

Islands, under the command of Hugh McCormick Smith

of the U.S. Bureau of Fisheries. It was conducted as

a series of cruises, with the addition of a final two months

in the Dutch East Indies (Indonesia) at the end. The

major biological reports are on the fishes (see Smith and

Williams, 1999). The track of the Albatross in Indonesia

is shown in purple in Figure 12).

Little is known about the objectives or why the cruise

was narrowly focused on obtaining collections and data

from deep stations in Sulawesi (Makassar Strait, Gulf of

Bone, and Gulf of Tomini) and the western margin of

Halmahera. Malaeologist Paul Bartsch, who had repre¬

sented the Smithsonian Institution in the preceding

Philippines cruises, left the expedition before it departed

for Indonesia. Hugh McCormick Smith also had de¬

parted, and there is no record of who was in charge. What

little we do know is from the autobiography of Roy

Chapman Andrews, who was recruited as a 24-year-old

graduate student and charged with terrestrial collecting.

He notes (Andrews, 1943: 67) that “It wasn't a ‘happy

ship and that he was “supposed to have no part in the

dredging operations” (p. 72). He makes no mention of

marine mollusks in his romanticized account.

In spite of what little is known samples were obtained

from 72 stations between 7 November and 30 December

1909. Specimens and station data were meticulously

processed and accessioned into the Smithsonian mollusk

collection and archives and are still being discovered and

studied by malaeologists with expertise in various taxo¬

nomic groups.

With the exception of the Siboga Expedition and its

monumental series of reports, oceanographic research

during colonial rule in Indonesia has been character¬

ized as a “gunboat science” (Van Aken, 2005). However,

Van Aken’s (2005) summary highlights many important

hydrographic, physical and chemical oceanographic, and

geophysical data that were obtained and published by

dedicated scientists who obtained private funding.

A limited biological sampling effort in 1922 in the vi¬

cinity of Kai and Tanimbar was aimed at evaluating

a proposed Danish tropical marine station (Mortensen,

C.S. Hickman, 2017

Page

10/

Figures 11-12. Simplified maps of marine Wallaces comparing occurrences of relictual deep-water taxa and tracks of major collecting

expeditions. 1 1 . Collecting sites for eataegid (red circles), calliotropid (green circles), gazid (bine circle) and abyssochrisid (yellow circles)

gastropods; and living fossil coelacanth fish (blue star). 12. Tracks of the Sihoga Expedition (orange) on the margins of Wallacea and the

Albatross Expedition (purple) within northwestern Wallacea.

1923) and contributed no deep-water gastropods perti¬

nent to this discussion. The KARUBAR joint Indonesian

and French cruise in this region, also sampled primarily at

shallower depths (< 500 m) with a strong focus on the

Arafura Sea (outside of Wallacea) rather than in the Banda

Sea (Crosnier et ah, 1997). An additional joint Indonesian

and French sampling effort (CORIN' DON 2) (Moosa,

1984) sampled deep-water stations on either sick' of the

Makassar Strait, adding a abyssoehrysid records, including

a new species (Bouehet, 1991).

Bioceographic Lines and The Evolving Concept of

Wallacea

M ore than enough has been written about the drawing of

lines by terrestrial biogeographers in the Indonesian re¬

gion (e.g., Mayr, 1944; Michaux, 2010). From a marine

perspective, the margin of the Sunda Shelf to the west and

margin of the Sahul shelf to the east define Wallacea as

a region in precisely the manner originally intended bv

Dickerson (1924) when be proposed the term. Wallacea is

illustrated here (Figure 13) as lying between the “Wallace

Line and Weber Line slightly modified" (see Dickerson,

1924; fig. 1).

The' Oceanographic Data Set for the Albatross sampling

in Indonesia includes 71 stations concentrated close to shore

in four separate basins adjacent to Sulawesi and Halmahera

(Makassar Strait, Gulf of Bone, Gulf ’of Tomini, and Molucca

sea). These basins are shown in blue (Figure 13). In each

setting, the 2(X) m isobath is veiy close to shore, and 56 of the

stations sampled were from depths >200 m, 30 stations were

from >1,000 m and 13 stations were from >1,500 m.

Restriction of marine fauna to individual basins, gulfs,

and straits within Wallacea is consonant with the complex

geological history of fragmentation and amalgamation in

marine Wallacea as a whole. Similar recognition of

multiple sub-regions of endemism in the terrestrial biota

(e.g., Michaux, 2010) advances biogeographic analysis as

an integration of distributional data with geoteetonie

history.

The Indonesian Throughflow and Shallow Oceanic

Circulation

Patterns of distribution of marine invertebrates in Wallacea

are typically examined in terms of the oceanic currents that

transport and disperse pelagic larvae. Considerable emphasis

has been placed on the potential importance of the In¬

donesian Throughflow (ITF) in determining patterns of

genetic connectivity. The gateway connection between the

North Pacific and the Indian Ocean (Figure 14) is a com¬

plex system of shallow and deep currents. The main route

is a shallow southward flow of the Mindanao Current

through the Celebese Sea and Makassar Strait. It enters

the Indian Ocean between Bali and Lombok with another

branch turning eastward through the Flores Sea, branching

again to How through the Banda Sea, entering the Indian

Ocean through the Ombai Strait and Timor Passage on

either side of Timor.

Minor secondary flow through the Molucca Sea and

into the Banda Sea is more complicated. North Pacific

water is initially deflected from reaching the Banda Sea by

eastward reversal of flow in the South Equatorial Current

(SEC) into the North Equatorial Counter Current

Page 158

THE NAUTILUS, Vol. 131, No. 3

Figures 13-16. Simplified maps of factors correlated with occurrences of relictual deep-water taxa. 13. Outline ol Wallacea hounded

on the west by Wallace’s Line (purple dashes) and the east by Lydekker’s Line (broken purple dashes), and the four deep basins (blue)

from which the Albatross gastropod specimens were dredged: Molucca Sea, Gulf of Bone, Makassar Strait, and Gulf of Tomini. 14.

Surface currents of the Indonesian Throughflow (ITF) from the North Pacific through the Southeast Asian Gateway to the Indian Ocean

and surf ace currents with no significant How through Wallacea (gray). 1 5. Nine major Wallacean basins underlain by oceanic lithosphere

(blue), the anomalously deep Weber Basin (deep blue), and historically active volcanoes (red triangles) defining basin boundaries. 16.

Fault systems and subduction zones (red) associated with major tectonic terranes, and position of the outer (light pink) and inner (dark-

pink) zones of the Banda Arc.

(NECC) influenced by several large semi-permanent

eddies (Arruda and Nof, 2003). Deeper How entering

the Banda Sea is further impeded by sills and physical

barriers separating sub-basins (Gordon et ah, 2003;

Van Aken et ah, 2009). The Banda Sea appears to con¬

tribute a relatively minor amount to total throughflow

(Gordon and Line, 1996). This is an oversimplification of

a temporally and spatially variable pattern that is in¬

completely characterized in spite of a growing body of

detailed oceanographic data. The patterns of flow vary on

time scales ranging from days and seasons to decades and

significant periods of geologic time (Tillinger, 201 1).

While the ITF (shown in Purple in Figure 14) and

interactions with the boundary between major gyres and

C.S. Hickman, 2017

Page 159

eddies (shown in gray in Figure 14) may be significant

factors in the dispersal and connectivity in the hyper-

diverse shallow marine biota of Wallacea, they are less

likely to play a role in understanding the endemism in

deep-water cataegid, ealliotropid, or gazid gastropods

(Hickman, 2012, 2016). From an oceanographic per¬

spective, this is because deep-water current systems are

different and from a biological perspective because deep¬

water vetigastropods lack a significant pelagic dispersal

stage in their life history. They likewise have been un¬

affected In' Pliocene and Pleistocene sea-level change and

because they occur in basins that are isolated from one

another by submarine topographic barriers that restrict

the dispersal of benthic adults.

Marine Basins and Volcanic Arcs

There are at least 26 hathymetrically distinct basins within

the Wallacean region that reach bathyal depths (>200 m).

The nine illustrated in blue (Figure 15) contain species of

basal marine gastropods in clades associated with

chemical energy sources. The Wallacean basins are al¬

ternatively identified in the literature and on maps as seas

(linil), straits ( selat ), or gulfs (teluk). Straits are narrow

passages between islands, while gulfs are deep inlets or

indentations in the larger emergent landmasses, notably

the Gulf of Tomini and the Gulf of Bone in Sulawesi.

Complex ridge systems create bathymetric isolates in

some of the basins. Examples in the Banda Sea include

a main North Basin (Sula) and a South Basin that is further

subdivided into the Wetar, Damar, and Weber basins.

The Weber Deep (in dark blue) is an anomalous, tec¬

tonically generated bathymetric isolate in the Banda

Forearc with depths exceeding 7 km. The Makassar Strait,

the Bone Basin, and the Gulf of Tomini also contain

hathymetrically distinct sub-basins. Although bathymetry

and submarine topography can confine or restrict benthic

taxa that lack a planktonic dispersal phase, bathymetric

complexity must be further integrated with the geologic

history. This is especially true of volcanic arc activity that

has shaped basin development and the sedimentary

component of deep-water ecosystems.

The link between geologic and ecologic settings in deep

Wallacea has not been explored. Understanding the role

of volcanic arcs in basin development, sedimentation,

underlying stratigraphy, structure, geophysics, and geo¬

chemical peculiarities is crucial to understanding the deep

benthic realm in which endemic and relictual taxa have

accumulated. The major abiotic features of magmatic arc

history (Hall and Smyth, 2008) provide a framework and

starting point for future integrative study.

The modern volcanic arcs in Wallacea (Figure 15) are

depicted by red circles representing volcanoes that have

been active historically (Smithsonian, 2016). There are

many more named volcanic edifices that are presumably

inactive, and little is known of submarine voleanism.

The subduetion zone system in Wallacea includes four

volcanic arcs (Figure 15). The Sunda and Banda volcanic

chains mark the zone of melting as the Indo-Australian

plate is subducted beneath the Eurasian Plate. To the

north, tlu' Sanghe (or North Sulawesi Arc) and the

I lalmahera Arc mark die convergent margins of the former!)

larger Molucca Sea Plate that is disappearing in a unique

modem example of double subduetion system. The con¬

striction of the formerly broad Molucca Seaway necessarily

represents a constriction of deep marine habitat and is

consistent with the occurrence of relictual molluscan taxa

(Hickman, 2012; 2016).

Much of the basin formation in Wallacea appears to be

driven by extensional rather than eompressional tectonic

forces (Charlton, 1991; Charlton et ah, 1991; Hall, 2013;

Pownall et ah, 2013). These include the opening of both of

the two deep interarm basins of Sulawesi: the Gorontolo

basin in the Gulf of Tomini (Pholbud et ah 2012) and the

Bone Basin in Sulawesi (Sudarmono, 2000). The im¬

portance of slab rollback and lithospheric extension in the

Banda Arc is perhaps most dramatic in the proposed

detachment model for formation of the Weber Deep

(Pownall et al. 2016).

Further understanding of basin history requires ex¬

amination of subsurface structure, which is difficult to

depict and interpret in simplified map view. The following

section highlights some of the Wallacean structural fea¬

tures that are especially germane to the peculiar geo¬

graphic distribution of deep-water marine gastropods.

Major structural Features and History

The major structural features of Wallacea (Figure 16)

provide a starting point for understanding the complex

history that is treated in a diffuse and evolving body of

literature and lively debate over alternative interpretations

(See Hall, 2011; 2013; Hall and Blundell, 1996); Hall and

Wilson, 2000; Hall et al. 2011 and references therein).

Full understanding of modern Wallacea requires going

back to the Paleozoic break-up of greater Gondwanaland

and tracing separation and movement of micro¬

continental fragments, the openings new ocean basins,

the collision and suturing of slivers, translations and

subductions, and disappearance of some of the features

ultimately responsible for the modern distribution of

tectonically bounded marine basins and terrestrial

blocks and fragments (e.g., Metcalfe, 2001; 2011).

Features depicted in red in Figure 16 include active

trenches, major sutures, thrust faults, and strike-slip

faults and fault zones.

The most prominent structural feature is the horseshoe¬

shaped Banda Arc (Figure 16), a complex suture zone in

which multiple Jurassic to Neogene pre-eollisional imbri¬

cate wedges are continuous with post-collisional evolution

of the modern forearc (Charlton et al., 1991 ). The zones of

folding and thrusting of the outer arc complex (successively

older from southeast to northwest) and the modem deep

forearc basin are represented in pink, in Figure 16). The

Islands and volcanoes of the active inner arc are repre¬

sented in salmon. The Banda Arc is an exemplar of an arc-

continent collision with multiple episodes of accretion of

crustal blocks and fragments and is treated in an extensive

Page 160

THE NAUTILUS, Vol. 131, No. 3

literature (e.g. Bowin et al., 19S0; Charlton et al, 1991; Hall

and Wilson, 2000; Harris, 2006; and references therein).

Three additional suture zones (Figure 16) are: (1) the

Sulawesi Suture, also a complex region of multiple col¬

lisions, (2) the Sorong Suture, a major east-west trending

fault zone of multiple collisions, and (3) the Molucca

Suture where two colliding volcanic arcs are converging to

close the ancient Molucca Seaway (see Hall and Wilson,

2000 for a review and references).

The relevance of the Wallacean suture zones to marine

biogeography, endemism, and the deep-water relietual

gastropod taxa resides in the clear evidence of geo-

chemically unusual and extreme environments of interest

to petroleum geologists, who have identified commercial

hydrocarbon occurrences (Charlton, 2004), hydrocarbon

seeps (Camplin and Hall, 2014) and mapped fields of mud

volcanoes (Barber et al.. 1986).

Biological expedition reports provide few clues to

geochemical and sedimentary settings at sampling sta¬

tions. However, the increasing geological evidence in

deep Wallacea for both explosive and diffuse expulsion of

fluids rich in sulfides and hydrocarbons (Hickman, 2016)

are consonant with the presence of (1) taxa that have been

identified elsewhere in the world as part of ehemo-

syntheticallv based communities, (2) adaptations to toxic

fluids and hypoxia, and (3) nutrition based on elevated

productivity of chemosynthetic microbes and microbial

mats.

CONCLUSIONS

Understanding of the marine biogeography of Deep

Wallacea is in its infancy. Distribution of relietual basal

gastropod taxa is linked to features mapped in this paper:

expedition tracks and biological sampling, positions of

discrete marine basins and their topographic boundaries,

patterns of ocean currents and circulation, distribution of

active volcanoes and volcanic arcs, and distribution of

what remains of the complex structural record of tecto-

nism over many millions of years. A full geobiological

history will emerge only through integrating data from the

disparate sources and disciplines implicated above. At the

same time, interdisciplinary exchange is likely to assist

both geologists and biologists in addressing separate,

discipline-specific questions. It is ironic that a paleontol¬

ogist should find herself attempting to unravel a deep¬

water geobiological history that has no preserved fossil

record in Wallacea. However, it is those with formal

training in both geology and biology who are most likely to

be challenged and attracted to this kind of integrative

effort.

AC K N OWLEDG N 1 E NTS

1 express my deepest gratitude to the many geologists,

paleontologists, and biologists who have influenced my

thinking about the history of the deep-sea biota in general

and, more specifically, my investigations of the mor¬

phology, ecology, relationships, and evolutionary' history

of basal marine gastropods. I am especially grateful to

Marla Coppolino for her patience and graphic expertise in

drafting the set of maps and to Dave Strauss for digital

images of specimens and assembly of final figures. Andrzej

Kaim provided a careful peer review with helpful com¬

ments. This is contribution number 2083 from the Uni¬

versity of California Museum of Paleontology.

LITERATUBE CITED

Aktipis, WAV. and G. Giribet. 2012. Testing relationships among

the vetigastropod taxa: A molecular approach. Journal of

Molluscan Studies 78: 12-27.

Andrews, B.C. 1943. Under a Lucky Star: A Lifetime of Ad¬

venture. Viking Press, New York, 300 pp.

Arruda, W. Z. and D. Nof. 2003. The Mindanao and Hahnahera

eddies - twin eddies induced by nonlinearities, journal ol

Physical Oceanography 33: 2815-2830.

Barber, A.| , S. Tjokrosapoetro, and T.R. Charlton. 1986. Mud

volcanoes, shale diapirs, wrench faults, and melanges in

accretionary complexes, Eastern Indonesia. American As¬

sociation of Petroleum Geologists Bulletin 70: 1729-1741.

Bouehet. P. 1991. New records and new species of Alnjssochrxjsos

(Mollusca, Caenogastropoda). Journal of Natural History 25:

30.5-313.

Bouehet, P. and J.-P. Rocroi (eds.). 2005. Classification and

Nomenclator of Gastropod Families. Malaeologia 47:

1-397.

Bowin, C., G.M. Purdy, C. Johnston, G. Shor. 1 , Lawver, H.M.S.

Hartono, and P. Jezek. 1980. Arc-continent collision in

Banda Sea region. American Association of Petroleum

Geologists Bulletin 64: 868-915.

Camplin, D.J. and R. Hall. 2014. Neogene history of Bone Gulf,

Sulawesi, Indonesia. Marine and Petroleum Geology 57:

88-108.

Cane, VI. and P. Molnar. 2001. Closing of the Indonesian seaway

as a precursor to east African aridification around 3-

4 million years ago. Nature 411: 157-162.

Carney, R.S. 1994. Consideration of the oasis analogy for che¬

mosynthetic communities at Gulf of Mexico hydrocarbon

vents. Geo-Marine Letters 14: 149-159.

Charlton, T.R. 1991. Postcollision extension in arc-continent

collision zones. Eastern Indonesia. Geology 19: 28-31.

Charlton, T.R. 2004. The petroleum potential of inversion an¬

ticlines in the Banda Arc. American Association of Petro¬

leum Geologists Bulletin 88: 565-585.

Charlton, T.R., A.J. Barber, and S.T. Barkham. 1991. The

structural evolution of the Timor collision complex, eastern

Indonesia. Journal of Structural Geology 13: 489-500.

Cordes, E.E., S. Hourdez, and H.H. Roberts. 2010. Unusual

habitats and organisms associated with the cold seeps of the

Gulf of Mexico In: The Vent and Seep Biota. Topics in

Geobiology 33: 315-331.

Crosnier, A.. B. Richer de Forges, and P. Bouehet. 1997. La

campagne KARUBAR en Indonesie, au large des lies Kai et

Tanimbar. In: A. Crosnier and P. Bouehet (eds.). Resultaats

des campagnes MUSORSTORM 16. Memoires du

Museum National d Histone Naturelle, Zoologie 172: 9-26.

D'Attilio, A. and G.E. Radvvin. 1971. The intritacalx, an unde¬

scribed shell layer in mollusks. The Veliger 13: 344-347.

C.S. Hickman, 2017

Page 161

Dickerson, R.E. 1924. Geologic aspects of Philippine hydrography.

Overdruk uit. llandelingen van het Derte Nederl.-Indisch

Naturwetenschappelijk Congres, gehouden te Buitenzorg

op 25, 26, 27 en 28 September 1924. Arehipel Drukkerij,

Buitenzorg, 28 pp.

Fu. I.-F. and C.-L. Sun. 2006. A new bathyal trochid from South

China Sea. Bulletin of Malacology, Taiwan 30: 17-20.

Gill, F.S., I.C. Harding, C.T.S. Little, and J.A. Todd. 2005. Pa¬

leogene and Neogene cold seep communities in Barbados,

Trinidad and Venezuela: an overview. Palaeogeography,

Palaeoclimatology, and Palaeoeeology 227: 191-209.

Gordon, A.L. and R. Fine. 1996. Pathways of water between the

Pacific and Indian oceans in the Indonesian seas. Nature

379(6561): 146-149.

Gordon, A.L., C.F. Giulivi, and A.G. Ilahude. 2003. Deep to¬

pographic barriers within the Indonesian Seas. Deep Sea

Research II 50: 2205-2228.

Gracia, A., N. Rangel- Buitrago, and | Sellanes. 2012. Methane

seep molluscs from the sinu-San Jacinto fold belt in the

Caribbean Sea of Colombia. |ournal of the Marine Bi¬

ological Association of the United Kingdom 92: 1367-1377.

Hall, R. 2001. Cenozoic reconstructions of Southeast Asia and

the SWi Pacific: changing patterns ol land and sea. In: I.

Metcalfe, J.M.B. Smith, M. Morwood and I. Davidson

(eds.). Faunal and Floral Migrations and Evolution in SF

Asia-Australasia. A. A. Balkema, Lisse, pp. 35-56.

Hall, R. 2013. The palaeogeography of Sundaland and Wallacea

since the Late Jurassic |ournal of Limnology 72(2s): 117.

Hall, R. and D.J. Blundell (eds.). 1996. Tectonic Evolution of SE

Asia. Special Publication 106. Geological Society of Lon¬

don, London, 556 pp.

Hall, R. and H R. Smyth. 2008. Cenozoic arc processes in

Indonesia: Identification of the key influences on the

stratigraphic record in active volcanic arcs. In: A.F. Draut,

P.D. Clift, and D.W. Scholl (eds.). Formation and Appli¬

cations of the Sedimentary Record in Arc Collision Zones.

Geological Society of America Special Paper 436: 27-54.

Hall, R. and M.E.J Wilson. 2000. Neogene sutures in eastern

Indonesia. Journal of Asian Earth Sciences 18: 781—808.

Hall, lb. M.A. Cot tarn, and M.E.J. Wilson. 2011. The SE Asian

gateway: history and tectonics of the Australia-Asia collision.

Geological Society, London, Special Publications 355: 1-6.

Harris, R. 2006. Rise and tall of the eastern Great Indonesian Arc

recorded by the assembly, dispersion and accretion ol the

Banda Terrane, Timor. Gondwana Research 10: 207-231.

Herbert, D.G. 2012. A revision of the Chilodontidae (Gastro¬

poda: Vetigastropoda: Seguenzioidea) of southern Africa

and the south-western Indian Ocean. African Invertebrates

53: 381-502.

Hickman, C.S. 2009a. Drawing lines in Wallacea: Historical

biogeography meets geophysics in the deep sea. Society for

Integrative and comparative Biology 2012 Annual Meeting

Abstracts, 1 1.2.

Hickman, C. S. 2009b. Relict deep-water gastropods in a dis¬

appearing seaway. Program and Abstracts, American

Malacological Society 75"' Annual Meeting. Paleontological

Research Institution Special Publication 37: 45.

Hickman, C.S. 2009c. Deep marine endemism in Wallacea: old

snails in a disappearing seaway. Southeast Asian Gateway

Evolution Programme and Abstracts. Southeast Asia Re¬

search Group, University of London, Royal Holloway

University, UK. p. 69.

Hickman, C.S. 2012. A new genus and two new species of deep-

sea gastropods (Gastropoda: Vetigastropoda: Gazidae). The

Nautilus 126: 57-67.

Hickman, G.S. 2016. New species ol deep-sea gastropods from

the Indo-Pacific Region (Vetigastropoda: Seguenzioidea:

Calliotropidae), The Nautilus 130: 83-100.

Hickman, G. S. and J. H. McLean. 1990. Systematic revision and

suprageneric classification of trochacean gastropods. Nat¬

ural History Museum of Los Angeles County Science Series

35: 1-169.'

Kano, V. 2007. Vetigastropod pliylogeny and a new concept of

Seguenzioidea: independent evolution of eopulatory organs

in the deep-sea habitats. Zoologica Seripta 37: 1-21.

Kano, Y., E. Chikyu, and A. Waren. 2009. Morphological,

ecological and molecular characterization of the enigmatic

planispiral snail genus Adeuomphalns (Vetigastropoda:

Seguenzioidea). Journal of Molluscan Studies 75:

397-418.

Kaim, A., R.G. Jenkins, and Y. llikida. 2009. Gastropods from

Late Cretaceous hydrocarbon seep deposits in Omagari and

Yasukawa, Nakagawa area, Hokkaido, |apan. Acta Palae-

ontologica Polonica 54: 463-490.

Kaim, A., R.G. Jenkins, K. Tanabe, and S. Kiel. 2014. Mollusks

from Late Mesozoic seep deposits, chiefly in California.

Zootaxa 3861(5): 401-440.

Kiel, S. 2010. On the potential generality of depth-related

ecologic structure in cold-seep communities: Evidence

from Cenozoic and Mesozoic examples. Palaeogeography,

Palaeoclimatology, Palaeoeeology 295: 245-257.

Kiel. S. and C.A. Campbell. 2005. Lithomphnlus enderlini gen.

et sp. nov. from cold-seep carbonates in California

a Cretaceous neomphalid gastropod? Palaeogeography,

Palaeoclimatology, Palaeoeeology 227: 232-241.

Kiel, S. and B.T. Hansen. 2015. Cenozoic methane-seep faunas

of the Caribbean region. PLoS ONE doi: 10. 1371/journal.

pone. 0140778.

Lydekker, R. 1896. A Geographical History’ of Mammals.

Cambridge University Press, Cambridge, 416 pp.

McLean, J.H. and | Quinn, J.F., Jr. 1987. Cataegis , new genus

and three new species from the continental slope (Tro-

chidae: Cataeginae new subfamily. The Nautilus 101:

111-116.

Mayr, E. 1944. Wallace’s line in the light of recent zoogeo¬

graphic studies. Quarterly Review of Biology’ 19: 1-14.

Metcalfe, 1. 2001. Paleozoic and Mesozoic tectonic evolution and

biogeography of SE Asia-Australasia. In: I. Metcalfe,

J.M.B. Smith, M. Morwood and I. Davidson (eds.). Faunal

and Floral Migrations and Evolution in SE Asia-Australasia.

A. A. Balkema, Lisse, pp. 15-34.

Metcalfe, 1. 2011. Paleozoic- Mesozoic history of SE Asia. In: R

Hall. M.A. Cottam and M.E. J. Wilson (eds.) The SE Asian

Gateway. History and Tectonics of the Australia - Asia

Collision. Geological Society’ of London, Special Publica¬

tions 355: 7-35.

Michaux, B. 2010. Biogeography of Wallacea: Geotectonie

models, areas of endemism, and natural biogeographical

units. Biological Journal of the Linnean Society 101:

193-212.

Moosa. M.K. 1984. Report on theCORINDON cruises. Marine

Research in Indonesia 24: 16.

Moroni, VI. A. 1966. Malacofauna del “Calcare a Lucine” di S.

Sofia, Forli. Palaeontographica Italica 60: 69-87.

Mortensen. 4’. 1923. The Danish expedition to the Kei Islands

1922. Videnskabelige fra Dansk Naturhistorisk Forening

76: 55-99.

Pholbud, P., R. Half E. Advokaat, P. Burgess, and A. Rudyawan.

2012. A new interpretation of Gorontolo Bay, Sulawesi.

Page 162

THE NAUTILUS, Vol. 131, No. 3

Proceedings ol the Indonesian Petroleum Association 36'1'

Annual Convention and Exhibition., 1 1 pp.. 15 figs.

Pownall. J.M., R Hall, and G.S. Lister. 2016. Rolling open

Earth’s deepest forearc basin. Geology 44: 947-950.

Pownall, J.M., R. Hall, and I.M. Watkinson. 2013. Extreme

extension across Seram and Ambon, eastern Indonesia:

Evidence for Randa slab rollback. Solid Earth 4: 277-314.

Schepinan, VI. M. 1908. The Prosobranehia of the “Siboga” Ex¬

pedition. Part 1. Rhipidoglossa and Docoglossa, with an

appendix by Prof. R. Bergh. In: Siboga Expeditie 1899-1900.

Vol. 49. Part 1. E.J. Brill, Leiden, pp. 1-107. pis. 1-9.

Schepinan, M M. 1909. The Prosobranehia of the “Siboga”

Expedition. Part 2. Taenioglossa and Ptenoglossa. In:

Siboga Expeditie 1899-1900. Vol. 49. Part 2. E. ]. Brill.

Leiden, pp. 1-123, pis. 10-16.

Smith, D.G. and J.T. Williams. 1999. The great Albatross

Philippine Expedition and its fishes. Marine Fisheries

Review 6 1 (4)>3 1 — 4 1 .

Smithsonian. 2016. Global Volcanic Program: http://www. volcano..

si.edu/. Holocene spread sheet 4.5.3 (December 2016).

Sudarmono. 2000. Tectonic and stratigraphic evolution of the

Bone Basin, Indonesia: insights into the Sulawesi collision

complex. In: Indonesian Petroleum Association, Pro¬

ceedings 27* 1 1 Annual Conventio, pp. 531-543.

Taviani, M. 1994. The “calcari a Lucina” macrofauna recon¬

sidered: Deep-sea faunal oases from Miocene-age cold

vents in the Romagna Apennine, Italy. Geo-Marine Letters

14: 185-191.

Tillinger, D. 2011. Physical oceanography of the present day

Indonesian ThroughHow. In: R. Hall, M.A. Cottam, and

M.E.J Wilson (eds). The SE Asian Gateway: History and

Tectonics of the Australia-Asia Collision Geological So¬

ciety of London. Special Publication 355: 267-281

Tydeman, G.F. 1903. Hydrographic results of the Siboga Ex¬

pedition. Siboga Expedition, Monograph III E.j. Brill.

Leiden, The Netherlands. 78 pp.

Van Aken, H.M. 2005. Dutch Oceanographic Research in

Indonesia in colonial times. Oceanography 18: 30—4 1 .

Van Aken, H.M.. I S. Brodjonegor. and I. Jaya. 2009. The deep¬

water motion through the Lifamatola Passage and its con¬

tribution to the Indonesian Throughflow. Deep-Sea Research

Part 1: Oceanographic Research Papers 56: 1203-1216.

Van Winkle, K. 1919. Remarks on some new species from

Trinidad. Bulletins of American Paleontology 8(33): 19-33.

Wallace, A.R. I860. On the zoological geography of the Malay

Archipelago. Journal of the Linnean Society Lotion 4:

172-184.

Wallace, A.R. 1863. VIII. On the Physical Geography of the

Mai ay Archipelago. Journal of the Royal Geographic So¬

ciety of London 33 (1863): 217-234.

Wallace, A.R. 1869. The .Malav Archipelago: The Land of the

Orang-utan, and the Bird of Paradise. A narrative of travel,

with studies of Man and Nature. London, Macmillan and

Co. 2 Volumes.

Waren, A. 201 1 . Molluscs on biogenic substrates. In: P. Bouchet.

H. Le Guvaderand O. Pascal (eds.) The Natural Historvof

Santo. Patrimonies Naturels, vol. 70. Museum national

d’Histoire naturelle, Paris; IRD, Marseille; Pro-Natura

International, Paris, pp. 438—448.

Waren. A. and P. Bouchet. 1993. New records of species, genera,

and a new family of gastropods from hydrothermal vents

and hydrocarbon seeps. Zoologica Scripta 22: 1-90.

Waren, A. and P. Bouchet. 2001. Gastropoda and Mono-

placophora from hydrothermal vents and seeps; New taxa

and records. The Veliger 44: 1 16-231.

Weber, M. 1902. Der Indo-australische Archipel und die /geschichte

seintr Tierwelt. Jena, 46 pp.

THE NAUTILUS 1 31 (3): 163-201, 2017

Page 163

A revision of the Hispaniolan genera Chondropomella ,

Chondropomium , and Clydonopoma (Gastropoda: Annulariidae),

with the recognition of a new genus, Superbipoma : phylogenetic,

radular, and conchologieal evidence

G. Thomas Watters

Department of Evolution, Ecology and Organismal Biology

Ohio State University

Columbus, OH 43212 USA

[email protected]

Paul Larson

Fish and Wildlife Research Institute

100 8th Ave SE

St. Petersburg FL, 33701 USA

Paul. I [email protected] n

ABSTRACT

The annulariid genera Chondropomella , Chondropomium ,

and Clydonopoma are believed to have originated from the

Tiburon/Barahona Peninsula in isolation lrom the rest ol

Hispaniola. Chondropomium has colonized the rift valley and

adjacent river valleys between the Tiburon Peninsula and the

remainder of Hispaniola. It is primarily limited to xeric low¬

lands, rarely found above 200 m elevation. Little is known

about the rare Chondropomella but they seem to occur in the

rift valley in xeric areas as well. In contrast, Clydonopoma is

endemic to the Sierra Baoruco with a single species in the

adjacent eastern Massif de la Selle and occupies the upland

mesic forests and pine savannahs between 200-4000 m. The

most widely distributed species, Chondropomium weinlandi ,

has been the subject of considerable confusion concerning the

nature of its many color forms or subspecies, as well as its valid

name. This species was investigated using phylogenetic

methods and compared to congeners and related genera. A

phylogenetic study aimed at elucidating relationships among

these taxa analyzed a partitioned matrix of nuclear (ITS 1) and

mitochondrial (COl. 12S, 16S) IONA sequences in a Maxi¬

mum-likelihood framework under the GTR + T substitution

model. Contrary to Bartseh’s 1946 assessment that C. wein¬

landi is a complex of subspecies, it is here shown to be a single,

highly polymorphic species for color. The new genus Super¬

bipoma is recognized based on phylogenetic, radular, and

conchologieal evidence. It contains two species: S. asymme-

tricum (Henderson and Simpson, 1902) and S. superbum

(Pilsbry, 1933). Eleven species of Chondropomium are rec¬

ognized including two new species: C. caelicum and C sar¬

donyx; three species of Chondropomella and nine species of

Clydonopoma , including one new species, Clydonopoma

titanium, are recognized. A calcified operculum is the ancestral

condition for the Annulariidae. In Clydonopoma and Chon¬

dropomella the operculum is a particularly complex calcified

structure termed the pseudolamella. This structure has been

lost in Chondropomium and Superbipoma.

Additional Keywords: Haiti, Dominican Republic, land snail,

DNA sequence, molecular systematics.

INTRODUCTION

Hispaniola consists of two historically separate geo¬

logical entities, the Tiburon Peninsula and the re¬

mainder of Hispaniola. The Tiburon Peninsula (termed

the Rarahona Peninsula in the Dominican Republic)

was an isolated island on the Caribbean Plate before

beginning its ongoing collision with Hispaniola in the

Pliocene (Miller and Miller, 2001). The intervening

space between the two land masses is now the rift valley

of Moya de Enriquillo in the Dominican Republic and

the Plain du Cul-de-Sac in Haiti. This includes several

lakes that may represent the last vestiges of ocean

between them: Lago Enriquillo, Lago del Rincon, and

Etang Saumatre.

The Annulariidae are widely distributed across both

of these parts of Hispaniola. However, phylogenetic

evidence (Skomroek, 2014) suggested that the faunas of

the two regions remained distinct, reflecting the ancient

isolation of the two parts and the highly endemic nature

of most species. The four genera reviewed here are part

of the Tiburon fauna and many are limited to narrow

ranges of the Sierra Raoruco on the Rarahona Penin-

sula. The species are overall quite similar to each other

and the genera were loosely defined by shell and

opercular characteristics. The phylogenetic relation¬

ships between the taxa were unknown. In particular,

several taxa display high degrees of shell color variation

that have been described as subspecies by some au¬

thors. Recent collections indicate that these pop¬

ulations are sympatric and cannot be subspecies, but the

question remained as to their relationships to each

other.

Like all annulariids, these species are calciphiles,

never found far from a limestone source. They com¬

monly occur in association with limestone knolls,

mogotes, fossilized reefs, and karst topography, which

are common in the Tiburon area. Recause of their

dependency on limestone many species are highly

Page 164

THE NAUTILUS, Vol. 131, No. 3

Figures 1-4. Opereula. 1. Clijdonopoma peasei (Pilsbry, 1933). Pseudolamellate operculum showing pseudolamella partially broken

away (rom underlying support, 7 mm length. 2. Clijdonopoma poloense (Bartseh, 1946). Pseudolamella at edge supported by erect

calcified ribs on corneous base. Scale bar = 400 [im 3, 4. Chondropomium mannoreum (Watters and Duffy, 2010). 3. Operculum with

calcareous deposit on outer surface, 5 mm length. 4. Calcareous deposit on outer surface. Scale bar = 100 |j.m, crystals at bottom of

micrograph are an artifact of fixation.

endemic, with outcrops acting as islands of speciation

among more inhospitable intervening areas. To the

north in central Hispaniola is the Cordillera Central,

which is predominately granitic and largely devoid of

annnlariids. But even there, species will be found in

calcareous areas where available.

Members of the four genera reviewed here are unusual

in several respects. They are large for the family (to ca.

35 mm in length) and have a shell sculpture consisting of

(usually) only polished axial ribs. This sculpture seems

adapted for a burrowing lifestyle beneath rocks and plants,

offering little resistance in the burrowing direction.

Clijdonopoma and Chondropomella have a highly spe¬

cialized type of operculum termed a pseudolamella

(Figures 1. 2). As in all annnlariids, the operculum is

attached to the foot as a thin, corneous layer. In Clydo-

nopoma and Chondropomella, the operculum is paucis-

piral, with closely-set, erect calcified ribs arranged in

a pin-wheel fashion on top of the corneous layer. The

distal edges of these ribs are fused to form a second

calcified surface parallel to the corneous base termed the

pseudolamella. In contrast, the operculum of Chon¬

dropomium and Superbipoma lacks a pseudolamella and

consists only of the corneous plate, although often

overlain with a layer of microscopic, calcified crystals

(Figures 3, 4). The genus Crossepoma (not reviewed here)

also has a pseudolamella, but in that genus there is a sulcus

separating the opercular whorls; the lip is also single in

Crossepoma rather than double as in Clijdonopoma and

Chondropomella .

This study reviewed these groups using phylogenetic,

radular, and conchological evidence. Although the family

as a whole was phylogenetically studied by Skomroek

(2014), this review looked at a much finer scale to dis¬

cern the' relationships not only of genera and species, but

individual populations.

Abbreviations and text conventions are: |): Used by

Thiele (1929) for the category Tribe; ANSP, Academy

of Natural Sciences, Philadelphia, USA; BMSM: Bailey-

Matthews National Shell Museum, Sanibel, USA; CMNH,

Carnegie Museum of Natural History, Pittsburgh, USA;

MCZ, Museum of Comparative Zoology, Harvard Uni¬

versity, Cambridge, USA; GTW, Collection of the author,

Columbus, USA; NI1MUK, Natural History Museum,

London, UK; USNM, National Museum of Natural

History, Washington, USA; OSUM, Ohio State University

Museum of Biological Diversity, Columbus, USA; UF,

Florida Museum of Natural History, Gainesville, USA;

ZMB, Zoologisehes Museum Berlin, Federal Republic of

Germany.

MATERIALS AND METHODS

Molecular Data Collection: Of the 24 taxa reviewed

here, suitable genetic material was only available for 25

specimens of seven species (Table 1). Most species have

either never been collected alive or if so, inadequately

preserved. The svmpatrie Crossepoma vermici datum

(Bartseh, 1946) is conchologically very similar to Chon¬

dropomium and, although the genus was not reviewed

here, was included in the phylogenetic analysis as a pos¬

sible' misplaced member of Chondropomium. A species of

Abhottella , now placed in its own subfamily (Abbottelli-

nae) within the Annulariidae (Watters, 2016b), was in¬

cluded as well. Members of the sister family Pomatiidae

were included as the outgroup.

DNA was extracted from foot tissues of preserved

specimens representing species from Pomatiidae (out¬

group) and Annulariidae (ingroup) using the DNeasy

(Qiagen Hiden, Germany) blood and tissue kit according

to the rodent tail protocol. Nuclear (ITS 1) and mito¬

chondrial gene regions (COl, 12S, 16S) were amplified

using new and previously published primers. Custom

degenerate primers were developed for 12S based on

those reported for Mollusca by Machida et al. (2012) as

follows: 12S forward primer sequence 5 -GTGCCAG-

CADYYGCGGTYA-3’; reverse primer sequence 5-

AGRGYGACGGGCGATDTGW-3’. Partial Cytochrome

Oxidase subunit 1 was amplified using universal primers

LCO1490, HC02198 (Folmeretal., 1994); 16S and ITS 1

were amplified with the mollusc-specific primers pub¬

lished by Pfenninger et al. (2005).

G.T. Watters and P. Larson, 2017

Page 165

Multiple-band PCR results occurred for ITS and 12S in

some but not all individuals. In these eases, the band

matching the length of single-band results in other

specimens (~600bp for 12S, ~300bp for ITS) was

extracted via Qiaquick gel extraction kit (Qiagen).

Extracted DNA was then subjected to a second round of

PCR and gel electrophoresis. PCR products were se¬

quenced by GeneWiz (South Plainfield, New Jersey). All

DNA sequences were searched against the NCBI nu¬

cleotide database using the BLAST (Altschul et al., 1990).

Any sequence with the top BLAST result not matching

a gastropod was excluded from the analysis.

Phylogenetic Analysis: Trimmed sequences for each

locus were aligned by MUSCLE (Edgar, 2004) imple¬

mented through Geneious v.7 1 .9 (Biomatters) and using

default parameters. A complete data matrix was tested for

an optimal partitioning scheme and a nucleotide sub¬

stitution model in PartitionFinder (Lanfear et al., 2012)

under the AICc. Possible partitions were defined as each

individual locus plus three codon positions of COl,

resulting in a maximum of six possible partitions. Using

the OSU Ohio Biodiversity Conservation Partnership

computing cluster, the sequence matrix was analyzed in

RAxML v. 8. 1.16 (Stamatakis, 2014). Twenty independent

runs were followed by 10,000 bootstrap replicates on the

best scoring tree to assess elade support. Each sequenced

gene region was also analyzed independently in the same

manner as the complete data matrix. A second analysis of

the complete matrix was carried out in MrBayes v. 3.2.6

(Ronquist and Huelsenbeck, 2003). The Bayesian analysis

was also run on the full, partitioned data set using the

GTR + E model for two million generations with 500,000

discarded as bum-in. Prior distributions were left as de¬

faults and trees were sampled every 500 generations.

Radulae: Radulae were available for eieht of the taxa.

O

Radulae were removed from ETOH preserved specimens

and placed in commercial bleach until all attached tissue

had been dissolved (typically 10-15 minutes). The

remaining radular ribbon was washed in two changes of

100% ETOH and placed on an SEM stub. Specimens

were sputter-coated with gold-palladium and viewed with

a Philips XL30 scanning electron microscope at an ac¬

celerating voltage of 20 KeV.

More than 4000 shells were examined. Descriptions and

measurements were based on shells oriented with the spire

up and the aperture facing the viewer. Measurements are

for adult shells unless noted otherwise. Length was mea¬

sured from the tip of the protoconch (or teleoconch of

decollate specimens) to the opposite anterionnost exten¬

sion of the outer lip. Subsets of the largest and smallest

adult specimens were selected by eye from all available

specimens and measured to determine the minimum and

maximum lengths. The number of whorls was determined

using the I D method of Van Osselaer (1999).

Original descriptions from non-English sources are

translated and given with each species review. Specimens

used in the phylogenetic portion are highlighted in bold

text in the “Other Material Examined" section of each

species. Numbers in parentheses after catalog numbers

are the number of specimens in the lot.

RESULTS

Sequences for two or more loci were obtained for 25

specimens. All sequences used in the analysis have been

submitted to GenBank (see Table 1 for accession num¬

bers). The complete data set was an aligned, concatenated

matrix of 2369 nucleotides in length. The optimal scheme

identified by PartitionFinder included all six separate

partitions and GTR + L as the favored partition scheme

and substitution model, respectively.

All 20 ML replicate runs of the full data set produced

an identical tree topology. Bootstrap support was gen¬

erally high at the genus level (supporting generic

monophyly) but poor at intermediate nodes. All sub¬

species (morphotypes) or putative species of Chon-

dropomium weinlandi formed a single monophvletic

elade, but morphotypes within the group did not segre¬

gate. We recovered high support for a sister relationship

between individuals of different types to the exclusion

of their comorphs (e.g., 94% bootstrap support for

C. weinlandi “ azuense ” and C. weinlandi " barahonense "

and 99% for another C. weinlandi “ azuense ” with C.

weinlandi) (Figure 5). This result is consistent with

a single diverse species rather than subspecific radiations.

The topology of all four ML single-gene trees differed

from each other with respect to genus-level relationships,

but in most cases the putative genera ( Crossepoma ,

Clijdonopoma, Chondropominm, and Snperbipoma new

genus) are recovered as monophvletic groups. Exceptions

are that in the 12S gene tree, two individuals of Chon-

dropomium nest within a poorly supported (21% boot¬

strap) Crossepoma elade, while Clijdonopoma species are

distributed among Snperbipoma and Chondropominm

species (20% bootstrap), and in the COl gene tree,

Snperbipoma new genus is recovered as a paraphyletic

grade at the base of the ingroup. As in the combined

analysis, putative species in the Chondropominm wein¬

landi elade fail to segregate based on shell morphology

regardless of the gene region analyzed. Only in the ITS

gene tree do individuals of Chondropominm superbum

and Chondropominm assymetricnm (= Snperbipoma new

genus) form a weakly supported sister- relationship with

Chondropominm weinlandi. All other genes support a

Chjdonopoma-Chondropomium weinlandi sister-relationship

(COl, 16S) or a Clydonopoma-Superbipoma sister-

relationship (12S). The Bayesian analysis was ended af¬

ter two million runs with a final average standard deviation

of split frequencies of O.OOfi.The pertinent results of the

Bayesian analysis agree with those of the ML analysis, with

a strongly supported (100% posterior probability) relation¬

ship between the Chondropomium weinlandi and Clydo-

nopoma clades and species of Chondropominm weinlandi

failing to segregate by morphotype.

Reconstruction of ancestral characters suggest that

calcification of the operculum and the presence of

Page 166

THE NAUTILUS, Vol. 131, No. 3

Figure 5. Inferred relationships

condition of the inner marginal rac!

Bar represents branch length at w

from maximum likelihood analysis of DNA sequence data. Symbols at the base of clades mark the

lular tooth for all terminals in that clade. Bootstrap support values appear above or left of branches,

hich 0.05 substitutions per nucleotide site have occurred.

a pseudolamella are both ancestrally present in the taxa of

Annulariidae reviewed here (pseudolamella does not occur

in Abbott ella). Two subsequent losses of flu1 pseudolamella

in Superbiponm and Chondnypomium (proportional like¬

lihood of 0.81 and 0.79, respectively, for “presence” in

nodes immediately preeeeding the loss' bars in Figure 5)

are inferred. While the calcification of the operculum has

been reconstructed as possibly a loss with subsequent gain

in Clydonopoma , it is more likely that the pseudolamella

persisted in that genus. The conchologically similar

Crosseporna vermieulatum, which possesses a pseudola¬

mella, included as potentially a Chondropomiwn , is shown

to belong to a genus apart from those reviewed here.

Radul ae delineated the same genera as identified here

by phylogenetic and conchological characteristics. The

inner marginal was the most variable between taxa and of

the most systematic importance. The remaining radular

teeth showed remarkable uniformity, with some ex¬

ceptions. A more extensive study of the radulae of 111

annulariid species found similar results (Watters, in

prep.). In that study, as here, Abbottella (and related

genera) differed in radular characteristics from most

other annulariids. Abbot t§Ua was raised to subfamily rank

within the Annulariidae by Watters (2016b).

SYSTEM ATICS

Family Annulariidae Henderson and Bartsch, 1920

Genus Chondropomium Henderson and Bartsch,

1920

Type Species: Cdiondropoma weinlaruh Pfeiffer, 1862,

by original designation.

Description: Shells medium to large for family (to ca.

25 mm length), elongate-conic, usually decollate. Proto¬

conch of 1.5 smooth, minute whorls. Final whorl shortly

detached from previous whorl. Axial sculpture of wide,

flattened, close-set ribs. Spiral sculpture absent on final

whorl but present in umbilicus; rarely on earlier whorls.

Overall sculpture very smooth, often polished. Suture

minutely serrate lacking fused tufts. Adult with very

narrowly reflected lip. Lip single or double; when double,

G.T. Watters and P. Larson, 2017

Page 167

inner lip usually fused to outer lip. Color pattern of hands

and blotches, continuous or interrupted; rarely lacking

any pattern. Operculum paucispiral. with uniform, thin

granular deposit. Taenioglossate radula with rachidian

tooth, single pair of lateral teeth, and two pairs of marginal

teeth. Rachidian and lateral teeth usually unicuspid. Inner

marginal tooth deeply multicuspid on outer side. Outer

marginal tooth pectinate. Animal with foot longitudinally

bisected into lobes. Locomotion ditaxic between lobes of

foot. Eves at base of tentacles; bifid snout produced into

short secondary tentacles.

Remarks: This genus differs from Clydonopoma and

Chondropomella in lacking a pseudolamellate operculum

and in having deeply incised inner marginal radular teeth.

Phylogenetic results corroborate the separation of Chon-

dropomium from the other taxa reviewed here (Figure 5).

Chondropomium was originally described as a subgenus

of Clmndropoma, but was raised to generic status by

Watters (2006). As defined here, Chondropomium is

a Tiburon peninsular group. Most Tiburon annulariids are

limited to that region but species of Chondropomium have

colonized the rift valley and associated river valleys now

separating that peninsula from the remainder of Hispaniola.

It is perhaps significant that, with one exception, these

colonizing taxa are limited to the xerie lowlands of these

\ alleys, generally below 200 m, and have not yet adapted to

the montane regions of remaining Hispaniola. The ex¬

ception is C. caelicum, which occurs from 200-800 m in the

mountains on either side of the Rio San |uan valley.

Bartseh (1946) included Chondroponui eusarcum

Pfeiffer, 1854, in Chondropomium. He recognized three

subspecies; one from Puerto Plata, one from Isla Cata-

linita, and the nominate subspecies from an unlocalized

region. A forth, unnamed relative occurs in Altagracia

Province. Puerto Plata and Isla Catalinita/Altagracia are

separated by >270 km and it is bighlv unlikely that

Bartsch’s taxa are conspecific. All of these taxa resemble

Chondropomium but are far removed from the Tiburon

area and adjacent valleys. These species differ from

Chondropomium in having lamellate axial sculpture, at

least on early whorls, rather than flattened threads or

cords. No material was available for phylogenetic study.

Pending additional study these are here regarded as

species of Chondroponui superficially similar to Chon¬

dropomium. Should they be found to belong to Chon¬

dropomium it would indicate that that group is much

more widespread on the island than described here.

With few exceptions, members of Chondropomium live

in very xerie habitats characterized by sparse vegetation

(cacti, agave, thorny bushes, small clumps of grass) and

rocky soils. They estivate under seemingly solidly em¬

bedded rocks or under dense mats of dead grasses and

agave. They probably emerge during periods of rain or

possibly at night. The nearly smooth shells have only weak-

axial sculpture that would minimize drag while burrowing

and which seems to be an adaptation for this lifestyle.

Individuals are rarely found alone; a half dozen or more

specimens of a species may be found under the same rock

and they may be locally abundant at a site. Estivating live

individuals are often accompanied by numerous dead

shells, suggesting that the snails either died of natural

causes while estivating or were killed by some burrowing

predator. Dead specimens often have drilled holes

through the shell that may be the result of predation by

the burrowing larvae of lampyrid and elaterid beetles.

Individuals of Chondropomium species show an un¬

usually broad range of shell colors for the family. Within

a population specimens may range from a nearly uniform

dark color (often brown or orangish) with conspicuous

markings to all white with no markings. This is particularly

evident in C. weinlandi, discussed at length below.

Based on its position in the phylogenetic tree presented

here (Figure 5), Chondropomium is one of the most

recently derived members of the genera reviewed here.

Like Superhiponm , the operculum has lost the ancestral

pseudolamella and been reduced to a fine granular

deposit.

Chondropomium beatense (Clench, 1932)

(Figures 6-11, 183)

Chresoxymy

Chondroponui (Chondropomium) beatensis Clench,

1932: 106; Bartseh, 1946: 33-34, pi. 5, fig. 3.

Chondroponui (Chondropomella) beatensis Clench, 1932:

Bartseh, 1932: 3, pi. 1, figs. 7, 9.

Chondroponui ( Chondropomium ) beatensis armouri

Clench, 1932: 106; Bartseh, 1932: 3 [in synonymy of

Chondroponui beatensis Clench, 1932]; Bartseh, 1946:

33 [in synonymy of Chondroponui beatensis Clench,

1932]; Watters, 2006: 167 [in synonymy of Chon¬

droponui beatensis Clench, 1932],

Chondroponui sp. indet.: Brooks, 1936: 124, text fig.

Chondropomium beatense (Clench, 1932): Watters, 2006:

63, 166.

Description: Shell medium for family (hu'gest adult

specimen 16.4 mm maximum length, decollate, including

peristome; smallest adult specimen 9.5 mm length, de¬

collate, including peristome), solid, conic, umbilicus small,

narrow. Protoconch of 1.5 smooth whorls, not well-

differentiated from teleoconeh, rarely retained in adults.

Teleoconeh of 3.5 4 whorls, final whorl detached for last 1/

6th of whorl. Suture narrow, channeled. Peristome single or

rarely double, tear drop-shaped. Outer lip narrow, weakly

expanded, minute posterior auricle present, narrowly de¬

tached from previous whorl. Spiral sculpture absent except

for few very weak, almost obsolete cords within umbilicus.

Axial sculpture of narrow, low ribs, separated by incised

grooves, ea. 120 ribs on final whorl, fonning minute ser¬

rations at suture. Background color very variable, from white

to clouded with orange or yellow, often overlain with con¬

tinuous or interrupted spiral brown bands, chevrons, and

blotches; orange individuals may have faint pale, vertical

zones; umbilicus and lip white. Operculum paucispiral with

thin calcareous deposit. Anatomy and radula unknown.

Page IBS

THE NAUTILUS, Vol. 131, No. 3

Figures 6-33. Chondropomium species. 6—11. Chondropomium beatense (Clench, 1932). 6, 7. USNM 414227, paratype, 13.4 mm.

8. UP’ 216536, 16.7 mm. 9. UF 216536, 17.4 mm. 10. Chondropoma beatensis armouri Clench, 1932. USNM 414228, paratype, 14.5

mm. 11. UF 216539, 9.7 mm. 12-17. Chondropomium blaineorum Watters, 2012. 12, 13. UP" 446070, holotype, 13.0 mm. 14. UF

45726, 17.1 mm. 15. UF 45726, 14.4 mm. 16. UP’ 45726, 17.9 mm. 17. UP" 216507. 17.2 mm. 18-24. Chondropomium caelicum new

species 18, 19. UP" 216868, holotype. 18.9 mm. 20. UF 505818, paratype 1. 17.1 mm. 21. UP’ 505818, paratype 2, 14.4 mm. 22.

UP" 505818, paratype 3, 17. 1 mm. 23. UP" 505818, paratype 5, 14.9 mm. 24. UP’ 216812, 16.3 mm. 25-28. Chondropomium gimbiense

(Bartsch, 1946). 25, 26. Chondropoma (Chondropomium) gimbiense gimbiense Bartsch, 1946. USNM 471935, holotype, 10.2 mm. 27,

28. Chondropoma (Chondropomium) gimbiense saltrouense Bartsch, 1946. USNM 471936, holotype, 14.0 mm. 29, 30. Chon¬

dropomium ignotum (Bartsch, 1946). USNM 471940, holotype, 16.6 mm. 31-33. Chondropomium lynx Watters, 2012. 31, 32. UF

446072, holotype, 19.6 mm. 33. OSUM 36519, paratype, 19.0 mm.

G.T. Watters and P. Larson, 2017

Page 169

Type Material: Chond ropoma beatensis Clench, 1932:

Holotype: MCZ 81493; Paratypes: MCZ81494(?); USNM

414227(2) (but not listed as types in database); ANSP

157702(2). Clench (1932) only listed four paratypes,

without catalog numbers or collection identification; it is

not clear which of the MCZ, USNM, or ANSP specimens

are actual paratypes; Chondropoma beatensis annouri

Clench, 1932: Holotype: MCZ 81495; Paratype: USNM

414228(1).

Type Locality: Chondropoma beatensis Clench, 1932:

“Beata Island, Santo Domingo. " Chondropoma beatensis

armouri Clench, 1932: “Beata Island.”

Type Figured: Chondropoma beatensis Clench, 1932:

unfigured. Chondropoma beatensis armouri Clench,

1932: un figured.

Distribution: Endemic to Isla Beata, a uninhabited is¬

land off the Barahona Peninsula.

Habitat: Coastal limestone cliffs.

Other Material Examined (152 Specimens): Do¬

minican Republic. GTW 7261a(l), NE part of Isla

Beata; USNM 414227(2), USNM 414228(1), UF 216536

(23), UF 216537(71), UF 216538(8), Isla Beata; UF

216539(46), N point, Isla Beata.

Variation Among Specimens: Specimens appear to

come in two color forms with intergradation. One form

has brown patterns on a white background, the other is

lacks a pattern or nearly so, with a yellow or orange

background.

Comparison with Other Species: This species is most

similar to C. blaineomm Watters, 2012, but differs in the

more diffuse, less defined color pattern that is not

arranged in a mosaic configuration.

Remarks: Chondropoma beatensis armouri was de¬

scribed for the orange, patternless color form; we do not

consider it distinct. Other Hispaniolan annulariid species,

originally believed to be endemic to an island, have sub¬

sequently been found on the adjacent mainland:

Abbottella milleacantha Watters and Duffy, 2010, de¬

scribed from Isla Saona, and Colonina dominicensis

(Pfeiffer, 1850) from Isla Catalina have both been found

on the coastal mainland (Watters and Frank-Fellner,

2017). H owever, C. beatense has not been found on

the adjacent mainland as yet. No material was available

for phylogenetic study.

Etymology: As the root endings -ponui and -pomium are

neuter, the correct orthography should be beatense.

Chondropoma beatense Clench, 1932: Isla Beata, Do¬

minican Republic. Chondropoma beatense armouri

Clench, 1932: Allison V. Armour (1863-1941), archeol¬

ogist, botanist, herpetologist associated with the Chicago

Field Museum; his yacht Utowana , a one time fishing

trawler and WAV I Atlantic escort craft, visited Isla Beata

on 14-15 February 1929 where Thomas Barbour of MCZ

collected the types of both of these taxa. An excellent

account of the expedition is found in Henderson and

Powell, 2004.

Chondropomium blaineorum Watters, 2012

(Figures 12-17, 182)

ChRESOXYMY:

Chondropomium blaineorum Watters, 2012: 9, figs. 34, 35.

Description: Shell medium to small for family (largest

adult specimen 18.0 mm maximum length, decollate,

including peristome; smallest adult specimen 12.9 mm

length, decollate, including peristome), solid, conic,

umbilicus minute. Protoconch whorls unknown, decollate

in all specimens examined. Teleoeoneh of 3-3.5 whorls,

final whorl detached for last 1 /6th of whorl. Suture nar¬

rowly channeled. Peristome single or weakly double, tear

drop-shaped. Outer lip narrow, not expanded, minute

posterior auricle present. Spiral sculpture present only as

7-8 very feeble cords within umbilicus. Axial sculpture of

narrow, closely-spaced, low ribs, nearly obsolete on final

whorl, forming minute cusps at suture. Suture serrate.

Background color pale tan to brown with narrow darker

bands having zigzag markings between them axially

arranged into tile-like pattern; umbilicus and lip white.

Operculum paucispiral with thin calcareous deposit.

Anatomy and radula unknown.

Type Material: Holotype: UF 446070; Paratype: OSUM

36516(1).

Type Locality: “ca. 9.3 km NW of Manuel Golla, ea. 140

m, off Highway 44, Pedernales Province, Dominican

Republic. 1 7.94 N, -71.65 W.”

t ype Figured: Watters, 2012, figs. 34, 35.

Distribution: With a single exception at Laguna de

Oviedo, all specimens have been collected along RD

Highway 44 between Pedernales and Oviedo. This road

runs along the southern foothills of the Sierra Baoruco,

from nearly sea level to 200 m.

Habitat: Under rocks on hillsides with limestone

outcrops.

Other Material Examined (86 Specimens): Domini¬

can Republic. Pedernales Province. UF 216457a(2),

8 km SF of Pedernales; UF 216400(2), 23 km SF of

Pedernales; UF 216505(21), 26 km NW of Oviedo;

UF 45726(7), UF 216507(9), 18 km NW of Oviedo; UF

216779(3), UF 216780(3), 17 km NW of Oviedo; UF

216778(26), 13 km NW of Oviedo; GTW 7082c(2), 10 km

NW of Oviedo; UF 216781(1), 8 km NW of Oviedo; UF

216773a(12), 7.5 km NW of Oviedo; UF 467467(2), S end

of Laguna de Oviedo.

Variation Among Specimens: The peculiar mosaic or

tile-like color pattern varies from well-developed to

Page 170

THE NAUTILUS, Vol. 131, No. 3

smudged and indistinct depending on the background

color.

Comparison With Other Species: This species is very

similar to Chain! ropoma quisquense Bartseh, 1946, in

color patterns and shape and may be sympatric with it in

some places, but lacks the spiral sculpture of that species.

It is also svmpatric with Chond ropomium marmoreum but

lacks the spiral threads on the spire. See also under C.

heatense.

Remarks: The original images of this species in 2012

were unfortunately vertically foreshortened by the

printer. No material was available for phylogenetic study.

Etymology: Matt and Dona Blaine (Delaware, USA),

who helped collect the tvpe lot.

Chondropotnium caelicum new species

(Figures 1 S— 24. 184)

Description: Shell medium to small for family (largest

adult specimen 19.8 mm maximum length, decollate,

including peristome; smallest adult specimen 12.0 mm

length, decollate, including peristome), solid, conic,

polished, umbilicus small, narrow. Protoconch unknown,

decollated in all examples seen, not retained in adults.

Teleoconeh of 3.75—4.25 whorls, final whorl detached for

last l/8th of whorl. Suture narrow, channeled. Peristome

single, tear drop-shaped. Lip narrow, very weakly ex¬

panded, posterior auricle absent, narrowly detached

from previous whorl, forming short keel posteriorly.

Spiral sculpture absent except for few very weak, almost

obsolete cords within umbilicus. Axial sculpture of flat,

low ribs, separated by incised grooves, ea. 120 ribs on

final whorl, forming minute serrations at suture; no fused

tufts. Axial ribs gathered into growth series having ea. 10

ribs/series. Background color very variable, base color

from white to clouded with orange or yellow; brown axial

bands on trailing edge of growth series have regular tri¬

angular or dashed markings that form interrupted spiral

pattern; shell usually pale overall but rarely darkly colored.

Umbilicus and lip white. Operculum paucispiral with thin

calcareous deposit. Anatomy and radula unknown.

Type Material: Holotype: UF 216868, 18.9 mm; Para-

type 1: UF 505818, 17.1 mm; Paratype 2: UF 505818,

14.4 mm; Paratype 3; UF 505818, 17.1 mm; Paratype 4:

UF 505818, 13.4 mm; Paratype 5: UF 505818, 14.9 mm.

Type Locality: 200 m elevation, 2 km NW of Cortes, San

Juan Province, Dominican Republic.

Distribution: From 200-800 m, in calcareous regions of

the Cordillera Central and the Sierra de Neiba bordering

the Rio San Juan and Rio Macasia valleys.

Habitat: Found under limestone blocks in sub-mesic

scrub and in mesic forest.

Other Material Examined (134 Specimens): Do¬

minican Republic. San Juan Province. UF 216868(42),

200 m, 2 km NW of Cortes; UF 216807(42), 450 m, 1 km

SSW of Cardon; UF 216841(1), 780 m, 5 km S of Vallejuelo;

UF 216819(7), 710 m, 4 km E of Vallejuelo; UF 216812(21),

540 m, 4 km SW of Cardon on road to Vallejuelo; UF

216818(1), 690 m, 15 km NNW of Juan de Herrera; UF

216820(1), 540 m, 4 km NW of Bohechio. La Estrelleta

Province. UF 216806(18), 790 m, 6 km SW of Hondo

Valle.

Variation Among Specimens: As in Chondropomium

weinlandi , this species shows great variation in color

pattern but unlike that species the patterns form a con¬

tinuum and do not seem to be separated into “forms.”

Comparison With Other Species: Chxmdrofmniium

caelicum new species is most similar to C. weinlandi. It differs

in having the color pattern axially arranged rather than spirally

aligned; C. weinlandi rarely has axial color elements.

Remarks: This species is unique in several regards. It lias

colonized the furthest from the Tiburon Peninsula of all of

the taxa reviewed here. It continues from the range of C.

weinlandi north into the valleys of the Rio San Juan and

Rio Macasia. But unlike C. weinlandi and other Chon-

d ropomium it occurs in upland sub-mesic and mesic

forests rather than xeric lowlands. No material was

available for phylogenetic study.

Etymology: L. caelicum. heavenly, for its comparatively

montane habitat.

Chondropomium gimbiense (Bartseh, 1946)

(Figures 25-28, 184)

Chresonymy

Chondropoma ( Chondropomium ) gimbiense gimbiense

Bartseh, 1946: 34, 35, pi. 6, fig. 3.

Chondropoma (Chondropomium) gimbiense saltrouense

Bartseh, 1946: 34, pi. 6, fig. 2.

Chondropomium gimbiense (Bartseh, 1946): Watters,

2006: 63.

Chondropomium gimbiense gimbiense (Bartseh, 1946):

Watters, 2006: 272.

Chondropomium gimbiense saltrouense (Bartseh. 1946):

Watters, 2006: 272.

Licina gimhiensis (Bartseh, 1946): Watters. 2013: map 1

Description: Shell small for family (largest adult speci¬

men 14.0 mm maximum length, decollate, including

peristome; smallest adult specimen 10.2 mm length, non-

decollate, including peristome), solid, conic, umbilicus

minute. Protoconch whorls of 1 .5 smooth, minute whorls,

usually decollate in adults. Teleoconeh of 4 whorls, final

whorl only shortly detached before aperture, forming

short posterior keel. Suture minutely channeled. Peri¬

stome weakly double, tear drop-shaped. Peristome dou¬

ble. Inner lip not erect, fused to outer lip; outer lip

narrow, not expanded, minute posterior auricle present.

Spiral sculpture present only as few very feeble cords

within umbilicus. Axial sculpture of numerous, narrow,

G.T. Watters and P. Larson, 2017

Page 171

closely-spaced, low ribs, separated by narrow incised

grooves, forming minute cusps at suture; slightly enlarged

cusps occur in groups of 5-6 at irregular intervals.

Background color white to brownish-purple with tan axial

zig-zags; these markings may be spirally aligned on earlier

whorls; early whorls may be darkly colored; umbilicus and

lip white; interior of aperture brown in some specimens.

Operculum paueispiral with thin calcareous deposit.

Anatomy and radula unknown.

Type Material: Chondroponui ( Chondropomium ) gim-

biense gimbiense Bartsch, 1946: Holotype: USNM

471935. Chondroponui ( Chondropomium ) gimbiense

saltrouense Bartsch, 1946: Holotype: USNM 471936.

Type Locality: Chondroponui ( Chondropomium ) gim¬

biense gimbiense Bartsch, 1946: “East bank of the Riviere

Gimbi in the vicinity of Saltrou.” Chondroponui ( Chon -

dropomium) gimbiense saltrouense Bartsch, 1946: “West

side of the Riviere Gimbi."

T>pe Figured: Chondroponui (Chondropomium) gim¬

biense gimbiense Bartsch, 1946: Bartsch, 1946, pi. 6, fig. 3.

Chondroponui ( Chondropomium ) gimbiense saltrouense

Bartsch, 1946: Bartsch, 1946, pi. 6, fig. 2.

Distribution: Known only from the Riviere Gimbi valley

in the vicinity of Saltrou (now Belle-Anse) in the Sud-Est

Department of Haiti on the southern coast of the Tiburon

Peninsula.

Habitat: Not reported.

Other Material Examined (19 Specimens): Haiti.

Departement du Sud-Est. USNM 471936(10), W side

of the Riviere Gimbi; USNM 402015(9), E bank of the

Riviere Gimbi in the vicinity of Saltrou.

Variation Among Specimens: Specimens differ in the

depth of background coloration and degree of elongation.

Comparison With Other Species: This species differs

from its congeners in its combination of silky texture and

vertically aligned color pattern. In this respect it re¬

sembles some Crossepoma , but is differentiated from that

genus by its non-pseudolamellate operculum. It is geo¬

graphically distant from any other known congener.

Remarks: No material was available for phylogenetic

study.

Etymology: Chondroponui (Chondropomium) gimbiense

gimbiense Bartsch, 1946. Riviere Gimbi, Haiti. Chon-

dropoma (Chondropomium) gimbiense saltrouense Bartsch,

1946. Saltrou, Haiti.

Chondropomium ignotum (Bartsch, 1946)

(Figures 29, 30)

Chresonymy

Chondroponui (Chondropomium) ignotum Bartsch, 1946:

31, pi. 5, fig. 7.

Chondropomium ignotum (Bartsch, 1946): Watters, 2006:

63, 300.

Description: Shell medium for family (only known

specimen 16.6 mm maximum length, decollate, in¬

cluding peristome), solid, inflated conic, umbilicus

minute. Protoconch unknown, decollate. Teleoeonch of

3.25 whorls, final whorl only shortly detached before

aperture, forming short posterior keel. Suture narrowly

channeled. Peristome double, oval. Inner lip narrowly

erect, not fused to outer lip; outer lip evenly expanded,

fairly wide, low posterior auricle present. Spiral sculp¬

ture present only as well-developed cords within um¬

bilicus. Axial sculpture of numerous (ea. 120), wide,

closely-spaced, flattened ribs, separated by narrow in¬

cised grooves, forming irregular, minute cusps at suture.

Background color white with seven tan, spiral bands

alternating between spots and nearly continuous bands on

final whorl; last band borders umbilicus; some bands

continue onto both sides of lip. Operculum, anatomy, and

radula unknown.

Type Material: Holotype: USNM 471940.

Type Locality: “Haiti?"

Type Figured: Bartsch, 1946, pi. 5, fig. 7.

Distribution: Described from “Haiti?" without further

information.

Habitat: Not reported.

Variation Among Specimens: Known only from the

holotype specimen.

Comparison With Other Species: This species is

known from the single, unloealized holotype specimen.

It clearly is related to taxa such as C. weinlandi and C.

caelicum but the expanded outer lip is unique. Super-

bipoma species also have an expanded lip, but the lip of

C. ignotum is double with a narrow, distinct lamella

rather than a reflected portion of a single lip as in

Superbipoma.

Remarks: Although it possibly is a teratological individual

of another species, we believe it is distinct but awaiting

rediscovery. No material was available for phylogenetic

study.

Etymology: L. ignotus , strange, unnoticed.

Chondropomium lynx Watters, 2012

(Figures 31-33, 182)

Chresonymy

Chondropomium lynx Watters, 2012: 10-11, figs. 39-41,

not 58 [habitat, mislabeled locality].

Description: Shell medium for family (largest adult

specimen 19.6 mm maximum length, decollate, including

peristome; smallest adult specimen 19.0 mm length.

Page 172

THE NAUTILUS, Vol. 131, No. 3

decollate, including peristome), solid, conic, umbilicus

minute. Protoconch whorls unknown, decollate in all

specimens examined. Teleoconch of 4 whorls, final whorl

detached for last 1 /4th of whorl. Suture narrowly chan¬

neled. Peristome single, tear drop-shaped. Lip narrow,

not expanded, minute posterior auricle present, detached

from previous whorl. Spiral sculpture present only as ea. 3

very feeble cords within umbilicus. Axial sculpture of

numerous (ea. 90) fairly wide, closely-spaced, low ribs,

strongest on last 14 of final whorl, forming minute cusps at

suture. Background color pale tan with 5 interrupted

bands, 3 above, one at, and one below periphery, com¬

posed of dark brown spots blurred and repeated; first

teleoconch whorl dark brown; lip white. Operculum

paueispiral with thin calcareous deposit. Anatomy and

radula unknown.

Type Material: Holotype: UL 446072; Paratypes:

OSUM 36518(1); OSUM 36519(1).

Type Locality: Originally described from the area of

Virgen de San Rafael and Los Patos in Barahona Province,

these specimens are now known to have been mislabeled.

The type locality (and that of all known specimens) is here

corrected to Dominican Republic, Peravia Province,

Punta Salina, 21 km W of Bani. This site is just north of

RD 2 (Carretera Lrancisco del Rosario Sanchez) on

a series of low hills on the west bank of the Rio Oeoa.

Type Figured: Watters, 2012, figs. 39, 40.

Distribution: Known only from the area of the type locality.

Habitat: Under rocks on low xeric hills (< 200 m) with

cacti and agave. This species co-ocurs with Tessaripoma

hooksi (Watters and Duffy, 2010) but appears to be much

rarer.

Other Material Examined (6 Specimens): GTW

16671a(2), GTW 16671b(2), GTW 16671c(2), all from the

type locality.

Variation Among Specimens: The few known speci¬

mens are very uniform in characteristics, varying mainly in

the degree of background coloration.

Comparison With Other Species: Chondropomium

lynx is very similar to C. weinlaruli , particularly the color form

known as barahonensi s. It differs in its more elongate shape,

dark earlv whorls, and different banding pattern. It occurs at

the extreme eastern end of the range of C. weinlaruli.

Remarks: Tl le original images of this species in 2012

were unfortunately vertically foreshortened by the

printer. No material was available for phylogenetic study.

Etymology: Lynx rufus, the American bobcat or lynx; the

shell has a similar color pattern. Used as a noun in

apposition.

Chondropomium marmoreum (Watters and Duflv, 2010)

(Ligures 3, 4, .'34—42, 184)

Chresonymy

Chondropoma ( Chondropoma ) marmoreum Watters and

Duffy, 2010: 6-7, figs. 16-19.

Chondropoma marmoreum Watters and Duffy, 2010:

Watters, 2012: 9, 14, figs. 51, 52, 55 [habitat], 56

[habitat].

Description: Shell medium for family (largest adult

specimen 18.3 mm maximum length, decollate, including

peristome; smallest adult specimen 14.0 mm length, de¬

collate. including peristome), solid, conic, umbilicus minute.

Protoconch whorls unknown, decollate in all specimens ex-

amined. Teleoconch of 4.25 whorls, final whorl detached

immediately before lip. Peristome double, tear drop-shaped.

Outer lip thin, narrow to moderately expanded, minute

posterior auricle present, lip detached from previous whorl.

Inner lip short, mostly fused with outer lip. Spiral sculpture of

numerous (ca. 36 on final whorl) low threads or cords, be¬

coming stronger and more widely separated towards umbi¬

licus; 3-6 very' feeble cords within umbilicus. Axial sculpture

of similar sized threads. Intersections of sculpture forming

minutely beaded surface. Suture minutely channeled. Axial

threads forming minute cusps at suture. Background color

tan, reddish brown, or grey, with dark brown ' IT-shaped

spots arranged in spiral bands, often aligned into axial stripes

as well; lip white. Operculum paueispiral with thin calcareous

deposit. Radula as in genus but with strong hook-shaped

denticle on inner side. Animal unknown.

Type Material: Holotype: UL 420735; Paratypes:

OSUM 32483(1); OSUM 32482(1); BMNH 1996347(1).

Type Locality: “Dominican Republic, Barahona Penin¬

sula, Pedernales Province, along Route 44 ca. 10 km SE of

Pedernales.

Type Figured: Watters and Duffy, 2006, figs. 16, 17.

Distribution: Low hills off the western Sierra Baoruco

from Pedernales down the coast to Cabo Rojo and

southeast almost to Oviedo.

Habitat: Under limestone debris in xeric areas with cacti,

often near the coast. Locally abundant.

Other Material Examined (688 Specimens): Do¬

minican Republic. Pedernales Province. UL 216776

(10), 5 km NW of Oviedo; UL 216774(26), 8 km NW of

Oviedo; UL 216419(9), 8 km SW of Las Mercedes; UL

216391(9), 6 km SW of Las Mercedes; UL 216460(66),

15 km SE of Pedernales; UL 216465(52), 20 m, 17 km

SE of Las Mercedes; UL 216459(74), 26 km NW of

Oviedo; GTW 717 0a( 1 ). 22.5 km W of Oviedo; GTW

717 0b ( 1 ) , Cabo Falso; UL 216510(32), Cabo Rojo; UL

216458(43), 13 km N of Cabo Rojo; GTW 7170c(15), 3.4

m, just N of Cabo Rojo airport; UF 216499(12), 6 km

SSL of Cabo Rojo; GTW 7170d(3), along RD 44, next to

large limestone outcrops, 16 km E of Cabo Rojo; UF

216388(150), UF 249174(19), 3 km SE of Cabo Rojo; UF

216457(69), UF 216514(25), 8 km SE of Pedernales;

GTW 7170e(3). 10 km SE of Pedernales off RD 44; UF

G.T. Watters and P. Larson, 2017

Page 173

Figures 34-48. Chondropomium species. 34—42. Chondropomium mamwreum (Watters and Duffy, 2010). 34, 35. UF 420735,

holotype, 18.3 nun. 36. UF 216459, 17.7 mm. 37. UF 216776, 15.4 mm. 38. BMNH 1996347, paratype, 18.3 mm. 39. OSUM 32483,

paratype, 15.4 mm. 40. Habitat (photo A. Gettleman). 41. Individuals in situ under rock (photo A. Gettleman). 42. GTW 771 70d,

radula. Scale bar = 100 pan. 43 — 48. Chondropomium sardonyx new species. 43, 44. UF 216454, holotype, 19.3 mm. 45. UF 505805,

paratype 1, 19.1 mm. 46. UF 505805, paratype 2, 17.3 mm. 47. UF 505805, paratype 5, 15.8 mm. 48. UF 467518, 20.1 mm.

216465(17), UF 216512(20), UF 257730(44), 17 km SE

of Pedernales; UF 216501(29), 22 km SE of Pedernales;

GTW 71701(3), 15-16 km E of Cabo Rojo off RD 44;

GTW 7170g(l), 0.7-1. 1 km along 5 km rough road that

runs from 1 .9 km S of old naval base to La Cueva, Cabo

Rojo.

Variation Among Specimens: The' dark brown color

pattern varies from a few axially aligned triangles to more

extensive mottling.

Comparison Witli Other Species: This species has

a color pattern similar to some specimens of C. blaineontm

(with which it may be sympatric) and C. weinlandi. It

differs from all other species in the genus by possessing

distinct spiral sculpture on at least the early whorls,

which may render the surface beaded or even minutely

serrate. It is also similar to Chonclropoma eyerdami

Bartseh, 1946, from theTiburon Peninsula of Haiti, and

Chondropoma brownianum Weinland, 1880, from lie

de la Gonave, Haiti. It differs from both in having

weaker sculpture.

Remarks: This species is placed in Chondropomium with

some reservation. The finely beaded sculpture is more

similar to Chondropoma, but the shell form and coloration

are more like Chondropomium. The radula differs from

other Chondropomium studied in having a prominent

Page 174

THE NAUTILUS, Vol. 131, No. 3

hook-shaped denticle on the inner side of the inner

marginal tooth; this is lacking in other species covered

here. Along with C. eyerdami and C. brownianum, it may

prove to he a member of an undescribed genus apart from

either Chondropoma or Chondropomium. Unfortunately,

no suitable genetic material was available for study for any

of these taxa.

Etymology: L. marmoreus , marbled.

Chondropomium sardonyx new species

(Figures 43-48, 182)

Description: Shell medium for family (largest adult

specimen 20.1 mm maximum length, decollate, in¬

cluding peristome; smallest adult specimen 15.0 mm

length, decollate, including peristome), translucent but

solid, stout, elongate conic, polished. Protoconch un¬

known; all specimens seen decollate. Teleoeonch of

3.75-4 whorls. Axial sculpture of final whorl of nu¬

merous (ca. 120) very fine, closely spaced, low, weak

threads, often obscure. Spiral sculpture absent except

forea. 15 cords within umbilicus. Overall sculpture very

weak, with polished aspect. Suture narrowly channeled.

Each axial thread slightly enlarged into bead at suture,

rendering suture minutely serrate. Aperture teardrop-

shaped, lip single, not or very slightly expanded,

posteriorly angulate but lacking auricle. Lip narrowly

detached from previous whorl for ca. 14 turn, angular

posterior of lip forming sharp crest. Color pattern of

final whorl reddish-orange with white umbilicus; pre¬

vious whorls tan posteriorly and dark purple anteriorly.

Lip white inside and out. Sutural beads white, con¬

trasting with purple of previous whorl and orange of

final whorl. Operculum paueispiral with thin calcareous

deposit. Animal and radula unknown.

Type Material: Holotype: UF 216454; Paratype 1: UF

505805; Paratype 2: UF 505805; Paratype 3: UF 505805;

Paratype 4: UF 505805; Paratype 5: UF 505805.

Type Locality: 20 m elevation, 3 km ESE of La Canoa,

Barahona Province, Dominican Republic.

Distribution: Known only from the western foothills of

the Sierra Martin Garcia, an isolated outlier of the Sierra

Neiba.

Habitat: Specimens were found in a xerie arroyo under

agave. Locally common.

Other Material Examined (43 Specimens): Domini¬

can Republic. Barahona Province. UF 505806(36),

from the type locality; UF 467518(5), 1 .6 km W of Fondo

Negro; UF 216822(2), 200 m, 12 km NF of La Canoa.

Variation Among Specimens: Specimens are very

uniform in most characteristics, but differ in the degree of

coloration.

Comparison With Other Species: The overall colora¬

tion of this rarely seen species is unique in the genus.

However, the color pattern is very similar to that found in

some Colonina, but C. sardonyx lacks the prominent

sutural tufts of those species.

Remarks: This species is sympatric in its very narrow

range with the' color forms azuense and barahonense of C.

weinlandi. It is known only from dead specimens.

Etymology: Named for the form of the mineral onyx

having red parallel bands: sardonyx; a noun in apposition.

Chondropomium weinlandi (Pfeiffer, 1862)

(Figures 49-73, 185)

Chresonymy

“ weinlandi form”

Cyclostoma semilabre “Lamarck" of Pfeiffer, 1850: 80

[misidentification]; Pfeiffer, 1854b: 271, pi. 37, figs. 1,

2; Pfeiffer, 1854d: pi. 49, fig. 17; Crosse, 1891: 172;

Bartsch, 1946: 27 [in synonymy of Chondropoma

weinlandi Pfeiffer, 1862] [non Lamarck, 1822].

Chondropoma semilabre “Lamarck” of Pfeiffer, 1851: 1 73

[misidentification]; Pfeiffer, 1852a: 286; Adams and

Adams, 1858: 295, pi. 86, figs. 1 la, b; Pfeiffer, 1858b:

139; Reeve, 1863: text to pi. 1, figs. 2a, b [in synonymy

of Chondropoma weinlandi Pfeiffer, 1862]; Bartsch,

1946: 27 [in synonymy of Chondropoma weinlandi

Pfeiffer, 1862] [non Lamarck, 1822],

Chondropoma weinlandi Pfeiffer, 1862: 96-97; Reeve,

1863: text to pi. 1, fig. 2a; Pfeiffer, 1865: 149; Pfeiffer,

1876: 193; Kobelt, 1880: 277; Weinland, 1880:

342-343, 346, 347; Crosse, 1891 167, 172-173; Clench

and Aguayo, 1948: 54.

Chondropoma ( Chondropomium ) weinlandi Pfeiffer,

1862: Henderson and Bartsch, 1920: 59-60; Pilsbry,

1933: 126, 129; Wenz, 1939: 542, fig. 1441.

Chondropoma j Chondropomium] weinlandi Pfeiffer,

1862: Thiele, 1929: 131.

Chondropoma (Chondropomium) swiftii swiftii “Shut-

tleworth” Bartsch, 1946: 24-26, 28, pi. 4. figs. 1, 2.

Chondropoma ( Chondropomium ) swiftii weinlandi

Pfeiffer, 1862: Bartsch, 1946: 27-28. pi. 4, fig. 5.

Chondropoma semilabris "Lamarck" of Pfeiffer, 1850:

Mermod, 1952: 42-44, fig. 109.

Chondropomium swiftii “Shuttleworth” of Bartsch. 1946:

Watters, 2006: 63, 498-499; Watters and Duffy, 2010:

7; Watters, 2012: 11.

Chondropomium siviftii weinlandi (Pfeiffer, 1862):

Watters, 2006: 63, 547-548; Watters, 2012: 9, fig. 42.

“ azuense form”

Cyclostoma semilabre “Lamarck” of Pfeiffer, 1854d: pi.

49, fig. 20 [misidentification, non Lamarck, 1822].

Chondropoma weinlandi Pfeiffer, 1862: 96-97; Reeve,

1863: text to pi. 1, fig. 2b.

Chondropoma (Chondropomium) swiftii azuense Bartsch,

1946: 26, 29, pi. 4, fig. 6.

Chondropomium swiftii azuense (Bartsch, 1946): Watters,

2006: 63, 499.

G.T. Watters and P. Larson, 2017

Page 175

Figures 49-73. Chondropomium weinlandi (Pleiller, 1862). 49-61 . “ weinlandi form”. 49. Cyclostoma semilabre “Lamarck” Pfeiffer,

1854b, pi. 37, figs. 1, 2. 50. Cyclostoma semilabre “Lamarck” Pfeiffer, 1854d: pi. 49. fig. 17. 51. UF 45658, 20.4 mm. 52. UF 45651, 19.0

mm. 53. UF 45684, 19.0 mm. 54. UF 216811, 15.5 mm. 55, 56. OSUM 42378. 1 2.0 mm. 57. OSUM 4236 1 . 18.8 mm. 58. GTW 7087e.

17.5 mm. 59. GTW 7087c, 20.7 mm. 60. Living individual. 61. Habitat. 62-67. “ barahonense form”. 62. Cyclostoma semilabre

“Lamarck” Pfeiffer, 1854d: pi. 49, figs. 18. 63. Chondropoma { Chondropomium ) swiftii barahonense Bartsch, 1946. USNM 354956,

holotype, 18.9 mm. 64. Chondropoma {Chondropomium) swiftii saturation Bartsch, 1946. USNM 354953, holotype, 16.6 mm. 65. UF

45651, 16.5 mm. 66. UF 45651, 19.2 mm. 67. OSUM 42362, radula. Scale bar = 200 (am. 68-73. “azuense form”. 68. Cyclostoma

semilabre “Lamarck” Pfeiffer, pi. 49, fig. 20. 69, 70. Chondropoma {Chondropomium) swiftii azuense Bartsch, 1946. USNM 493298,

holotype, 19.0 mm. 71, 72. UP' 216834, 18.3 mm. 73. Living individual.

Page 176

THE NAUTILUS, Vol. 131, No. 3

“/?a ra h o nense/sa turatum form ”

Cyclostoma semilabre “Lamarck” of Pfeiffer, 1854d:

Pfeiffer, lS54d: pi. 49, ffg. IS [misidentification, non

Lamarck, 1822].

Chondropoma ( Chondropomium ) swift ii barahonense

Bartsch, 1946: 26-27, pi. 4, fig. 3.

Chondropomium swift ii barahonense (Bartsch, 1946):

Watters, 2006: 63, 499; Watters, 2012: 11.

Chondropoma (Chondropomium) swiftii saturation

Bartsch, 1946: 26, pi. 4, fig. 4.

Chondropomium swiftii saturation (Bartsch, 1946):

Watters, 2006: 63, 499; Watters, 2012: 11

Description: Shell medium for family (largest adult

specimen 25.5 mm maximum length, decollate, including

peristome; smallest adult specimen 14. 1 mm length, de¬

collate, including peristome), solid, polished, almost trans¬

lucent, conic to fairly high-spired, umbilicus narrow,

compressed, spire Vi- 1/3 total length. Protoconch of 1.5

minute, smooth, pale tan whorls, decollated in adults. De¬

collate specimens have jagged, irregular break from earlier

whorls, which seems to involve more than just protoconch

whorls. Teleoconch of 3.75—4.5 whorls, final whorl narrowly

detached for last l/4th of whorl. Suture narrow, channeled.

Peristome single, tear drop-shaped. Lip barely expanded,

narrowest facing umbilicus, with minute triangular posterior

auricle, narrowly detached from previous whorl, forming

sharp posterior keel. Spiral sculpture absent except for 8-17

narrow, fine cords within umbilicus. Axial sculpture of

flattened, low ribs, separated by incised grooves, often

grouped in growth series, ca. 130 ribs on final whorl, each

forming minute bead at suture; no fused tufts present. Color

pattern very variable; see Remarks below. Operculum

paucispiral with thin calcareous deposit. Radula as in genus.

Type Material: Chondropoma weinlandi Pfeiffer, 1862:

three specimens from NHMUK, unnumbered, from

Cuming collection from “Haiti L, none bear an “X,” but

may be tbe syntvpe lot. However, (. Ablett (pers. comm.,

2016, NHMUK) does not believe that any of these are

Pfeiffer’s material as they lack Pfeiffer’s handwriting and

the label does not specifiy that they were collected by

Salle. Pfeiffer originally identified his specimens as

Cyclostoma semilabre “Lamarck" (see Remarks below)

but these specimens have not been found at NHMUK.

Three other specimens glued on a board at NHMUK, also

unnumbered, from Cuming collection labeled "weinlandi

VAR may be syntypes of Weinland’s variety major or

variety malleata but this cannot be determined. A spec¬

imen at ZMB of C. weinlandi (65684) is from Pfeiffer but

is labeled “kleiner ills typus” (smaller than the type) on

a second label; a second specimen labeled C. weinlandi

(ZMB unnumbered) is from Paetel rather than Salle;

neither can be identified as type material (C. Zorn, pers.

comm., 2016, ZMB), although apparently a type existed at

some point. Chondropoma ( Chondropomium ) swiftii

azuense Bartsch, 1946: Holotype: USNM 493298.

Chondropoma ( Chondropomium ) swiftii barahonense

Bartsch, 1946: Holotype: USNM 354956. Chondropoma

(Chondropomium) swiftii saturation Bartsch, 1946: Ho¬

lotype: USNM 493299; Paratype: USNM 354953.

Type Locality: Chondropoma weinlandi Pfeiffer, 1862:

“Haiti.” Chondropoma (Chondropomium) swiftii azuense

Bartsch. 1946: unknown, but collected by Salle, who only

collected in the Dominican Republic portion of Hispaniola.

Bartsch (1946) listed Azua, Dominican Republic, fora non-

tvpe specimen. Chondropoma (Chondropomium) swiftii

barahonense Bartsch, 1946: “Barahona.” Chondropoma

( Chondropomium ) swiftii saturation Bartsch, 1946:

unknown.

Type Figured: Chondropoma weinlandi Pfeiffer, 1862:

Pfeiffer (1862), in a list of misidentifieations of C. semi-

lahris that he subsequently reidentified as C. weinlandi ,

gave Pfeiffer, 1854b, pi. 37, figs. 1, 2 and 1854d, pi. 49,

figs. 17-20, as the only figured specimens, but see Re¬

marks. No one figure can be identified with Pfeiffer's

description. Chondropoma ( Chondropomium ) swiftii

azuense Bartsch, 1946: Bartsch, 1946, pi. 4, fig. 6.

Chondropoma (Chondropomium) swiftii barahonense

Bartsch, 1946: Bartsch, 1946, pi. 4, fig. 3. Chondropoma

(Chondropomium) swiftii saturation Bartsch, 1946:

Bartsch, 1946, pi. 4, fig. 4.

Distribution: Rift valley of Hoya de Enriquillo/Plain du

Cul-de-Sae from the western shore of Etang Saumatre,

around Lago Enriquillo, east to Agua.

Habitat: Generally below 200 m under fossilized coral

rubble and dead agave and grasses in xeric areas. Locally

common.

Other Material Examined:

“ weinlandi form” (566 Specimens): Dominican Re¬

public. Azua Province. UF 216837(3), 3 km W of

Ilatillo; UF 45565(7), UE 45857(1), 12 km ESE of Las

Charcas; UF 216805(1), UE 216833(1), 50 m, 8 km ESE

of Las Charcas; UF 23180(7), 80 m, 2 km N of Las

Charcas; UF 23162(1 1), 80 m, 2 km N of Las Charcas; UF

216834(62), 1 km N of Estebama; UF 216830(83), 2 km

SE of Azua; UF 216836(10), 3.5 km SSE of Azua; UF

216840(1), 25 km W of Azua; UE 45853(22), 70 m, 7 km E

of Azua; UF 216839(1). 160 m, 15 km SW of Las Yayitas;

UF 216827(9), 12 km E of Quita Coraza; OSUM 42361

(32), 80 m, off road to Galindo Adrento, 2.8 km ESE of

Galindo Adrento, 5.7 km VVSW of Isura, 18.4062° N,

-70.8909° W; UF 216809(16), 2 km NW of Cortes.

Peravia Province. UF 216811(10), 20 m, 5 km N of

Cruee de Ocoa. Barahona Province. UF 45584(4),

27 km W of Azua; UF 216835(7), 140 m, 14 km NE of La

Canoa; GTW 7087b(2), Fondo Negro; UF 216824(25),

13 km ESE of La Canoa; UF 216828(6), Cerro de Sal,

2 km SW of Las Salinas; UF 216831(13), 9 km W of

Cabral; UE 45644(9), SO m, 1 km E of Las Salinas; GTW

7087c(l), OSUM 42365(13), 37 m, top of small bill, off

road from Puerto Alejandro to La Canoa, 3.4 km NE of

Bombita, 18.3482° N, -71.1225° W. Independencia

Province. UF 45658(18), SO m, 1 km WSW of Los

G.T. Watters and P. Larson, 2017

Page 177

Saladillos; GTW 7Q87a(l), Mella; UF 216832(4), UF

249181(13); 1 km S of Mella; GTW 7087s(19), 42 m, along

road to Angostura, 1 km NNW of Angostura, 18.3412° N,

-71.4207° W; UF 45651(14), UF 45783(3), 60 m, 2 km

NW of Angostura; UF 45684(13), 50 m, 3 km NW of

Angostura; UF 216844(2), 250 m, 4 km SSE of Los Pinos

del Eden; UF 216829(4), 2 km WNW of Postrer Rfo; UF

216826b(18), 7 km E of Las Descubierta; UF 45793(13),

17 km ESE of El Limon; UF 216848(26), 14 km WNW of

Duverge; GTW 7087g(3), ridge NW of Jimanf, Inde-

pendeneia Province; GTW 7087d(5), S side of highway,

a few km E of Jimanf, Independence Province; OSUM

42379(7), Bartolome, Independeneia Province; OSUM

42377(2), 27 m, off RD 46, Independeneia Province.

18.4167° N, -71.7075° W; OSUM 42378(31), 10 m, off RD

48, Independeneia Province. 18.5204° N, -71.7614° W.

“azuense form” (194 Specimens): Dominican Re¬

public. Azua Province. GTW7 7087k(l), 191 m, top of

small hill, off RD 2, 1.86 km ENE of Agua, 1.6 km W of

Las Carreras, 18,3586° N, -70.5016° W; UF 45565(1),

12 km ESE of Las Charcas; UF 216837(2), 3 km W of

Hatillo; UF 216823(6), 7 km SEofHatillo; UF 216809(4),

2 km NW of Cortes; UF 216462(12), 3 km ESE of Cortes;

UF 23180(2), 80 m, 2 km N of Las Charcas; UF 216805

(4), 8 km SE of Las Charcas; UF 216834(20). 1 km N of

Estebanfa; UF 216866(7), 3 km SE of Azua; UF 216836

(4) , 3,5 km SSE of Azua. Barahona Province. GTW

70S7p(4), 20 m, off road from Puerto Alejandro to La

Canoa, 2.8 km NE of Bombita, 18.3478° N, -71. 1298° W;

OSUM 42364(5), 37 m, top of small hill, off road from

Puerto Alejandro to La Canoa, 3.4 km NE of Bombita,

18.3482° N, -71.1225° W; UF 216846(27), 2 km NE of La

Canoa; UF 216821(6), 3 km NE of La Canoa; UF 216454

(5) , UF 216847(1), 3 km ESE of La Canoa; UF 216822

(15), 20 m, 12 km NE of La Canoa; UF 216831(1), 9 km

W of Cabral; UF 45644(1), I km E of Las Salinas.

Independeneia Province. GTW 7087r(4), 42 m,

quarried area along road to Angostura, 1 km NNW of

Angostura, 18.3412° N, -71.4207° W; UF 45684(7), 60 m,

2 km NW of Angostura; UF 45684(7), 3 km NW of

Angostura; GTW 7087w(5), semi-cleared pasture with

piles of rocks, off RD 48, 18,5204° N, -71.7614° W; UF

45793(2), 17 km ESE of El Limon; UF 216832(4), I km S

of Mella; OSUM 42359(19), Bartolome; UF 216829(4),

2 km WNW of Postrer Rfo; GTW 7087f(15), Los Rios, N

shore Lake Enriqnillo.

“barahonense/saturatum form” (421 Specimens):

Dominican Republic. Azua Province. UF 216823(20),

7 km SE of Hatillo; UF 216462(28), 3 km ESE of Cortes;

UF 216809(3), 2 km NW of Cortes; GTW 7087m(l), 191

m, top of small hill, off RD 2, 1.86 km ENE of Agua,

1.6 km W of Las Carreras, 18.3586° N, -70.5016° W; UF

45565(34), UF 45857(11), 12 km ESE of Las Charcas; UF

216805(4), UF 216833(7), 50 m, 8 km SE of Las Charcas;

UF 216834a(9), 1 km N of Estebania; UF 216827(9),

Monte Rio; UF 216836(9), 3,5 km SSE of Azua; UF

216830(1), 12 km E of Quita Coraza. Barahona

Province. UF 45589(2), 270 m, 9 km NE of La Canoa;

UF 216838(3), 170 m, 9 km NE of Fondo Negro; UF

467518(6), 1 .6 km W of Fondo Negro; GTW 7087x(5), 20

m, off road from Puerto Alejandro to La Canoa, 2.8 km

NE of Bombita, 18.3478° N,'-71 1298° W; OSUM 42363

(3), 37 m, top of small hill, off road from Puerto Alejandro

to La Canoa, 3.4 km NE of Bombita, 18,3482° N,

-71.1225° W; OSUM 42362(20), 20 m, off road from

Puerto Alejandro to La Canoa, 2.8 km NE of Bombita,

18.3478° N, -71.1298° W; UF 216846(9), 2 km NE of La

Canoa; UF 216821(11), 3 km NE of La Canoa; UF

216847(51), 3 km ESE of La Canoa; UF 216824(5), 13 km

ESE of La Canoa; UF 216822(68), 12 km NE of La

Canoa; UF 216831(13), 9 km W of Cabral; UF 45644(2),

1 km E of Las Salinas; UF 45633(3), 3 km E of Las Salinas.

Independeneia Province. UF 45651(7), UF 45684(3),

60 m, 2 km NW of Angostura; UF 45684(3), 3 km NW of

Angostura; UF 45661(3), 5 km W of Los Saladillos; UF

21 6832(31 ), 1 km S of Mella; UF 45658(8), SO m, I km

SSW of Los Salidillos; UF 216829(8), 2 km WNW of

Postrer Rfo; UF 216817(1), 150 m, 6 km NW of Postrer

Rfo; UF 216842(2), 10 m, 4 km E of La Descubierta; UF

216843(3), 100 m, 3 km NNE of La Descubierta.

Variation Among Specimens: See under “Remarks”

below.

Comparison With Other Species: The unexpanded lip

differentiates this species from S. superlmm and S.

asymmetricum. Chondropomium bealense , C. blaineo-

rum, and C. lynx are similar but occur on the opposite

slopes of the Sierra Baorueo. In addition, C. blaineorum

and C. lynx have color patterns not seen in the nearly 1 100

specimens of C. weinlancli examined. Chondropomium

blaineorum has a peculiar mosaic pattern whereas C lynx

has smeared individual spots. Some specimens of C.

beatense in particular resemble the barahonense form of

C. weinlandi (but is geographically the farthest away from

that species); they have a shorter, more conic, less inflated

shell than does C. weinlandi. Chondropomium sardonyx

is sympatric with most of the C. weinlandi color forms but

the striking orange and black two-tone color pattern is

unique to it.

Dead specimens of Crossepoma vermiculatum Bartseh,

1946 lacking an operculum are very similar to the "azuense

form” and are sympatric. That species has a unique un¬

derlying pattern of vertical grey markings and regular

sutural spots not found in the "azuense form”. Crossepoma

vermiculatum has a pseudolamella.

Remarks: Three groups of similar shells may be recog¬

nized based on color pattern. Bartseh (1946), as “Chon-

dropoma swiftii,’’ described them as five subspecies.

(However, as explained below, this species is not

Cyclostoma swiftii Shuttleworth, 1854.) Pfeiffer (1862)

had referred to his 1854b, pi. 37, figs. I and 2 and his

1854d, pi. 49, figs. 17-20 all as variants of C. weinlandi.

Bartseh (1946) separated Pfeiffer’s pi. 49, fig. 19 as an

Page 178

THE NAUTILUS, Vol. 131, No. 3

example of C. vemiiculatum Bartsch, 1946, relegating his

remaining figures to various subspecies of C. “swiftii.

Bartsch s breakdown was:

1) “ swiftii swift ii": Narrow interrupted spiral brown

bands on a “buff’ background (Figure 59). This is the least

common of the forms.

2) “swiftii weinlancJi ”: Narrow interrupted spiral brown

bands on a “flesh-colored’’ background (Figures 51-58).

The differences in background color do not seem sig¬

nificant and th e swifti and the ” weinlanrii forms are here

considered the same. This is the most common variant.

3) “swiftii saturation" : Broad, more or less continuous

brown bands, which may appear as adjacent blotches

(Figure 65).

4) “ swiftii barahonense Narrower continuous brown

bands. Based on the material studied here, the saturation

and barahonense morphs cannot be differentiated (Fig¬

ures 63, 64, 66).

5) “ swiftii azuense": Bands absent or nearly so, back¬

ground white (Figures 69-72).

Bartsch, working with a very small sample size, saw not

only differences in color but differences in ranges, with

the subspecies being allopatrieally distributed. However,

subsequent, more intensive collecting clearly demon¬

strates that these forms are not allopatric. All generally

overlap (Figure 185) anti are widely distributed in the rift

valley of Hova de Enriquillo/Plain du Cul-de-Sac where all

three forms may be found under the same rock. Although

the three forms cannot be subspecies, they are (usually)

separable. What is their relationship to each other and are

they distinct species or a polymorphic species for color?

In our phylogenetic analysis all three morphs formed

a single monophvletic clade. I Iowever, the color forms did

not segregate together (Figure 5). Neither w'as there any

apparent geographic pattern. We conclude that C.

weinlandi does not consist of phylogenetically separable

units based on color forms but rather represents co¬

occurring polymorphic morphotypes.

Differences in shell color may extend to animal color as

well. In the “weinlandi color form” the animal is pale tan

with grey mottling between the eyes and on the snout and

the tentacles. In the “azuense color form the entire

animal, including the tentacles, is a uniform pale tan,

slightly darker on the foot, lacking the grey coloration; this

might be considered a leucistic phenotype. The animal of

the “barahonense color form” has not been recorded.

Tl le taxonomic history of C. weinlandi is unusually

complicated, beginning with Lamarck, entwining the

taxonomy of several nominal species. In 1822, Lamarck

described but did not illustrate Cyclostonm semilabris

from a unknown locality; neither did he refer to any

published figure. He described its sculpture as “minu-

tissime cancellated Pfeiffer (1862) proposed that it

originated from the Bahamas. Bartsch (1946) did not

concur with Pfeiffer and identified Lamarck’s species as

a Hispaniolan taxon. In 1952, Mermod figured (his figure

109) the tvpe of Cyclostoma semilabris in the Museum

d histoire naturelle de Geneve (unnumbered). This

specimen proved to be different from Bartseh’s

Hispaniolan species but very similar to several Bahamian/

Caicos taxa, especially Chondropoma hjalmarsoni Pfeiffer,

1858, from Turks and Caicos. This left Bartsch’s Haitian

species without a valid name, which prompted it to be de¬

scribed as Chondropoma bellevittatum Watters, 2016a.

Shuttleworth (1854) described but did not illustrate

Cyclostoma swiftii. He stated that it was collected from

Ponce, southern Puerto Rico, by Swift and had been in the

collection of Bland. Indeed, Dali and Simpson (1901) and

Baker (1941) listed this species from Puerto Rico and

w'ere followed in this identification by van der Sehalie

(1948), who illustrated a specimen as C. swiftii from

Ponce. However, that shell bears no resemblance to the

species complex discussed here from Hispaniola. This

Ponce taxon was later described as Chondropoma schaliei

Baker, 1950; Baker did not comment on why he discarded

C. swiftii in favor of the new species name. Shuttleworth,

in the description of C. swiftii , mentioned “obtuse

decussate ,” a sculpture not found in the Hispaniolan

species complex discussed here. He compared C. swiftii

to his Puerto Rican Cyclostoma blauneri Shuttleworth,

1854, also having decussate sculpture.

Apparently based solely on Lamarck’s brief description

of his "jolie coquille,” Sowerby (1843: pi. 24, fig. 60) il¬

lustrated a specimen supposedly from Cuba as Lamarck’s

C. semilabris. Like Shuttleworth’s description of C. swiftii

and Lamarck’s description of C. semilabris , Sowerby’s

illustration and description also indicated a cancellate

sculpture. The Hispaniolan species complex described

here lacks such sculpture and that complex would appear

to be neither C. swiftii nor C. semilabris.

Later, Pfeiffer (1852a) redescribed his “Chondropoma

semilabre Lamarck,” giving Haiti as the provenance rather

than the Bahamas, Puerto Rico, or Cuba. He added that

Salle had collected the specimen(s). As Salle only col¬

lected in the Dominican Republic portion of Hispaniola,

Pfeiffer’s use of “Haiti” referred to the entire island.

However, Pfeiffer described the species (p. 286) as having

“lineis spiralibus obsoletissimis interdum decussatula ,”

which, again, is a sculpture lacking in the Hispaniolan taxa.

But in 1862 he retracted this line of description as in¬

applicable stating that he had copied it. apparently in¬

advertently, from Lamarck s original description. Clearly,

whatever their valid name, by this time Pfeiffer was

working with specimens in the Hispaniolan species

complex described here.

In 1862, Pfeiffer had again changed his mind con¬

cerning the identification of C. semilabris. In 1854 Pfeiffer

had figured his Hispaniolan specimens of “C. semilabris ’

(1854b, pi. 37, figs. 1 and 2 and 1854 cl, pi. 49, figs. 17-20).

These shells undoubtedly belong to the species complex

recognized here and not to any Bahamian, Cuban, or

Puerto Rican species. By 1862, Pfeiffer had come to

believe they were not Lamarck’s C. semilabris , which

Pfeiffer now believed was a Bahamian species. Neither

did he include C. swiftii in his deliberations, apparently

dismissed as a Puerto Rican taxon and not under con¬

sideration for this Hispaniolan species. In his mind this

left the Hispaniolan species without a valid name. He

G.T. Watters and P. Larson, 2017

Page 179

named it Chondropoma weinlandi Pfeiffer, 1862, from

“Haiti” referring back to his 1854b and 1854d figures of

“C. semilabris .” Reeve (1863c) figured it as well under C.

weinlandi.

The description of decussate sculpture in the original

description of C. swiftii, and the subsequent mention of

such sculpture by Sowerby, suggest that Shuttleworth’s

species is not the Hispaniolan species, i.e., the “C. swiftii ”

of Bartsch. Mermod (1952) mentioned, but did not il¬

lustrate, the type of C. swiftii in the Naturhistorische

Museum, Bern. He compared it with presumed speci¬

mens of C. semilabris and pronounced that they were the

same species. Swift himself either collected or employed

someone else to collect in Puerto Rico (Clench, 1938), but

is not listed by Bartsch (1946) as having collected in

Hispaniola. How then did C. swiftii come to be associated

with Hispaniola?

The key point occurred when Bartsch (1946) figured

the stated type of C. swiftii , which matched the Hispa¬

niolan species complex studied here, but lacked the de¬

cussate sculpture described by Shuttleworth. Bartsch

recognized C. weinlandi as a subspecies of C. swiftii

differing only in the background color of the shell. He

further separated Pfeiffer’s 1854b and 1854d figures into

several subspecies of swiftii and renamed the 1854d figure

19 as Chondropoma vermicidatnm Bartsch, 1946. This

might have ended the confusion surrounding the identity

of C. swiftii , although leaving some nagging questions

unanswered concerning sculpture and the fact that Swift

may never have collected in Hispaniola.

In preparation of this study, we contacted E. Neubert at

the Naturhistorische Museum, Bern, for a digital pho¬

tograph of the type of C. swiftii. The photograph of the

specimen sent to us is not only not the same specimen as

figured by Bartsch but represents a different species

entirely: Chondropoma pictum Pfeiffer, 1839, from Cuba.

This same specimen was also illustrated as one of the

syntypes (NMBE 19109/4) for C. swiftii by Neubert and

Gosteli (2005: pi. 15, fig. 1). It should be noted that C.

pictum has predominantly spiral threads with irregular

axial growth marks that could be construed as decussate

sculpture. Specimens may also be colored as described by

Shuttleworth for C. swiftii , particularly the subspecies or

“color form" C. celsum Torre and Bartsch, 1938.

Shuttleworth indicated that his specimen of C. swiftii had

been sent to him by Bland. Bland possessed a large

collection of land snails from the Caribbean, including

Cuba, and it is possible that Bland had inadvertantly

switched a label from Cuba for that of Puerto Rico. But it

is unclear if Swift ever collected in Cuba.

Several explanations are possible. Perhaps the speci¬

men shown to us was part of a syntype lot that may consist

of mixed species, of which a different specimen and

species was the one figured by Bartsch. However, E.

Neubert (pers. comm., 2015), upon examination stated

that none of the specimens match that shown by Bartsch

and all appear to be C. pictum. Was it possible that

Bartsch never returned the specimen? Bartsch stated

(1946: 28): “Through the kindness of the Naturhistorische

Museum at Berne, Switzerland, I have been able to ex¬

amine and figure the type." But did Bartsch have the

specimen in hand? His figure of it, plate 4, figure 2, seems

of a different quality than all of his other figures: it is

slightly out of focus, appears washed out, and is oriented

slightly differently. It appears to be a photograph of

a photograph, suggesting he did not have the specimen in

hand but only a photograph supplied bv the Bern mu¬

seum. Furthermore, Mermod (1952) said "I was able,

thanks to Professor Baumann, the Museum of Bern, to

compare swifti types of Shuttleworth and his named

semilabris examples” [from the French] indicating that

the types were back in Bern by 1952, although perhaps

still missing Bartsch s possible loan. None of these sce¬

narios would seem to explain the discrepancy.

In summary: 1) the specimen illustrated as the type of

C. swiftii by Bartsch does not match any of the existing

syntypes; and 2) Shuttleworth’s description of C. swiftii

does not match the Hispaniolan species of Bartsch. The

question of where Bartsch s specimen originated (and

where it went) remains a mystery. But based on the ev¬

idence presented here C. swiftii is not the Hispaniolan

species and the earliest available name is Chondropoma

weinlandi Pfeiffer, 1862. The existing syntypes of C.

swiftii are specimens of the Cuban Cyclostonm pictum ,

and C. swiftii is a junior synonym of that name. However,

the types of C. swiftii are verv similar to C. pictum celsum

Torre and Bartsch, 1938, and may represent an earlier

name for that subspecies.

Weinland (1880) listed two varieties, both unfigured

and undescribed, from “St. Domingo:” Chondropoma

weinland var. major (p. 346) and Chondropoma weinlandi

var. malleata (pp. 343, 346). They are nomina dubia.

Original Description: Chondropoma weinlandi Pfeiffer,

1862. Pfeiffer’s 1852a description of “Ch. semilabre

Lamarck” appears to be the earliest description of

C. weinlandi, although apparently conflating the two

species. From the Latin. “Shell subumbilieate, ovate-

elongate, truncate, somewhat solid, longitudinally pli¬

cate, very obsolete spiral lines sometimes decussate, shiny,

diaphanous, white, ornated with interrupted lines perhaps

narrow and numerous brown bands; suture very minutely

crenulated; 4 remaining whorls a little convex, the front of

the last a little solute, the top carinate, base concentrically

striate; aperture vertical, obliquely aeuminate-ovale;

peristome obtuse, everywhere narrowly expanded.

Operculum white.” In 1862, he removed the phrase “very

obsolete spiral lines sometimes decussate" from the de¬

scription, which he had based on Lamarck’s description.

Etymology: Chondropoma weinlandi Pfeiffer, 1862:

Cristoph David Frederick Weinland (1829-1914), German/

American conchologist, physician. Chondropoma ( Chon -

dropomium) sw iftii azuense Bartsch, 1946: Azua. Dominican

Republic. Cliondroj>oma ( Chondropomium ) swiftii bar-

ahonense Bartsch, 1946: Barahona, Dominican Republic.

Chondropoma (Chondropomium) swiftii saturation Bartsch,

1946: L. saturatus , fill, glut; perhaps “filled with dark color.

Page ISO

THE NAUTILUS, Vol. 131, No. 3

Genus Chondropomella Bartsch, 1932

Type Species: Ctjclostoma magnificum Pfeiffer, 1852. bv

original designation.

Description: Shells very' large for family (to ca. 32 mm

length), turbinoid ratlier than elongate, usually decollate.

Protoconch of 1.5 smooth, minute whorls. Final whorl barely

detached for short distance from previous whorl. Axial

sculpture of wide, flattened, close-set ribs. Spiral sculpture

absent except for threads in umbilicus. Overall sculpture

very smooth, often polished. Suture serrate but lacks fused

tufts. Adult shell with reflected lip. Lip double, inner lip

usually erect and separate from outer lip. Outer lip broadly

expanded but narrowest facing umbilicus; adnate or very

narrowly separated f rom previous whorl. Color pattern of

spiral bands and blotches, continuous or interrupted; pattern

unknown in Chondrojiomella platychilum . Operculum only

known for C. magnificum : paueispiral with pseudolamella

occupying inner proximal half of each opercular whorl, outer

half with erect lamellae oriented along growth lines. Radula

known only from C. magnificum: inner marginal tooth of

radula broadly triangular, long side being proximal, with

irregular coarse serrations, lacking denticle on lateral side.

Animal as in Chondropomium.

Remarks: Chondropomella was introduced as Chron-

dropornella on p. 2, but spelled Chondropomella in all other

instances. Watters and Duffy (2010) placed Chondropoma

asymmetricum Pilsbry, 1922, and Chondropoma inaequi-

lahrum Bartsch, 1946, in Chondropomella . These two are

here shown to be synonymous anti to belong to the new,

distinctive genus Superbipoma, based on phylogenetic,

radular, and conchological evidence.

When Bartsch named Chondropomella (1946) he was

unaware of the operculum of either of the two species that

he placed in the genus. Pfeiffer (1854a) described

a ehondropomine operculum for Chondropoma magnif¬

icum Pfeiffer, 1852, which nevertheless appears pseu-

dolamellate a subsequent publication of bis (Pfeiffer,

1854a: fig. 3).

Chondropomella is known from three rare species.

Chondropomella magnified is the most common but still rare

outside of a few collections; C. playchilum is only known

from a handful of worn shells, and C. virilis is known from

two collections of unknown provenance. The habitat is only

known for C. magnified. Like Chondropomium, it lives in

very xeric limestone foothills with sparse vegetation. Genetic

material was only available for C. magnified, apparently the

only species of the genus that has been collected alive.

Chondropomella magnified (Pfeiffer, 1852)

(Figures 74-88, 183)

Chresoxymy

Chondropoma magnificum “Salle” Pfeiffer, 1852a:

278-279; Pfeiffer, 1853a: 197; Pfeiffer, 1854c: pi. 47,

figs. 20-22; Pfeiffer, 1854d: 365-366; Pfeiffer,

1858b: 136.

Cyclostonui {Chondropoma) magnificum "Salle" Pfeiffer,

1852: Pfeiffer, 1854a: 142, pi. 13, figs. 3a-c.

Cyclostoma magnificum Pfeiffer, 1852: Pfeiffer. 1854c: pi.

47, figs. 20-22; Pfeiffer, 1854d: 365-366.

Chondropoma magnificum “Salle”: Adams and Adams,

1856: 295; Chenu, 1859: 493, fig. 3650; Bland, 1861:

355; Pfeiffer, 1865: 145; Pfeiffer, 1876: 190; Kobelt,

1879: 198; Tryon, 1883: 284, pi. 76, fig. 82; Crosse,

1891: 170.

Chondropoma magnified (Pfeiffer, 1852): Reeve, 1863: pi.

1. figs. lb,c [not la, see below],

Chondropoma ( Chondropomium ) magnified (Pfeiffer,

1852): Henderson and Bartsch, 1920: 60.

Chondropoma (Chondropomella) magnified (“Salle”

Pfeiffer, 1852): Bartsch, 1932: 3.

Chondropoma (Chondropomella) magnificum (Pfeiffer,

1852): Bartsch, 1946: 38-39, pi. 8, fig. 7.

Chondropomella magnified (Pfeiffer, 1852): Watters,

2006: 59, 60, 341-342.

Chondropomella elegans Watters and Duffy, 2010: 10-11,

figs. 32-36.

Chondropomella magnificum [.sic] (Pfeiffer, 1854):

Watters and Duffy, 2010: 10, 11, fig. 31.

non Chondropoma magnificum var. Reeve, 1863: pi. 1, fig.

la[= Cyclostoma gutierrezi “Poey” Pfeiffer, 1858a ,fide

Arango v Molina, 1878: 32 and Crosse, 1890: 296].

Description: Shell very large for family (largest adult

specimen 31.5 mm maximum length, decollate, including

peristome; smallest adult specimen 22.7 mm length,

decollate, including peristome), solid, polished, conic,

umbilicus minute. Protoconch of 1 .5 white, smooth

whorls. Adult specimens usually decollate. Teleoeoneh of

4-5.5 whorls, final whorl adnate or detached for last l/6th

of whorl. Suture indented, shallowly channeled. Peri¬

stome double, tear drop-shaped. Outer lip thickened,

composed of numerous lamellae, rolled back abapertur-

allv. undulating, folded over umbilicus, narrowest facing

umbilicus, adnate or verv narrowly detached from pre¬

vious whorl, auriculate posteriorly. Inner lip adherent to

outer lip or barely exserted. Spiral sculpture present only

as 4-5 very feeble cords within umbilicus. Axial sculpture

of numerous (160-170) narrow, widely spaced, very low

undulating ribs, forming minute serrations at suture.

Background color white to tan with broken spiral bands of

spots and chevrons; bands extend onto face of outer lip but

not onto inner lip. Operculum with thick pseudolamella

divided into two regions: outer half of spiral with very-

coarse erect lamellae, inner half fused into smooth plate.

Radula as in genus. Animal tan to yellowish with dark-

brown band between eyes and across base of snout.

Type Material: Chondropoma magnificum Pfeiffer,

1852: ? Syntypes (3): NHMUK. Three specimens glued to

a board at NHMUK, unnumbered, of "magnificum var.

W. Indies” from Cuming collection; back of board reads

“No. 132, Cycl magnificans var: Salle. St. Domingo.”

These may! be syntypes but do not match Pfeiffer (1854a:

pi. 13, figs. 3a, b). They are not Chondropoma magnificum

G.T. Watters and P. Larson, 2017

Page 1S1

Figures 74-101 . Clwndropornella species. 74—88. Chondropomella magnified (Pfeiffer, 1852). 74, 75. Pfeiffer, 1854a: pi. 13, figs. 3a,b. 76.

Chondrojxmiella elegans Watters and Duffy, 2010. UP’ 420732, holotype, 27.2 min. 77, 78. UF 216670b, 26.4 mm. 79. UF 216387, 23.2 mm.

80. UF 216387, 22.7 mm. 81. UF 216671,' 26.2 mm. 82. UF 216671, 24,8 mm. 83. GTW 7639a, 26.1 mm. 84. OSUM 42368, 28.6 mm. 85.

UF 217950, 27.5 mm. 86. 1 labitat. 87. 1 jving individual. 88. OSUM 42368, radula. Scale bar = 200 pm. 89-95. Chondropomella platychilum

(Pfeiffer, 1851). 89, 90. Cijclostoma Uitilabre “d’Orbigny” of Pfeiffer, 1847: pi. 10, figs. 26, 27. 91, 92. UF 216766, 29.6 mm. 93. UP’ 216767,

26.3 mm. 94. UF 216768, 24.3 mm. 95. USNM 354984, 28.6 mm. 96-101. Chondropomella mrilis (Bartsch, 1946). 96, 97. USNM 504145,

holotype, 18.7 mm. 98, 99. USNM 425379, 21.8 mm. 100. USNM 425379, 13.4 mm. 101. USNM 425379, 18,5 mm.

Page 1S2

THE NAUTILUS, Vol. 131, No. 3

var. Reeve, 1863. Chond ropornella elegant; Watters and

Duffy, 2010: Holotype: UF 420732; Paratype: OSUM

3 2485 ( 1); NIIMUK 1 99349( 1 ) .

Tvpe Locality: Clwndroponui magnificum Pfeiffer, 1852:

“Habitat in insula Haiti” [“Rocks at the entrance of

a cave at Barrera, Dominican Republic [from the

French], fide Crosse, 1890.] [“This locality is in the

Province of Azua about 16 miles southwest of the town of

Azua,” fide Bartsch, 1946]. Chond ropornella elmans

Watters and Duffy, 2010: “Dominican Republic, Inde-

pendencia Province, ~ 8 km SW of Duverge, Puerto

Escondita.”

Type Figured: Chond roponui magnificum Pfeiffer,

1852: Pfeiffer, 1854a, pi. 13, figs. 3a-e. Chondropomella

elegant Watters and Duffy, 2010: Watters and Duffy,

2010, figs. 32, 33.

Distribution: Along the slopes facing Lago Enriquillo

from the northwest shore along the southern shore to the

isolated Sierra Martin Garcia and the lowlands sur¬

rounding the Bahia de Ocoa. Locally common.

Habitat: Hillsides under fossilized coralline blocks in

open xerie areas with sparse vegetation. It has also been

observed suspending itself from short mucus threads from

rocks.

Other Material Examined (517 Specimens): Do¬

minican Republic. Independencia Province. OSUM

42358(16), Bartolome; UF 216850a(15), 7 km SF of

|imani; GTW 1 603 1 e ( 3 ) , ca. 0.8 km from Puerto

Escondido, S of Duverge; UF 217956(17), 520 m. 3 km

SF of Puerto Escondido; UF 217950(27), UF 217950a

(12), 830 m, 4 km ESE of Puerto Escondido; UF 35087

(1) , 7.5 km S of Puerto Escondido; UF 217960(4), 8 km

WNW of Puerto Escondido; UF 217954(1), 490 m, 11 km

WNW of Puerto Escondido; UF 217953(6), UF 217959

(6), 2 km SW of Duverge; UF 217951(12), UF 217955

(12), UF 217957(20), 9 km WNW of Duverge; UF 217982

(2) , 12 km WNW of Duverge; OSUM 42368(17), 466 m,

on footpath up hill, 1.3 km E of center of Puerto

Escondido, 18.3290° N, -71.5593° W; GTW 16031e(l),

42 m, quarried area along road to Angostura, 1 km NNW

of Angostura, 18.3412° N , -71.4207° W; GTW 16031d(3),

473 rn, along RD 541, 2.3 km NNW of El Naranjo,

18.3514° N, -71 .6279° W; UF 45679(3), 2 km F 2 km S of

Angostura; UF 45669(3), 1 km F 2 km S of Angostura.

Azua Province. GTW 7639a(l), 244 m, on trail at

Barrera; UF 216669(30), UF 216673(2), 150 m, Barrera;

UF 216670b(39), UF 216671(37), 160 m, Barrera; UF

216769(1), 560 m. Sierra Martin Garcia, 5 km NNW of

Barrera; UF 216674(1), W slope of Loma del Aguacate;

UF 216387(163), 80 m, 7 km E of Azua; UF 216506(2),

UF 216521(10), UF 216522(16), UF 216523(21), Sierra

Fl Numbrero, 8 km SF of Las Charcas; UF 46258(1),

UF 216524(2), 100 m, 12 km ESE of Las Charcas; UF

216520(24), UF 216519(21), 3 km W of Hatillo. Bar-

ahona Province. UF 216672(4), 3 km NE of Puerto

Alejandro.

Variation Among Specimens: Specimens differ in the

degree of coloration and the degree of expansion of the

lip. Rare specimens may be without any color pattern.

Comparison With Other Species: The lip may be

barely adnate to the previous whorl in this species hut it is

broadly fused in C. platychilum and C. virilis. Chon¬

dropomella platychilum has a shorter spire and more

rounded whorls than the other species. Chondropomella

virilis differs from the other species in having a white

shell with a few spiral brown bands that extend over the

greatly expanded lip and a coarser sculpture of raised

narrow lamellae rather than flattened ribs.

Remarks: Chondropoma magnificum was recognized by

Bartsch (1946) as the population from the isolated Sierra

Martin Garcia having a wide, wholly adnate outer lip.

Chondropomella elegant was differentiated as the western

population in the Lago Enriquillo rift valley having

a narrower, solute outer lip. A third population occurs

around the Bahia de Ocoa that resembles C. elegant.

Examination of over 500 specimens suggests the variation

in shell moqvhologv within a population is as great as that

between populations and the three are considered a single

taxon pending further genetic studies.

Original Description: Pfeiffer, 1852a: 278-279. From

the Latin. “Shell umbilicate, ovate-conic, thin, longitu¬

dinally plicate-striated, diaphanous, a little shining, white,

variously interrupted brown bands, middle widest from

angular striped shapes, ornate; spire inflated, apex sub-

truncate; suture simple; remaining 5 whorls a little convex;

last one rounded; aperture vertical, oval; peristome

simple, shining, white, brown-spotted, top with expanded

hood, narrowly adnate to previous whorl, narrow, sinuate

at the umbilicus, then broadened into a open tongue, right

margin and base wide reflexed arching. Operculum car¬

tilaginous, fiat, pale corneous.

Etymology: Chondropoma magnificum Pfeiffer, 1852:

L. magnificus , magnificent. Chondropomella elegant

Watters and Duffy, 2010: L. elegant , elegant.

Chondropomella platychilum (Pfeiffer, 1851)

(Figures 89-95, 183)

Chresonvmy

Cyclostoma latilahre “d’Orbigny” Pfeiffer, 1S47: 78-79,

pi. 10, figs. 26, 27 [misidentification, non Cyclostoma

latilahris d’Orbigny, 1842]; Pfeiffer, 1851: 169 [in

synonymy of Cyclostoma platychilum Pfeiffer, 1851];

Pfeiffer, 1852a: 261, 294 [in synonymy of Cyclostoma

platychilum Pfeiffer, 1851]; Pfeiffer, 1852b: 41 [in

synonymy of Cyclostoma platychilum Pfeiffer, 1851];

Bartsch, 1946: 37 [in synonymy of Cyclostoma pla¬

tychilum Pfeiffer, 1851].

Cistula ? platychila Pfeiffer, 1851: 169; Pfeiffer, 1852a:

26L Pfeiffer, 1852b: 41; Pfeiffer, 1853a: 182; Pfeiffer,

1858b: 130; Pfeiffer, 1865: 140; Pfeiffer, 1 876: 187.

G.T. Watters and P. Larson, 2017

Page 1S3

Cyclostoma platychilum (Pfeiffer, 1851): Pfeiffer, 1853b:

266; Pfeiffer, 1858b: 130.

Cistula plotychilo (Pfeiffer, 1851): Adams and Adams,

1856: 294.

Cistula platychila? (Pfeiffer, 1851): Arango y Molina,

1878: 31.

Chondropoma (Chond roponiella) platychilum (Pfeiffer,

1851): Bartsch, 1946: 37, pi. 7, fig. 1.

Chondropoimlla platychila [sic] (Pfeiffer, 1851): Watters,

2006: 60, 410—41 1 .

Chond roponiella platychilum (Pfeiffer, 1851): Watters

and Duffy, 2010: 10, 11.

non ?Clstula platychila “Pfeiffer” Arango y Molina, 1878:

31 [= Cyclostoma latilahris d’Orbigny, 1842, fide

Bartsch, 1946: 37].

Description: Shell very large for family (largest adult

specimen 29.6 mm maximum length, decollate, including

peristome; smallest adult specimen 24.3 mm length,

decollate, including peristome), solid, low-spired, umbi¬

licus wide. Protoconch decollated from all examples.

Adult specimens decollate. Teleoconeh of 3.5 whorls, final

whorl barely detached for last l/6th of whorl. Suture

indented, shallowly channeled. Peristome double, oval.

Outer lip thickened, widely expanded, composed of nu¬

merous lamellae, rolled back abaperturallv, undulating,

folded over umbilicus, narrowest facing umbilicus,

broadly adnate with previous whorl, widely auriculate

posteriorly and at 8 o'clock position. Inner lip narrowly

erect. Axial sculpture absent but perhaps worn away in all

specimens. Spiral sculpture of few weak threads in um¬

bilicus. Suture smooth. All specimen worn and faded with

no indication of color patterns. Operculum, anatomy, and

radula unknown.

Type Material: Not located. On his death, many of

Pfeiffer’s specimens passed to Heinrich Dohm

(1838-1913). Dohrn’s collection was incorporated into

the Museum Naradone in Szczecin, Poland. Although

some of the other collections of that museum were moved

to the Pomfnersches Landesmuseum, Greifswald, Ger¬

many, the natural history collections are not there (pers.

comm., B. Frenssen, 2016). Dohm’s collection, and

Pfeiffer’s types, were probably destroyed by Allied

bombing of the Museum Naradone in WW II.

Type Locality: “Cuba,” for Cyclostoma latilabre "d’Or¬

bigny” of Pfeiffer, 1846. Cistula? platychila Pfeiffer, 1851:

“Habitat...?” H ere restricted to 8 km NW of Oviedo,

Pedernales Province, Dominican Republic.

Type Figured: Pfeiffer, 1847, pi. 10, figs. 26, 27?

Distribution: All records are for the Oviedo region in the

southern Sierra Baoruco foothills, one record around the

Lago de Oviedo.

Habitat: This species is very rare in collections. It is likely

that its exact habitat has not been discovered.

Other Material Examined (5 Specimens): Dominican

Republic. Pedernales Province. UF 216768(1), 8 km

NW of Oviedo; UF 216767(1), along shore of Lago de

Oviedo, 1 km SE of Oviedo Viejo; UF 216756(1), 210 m,

17 km NW of Oviedo; USNM 354984(2), Trnjin [Lago de

Oviedo].

Variation Among Specimens: The few known speci¬

mens are remarkably uniform in shell characteristics.

Comparison With Other Species: See under C

nmgnificum.

Remarks: Tl lis is one of the largest species in the family -

to ea. 30 mm length. Pfeiffer seems to have had a fresh

specimen before him but all other shells known to us are

weathered and faded. The color pattern, if any, is un¬

known and Pfeiffer only described it as “diaphanous,

hyaline-white. No material was available for phylogenetic

study.

Original Description: Cyclostoma latilabre “d’Orbigny’

Pfeiffer, 1847: 78-79. From the Latin. “Shell umbilieate,

ovate-oblong, decollate, a little thin, smooth, diaphanous,

hyaline-white; 4 convex whorls, base of last spirally and

obsoletely subsulcate; aperture vertical, oval; peristome

double, inner subexpanded, hardly prominent, external

slightly thickened, angle widely reflexed, bordered to the

penultimate whorl with thick narrow angulate junction

columella expanded in middle angle.”

Etymology: G. platys , broad + G. cheilos , mouth. A noun

in apposition.

Chondropomella virilis (Bartsch, 1946)

(Figures 96-101)

Chresowmy

Incertipoma virile Bartsch, 1946: 171, 174-175, pi. 30,

fig. S.

Chondropomella virilis (Bartsch, 1946): Watters, 2006:

60, 547.

Chondropomella virile [sic] (Bartsch, 1946): Watters and

Duffy, 2010: 10, 11.

Description: Shell medium to large for family (largest

adult specimen 21.8 mm maximum length, decollate,

including peristome; smallest adult specimen 13.4 mm

length, decollate, including peristome), solid, low-spired,

umbilicus wide. Protoconch decollated from all examples.

Adult specimens decollate. Teleoconeh of 3.5 whorls, final

whorl almost completely adnate with previous whorl.

Suture indented, channeled. Peristome double, oval.

Outer lip thickened, widely expanded, composed of nu¬

merous lamellae, rolled back abaperturallv, undulating,

folded over umbilicus, narrowest facing umbilicus, weakly

adnate with previous whorl, widely auriculate posteriorly

and at 8 o’clock position. Inner lip narrowly erect. Axial

sculpture of ea. 110-120 narrow, erect, widely spaced

lamellae. Spiral sculpture of ea. 3 cords in umbilicus.

Page 184

THE NAUTILUS, Vol. 131, No. 3

Suture smooth. The holotvpe has vestiges of color pattern

hut all other specimens are worn and faded. The holotvpe

has 9 nearly continuous, narrow, spiral tan bands between

suture and base; hands extend onto ahapertural face of

outer lip hut only faintly onto adapertural side. Opercu¬

lum, anatomy, and radula unknown.

Type Material: Holotvpe: USNM 504145.

Tvpe Locality: “Haiti.

Type Figured: Bartseh, 1946, pi. 30, fig. 8.

Distribution: Unknown. The two lots of this species are

stated to come from “Haiti” hut it is not clear if this refers

to tin' country of Haiti or the island of Hispaniola. We

expect that, like other members of the genus, it is endemic

to the Barahona Peninsula awaiting rediscovery.

Habitat: Not recorded.

Other Material Examined (24 specimens): “Haiti.”

USNM 504145(1); USNM 425379(23), “probably Haiti."

Variation Among Specimens: The few known speci¬

mens vary in the degree of expansion of the lip.

Comparison with Other Species: See under C mag-

nifica. The lamellate sculpture on the whorls is unique.

Remarks: This rare, distinctive species, like C. pla-

tychilwn , is known mainly from worn specimens. The

lamellate sculpture is unlike other Chondropornella and

discovery of specimens with opereula (or live ones with

genetic material) may necessitate reallocation to a differ¬

ent genus. It hears some resemblance to members of

Sallepoma from the western Tiburon peninsula.

Etymology: L. virilis, manly.

Genus Clydonopom a Pilshry, 1933

Type Species: Ttidora nobilis Pfeiffer, 1852. hv original

designation.

Parachondria (Clydonopoma) Pilsbiy, 1933: 127.

Parachondria (Eccritoponui) Pilshry, 1933: 128 [type

species Parachondria peasei Pilshry, 1933, bv original

designation].

Description: Shells medium to very large for family (to

ea. 33 mm length), turbinoid to elongate conic, often

decollate. Protoconch of 1.5 smooth or microscopically

granulate, minute whorls. Final whorl barely detached

from previous whorl for short length. Axial sculpture of

wide, flattened, close-set ribs. Clijdonopoma nobilis, the

type species, and C. poloensis lack spiral sculpture outside

of umbilicus. Remaining species show varying degrees of

very weak spiral sculpture, usually limited to subsutural

hands and often present only as beads or scalloped edges

to axial ribs. Overall sculpture appears very smooth, often

shiny, undulating below suture. Suture minutely serrate

but lacks fused tufts. Lip double, inner lip usually erect

and separate from outer lip. Adult shell with reflected lip.

Lip usually not widely expanded, narrowest facing um¬

bilicus; adnate or very narrowly separated from previous

whorl. Color pattern of spiral hands and chevrons, con¬

tinuous or interrupted, nearly unicolored in some species.

Operculum with pseudolamella. Inner marginal tooth

of radula broadly rounded with obsolete serrations

and very weak denticle on outer side. Animal as in

Chondropomium.

Remarks: Although synonymized with Chondropomium

by Watters (2006), Clydonopoma is here shown to be

a distinct genus based on phylogenetic and opercular

evidence. Bartseh (1946) placed four species into his

Kisslingia : K. hinchensis, K. poloen.se, K. bahorucensis (all

Bartseh, 1946), and K. clenchi (Pilshry, 1933) [= ber-

mudezi Jaume, 1984]. The latter three are endemic to the

Barahona Peninsula. Kisslingia hinchensis , the tvpe spe¬

cies of Kisslingia, occurs in northern Haiti. Examination

shows that the three Barahona taxa are not congeneric

wi th hi h inchensis and are better placed in Clydonopoma.

Articmlipoma Bartseh, 1946, is very similar to Clydono¬

poma but lacks a pseudolamella.

Clydonopoma is endemic to the Sierra de Baoruco and

eastern-most Massif de la Selle. Species occur in different

habitats than do those of Chondropornella and Chon¬

dropomium. Clydonopoma lives in mesic upland forests

and pine savannahs whereas Chondropomium and

Chondropornella live on the highly xeric foothills and

valleys below among cacti and agave.

The radula differs from that of Chondropomium in

having the inner marginal very weakly serrate rather

than deeply incised. The radula is very similar to

Chondropornella .

The extent to which the pseudolamella covers the basal

portion and radiating ribs of the operculum varies greatly

even among individuals of the same population. The

pseudolamellate layer may be completely worn away in

some specimens revealing the pin-wheel radiating, erect

calcareous ribs, rendering the operculum “rhytidopo-

mine” according to the terminology of Henderson and

Bartseh (1920). It is probable that the pseudolamellate

condition is an extension of that opercular type.

Clydonopoma bahorucense (Bartseh, 1946)

(Figures 102-107, 183)

Chresonymy

Kisslingia bahorucensis Bartseh, 1946: 112, 113-114, pi.

17, fig. 2.

Licina barorucensis (Bartseh, 1946): Watters, 2013:

map 1 .

Description: Shell medium for family (largest adult

specimen 15.4 mm maximum length, decollate, including

peristome; smallest adult specimen 13.4 mm length,

decollate, including peristome), thin, low-spired, umbi¬

licus open but narrow. Protoconch retained in many adult

specimens; of 1.5-2 microscopically granulate whorls.

G.T. Watters and P. Larson, 2017

Page 185

Figures 102-128. Clydonopoma species 102-107. Clydonopoma bahonicense (Bartsch, 1946). 102, 103. USNM 504038. holotype, 14.7

mm. 104. UK 217965. 15.4 mm. 105. UK 217965. 13.4 mm. 106. UP 217965, 14.5 mm. 107. UF 217965, 14.0 mm. 108-1 13. Clydonopoma

bartschi (Watters, 2012). 108, 109. UF 446073, holotype, 20.1 mm. 110. OSUM 36520, paratype, 20.6 mm. Ill, 112. UF 216467, 18.7 mm.

113. UF 216463, 16.7 mm. 114—117. Clydonopoma bennudezi (Jaume, 1984). 114, 115. ANSP 160995a, holotype of Chondnrpoma clenchi

Pilsbry, 1933, 18.0 mm [photo courtesy of ANSP]. 116, 117. USNM 426036, paratype, 14.3 mm. 1 18-128. Clydonopoma nobile (Pfeiffer, 1852).

1 18. OSUM 42366, radula. Scale bar = 4(H) pan. 1 19, 1 20. Pfeiffer, 1854a: pi. 13, figs. 2a, b. 121, 122. GTW 7089a, 28.9 mm. 123. OSUM 42366,

28.2 mm. 124. OSUM 42366, 32.4 mm. 125. GTW 7089b, 26.9 mm. 126. Li\ing individual. 127. Individuals mating. 128. Type locality.

Page 186

THE NAUTILUS, Vol. 131, No. 3

Teleoconcli of 3.5-4 whorls, final whorl narrowly adnate

with previous whorl. Suture indented, channeled. Peri¬

stome single, oval. Outer lip narrow, barely expanded,

narrowest facing umbilicus, weakly adnate or just touching

previous whorl, minutely or not auriculate posteriorly.

Axial sculpture of ca. 150 narrow, erect, widely spaced

lamellae. Spiral sculpture of variable number of weak

cords extending from suture to mid-whorl in some

specimens or covering entire shell in others, weak threads

in umbilicus. Intersections of axial and spiral sculpture

form weaklv scalloped surface. Axial elements render

suture finely serrate. Base color white with overlain weak

pattern of brown zig-zags and spots, most prominent

below suture and on base where they form interrupted

bands; overall appearance is pale with limited markings.

Markings may appear on lip as well. Operculum with

pseudolamella. Anatomy and radula unknown.

Type Material: Holotvpe: USNM 504038 (2 specimens

in lot, holotype listed as 14.7 mm long specimen).

Type Locality: “Polo District, Bahoruco Mountains,

Dominican Republic, at an elevation of 2,000 feet.”

Type Figured: Bartsch, 1946, pi. 17. fig. 2.

Distribution: Bartsch (1946) listed specimens from the

Polo region, Loma de Cielo (Capo de Cielo), and 1.6 km

north of Maniel Viejo. We add the area of Angostura in

Independeneia Province. With the exception of the

Angostura specimen at 42 m elevation, all localities are

between 600-1070 m in the eastern and northeastern

Sierra de Baoruco.

Habitat: Mesie limestone ravines; all of the UF sites

(below) have been cleared for coffee groves. The

Angostura site has been quarried.

Other Material Examined (46 Specimens): Domini¬

can Republic. Barahona Province. UF 217966(19),

910 m, 5 km NNF of Polo; UF 217974(5), 910 m, 7 km

NNE of Polo; UF 217968(3), 710 m, 7 km NNE of Polo;

UF 217965(18), 910 m, 14 km S of Cabral. Inde¬

pendeneia Province. OSUM 42371(1), 42 m, quarried

area along road to Angostura. 1 km NNW of Angostura,

18.3412° N, -71.4207° W.

Variation Among Specimens: This species and C

titanum show more variation in color than any other

Clydonopoma , ranging from heavily patterned with brown

chevrons and blotches to nearly all white. The subsutural

row of evenly spaced brown markings seems to be

consistent.

Comparison with Other Species: This species has the

most developed sculpture of the species of Ch/donopoma .

The intersections of the spiral and axial sculpture form an

extensive scalloped surface that is not seen in other

species. Clydonopoma bermudezi is most similar to this

species but only has a few weak scalloped spiral cords

below the suture; it is less elongate and smaller as well.

Clydonopoma bahorucense is svmpatrie with C. poloense

but is smaller and has a more channeled suture; C.

poloense lacks spiral sculpture outside of the umbilicus.

Etymology: Bahoruco Mountains, Dominican Republic.

Clydonopoma bartschi (Watters, 2012)

(Figures 108-113, 183)

Chresoxymy

Licina bartschi Watters, 2012: 13, figs. 45-47; Watters,

2013; map 1, plate 1, fig. 5.

Description: Shell medium for family (largest adult

specimen 19.9 mm maximum length, decollate, including

peristome; smallest adult specimen 10.3 mm maximum

length, including peristome), solid, conic, umbilicus

minute. Protoconch whorls decollate in adults, consisting

of 1.5 smooth, prominent whorls, well-delimited from

teleoconcli. Teleoconcli of 4 whorls, final whorl detached

just before peristome. Suture deeply channeled. Peri¬

stome single, thick, tear drop-shaped. Lip narrow, barely

expanded on outer side, auricle absent. Spiral sculpture

present only as 3-5 very feeble cords below suture ren¬

dering axial sculpture scalloped, 7-9 cords in umbilicus.

Axial sculpture of narrow, fine ribs separated by 2-3x their

width, ca. 150 on final whorl, forming minute cusps at

suture. Suture serrate. Background color glossy pale tan

with complex pattern of dense, brown, very fine, zigzag

markings axially aligned, sometimes forming interrupted

spiral bands on base and within umbilicus; growth rests

marked by axial rows of tiny dark spots; suture with

pattern of alternating white and dark brown patches;

markings persist on both sides of peristome and are visible

through inside of shell. Operculum paucispiral with

pseudolamella that extends 3/5ths of way to outer margin,

corneous portion with fine granular deposit. Anatomy and

radula unknown.

Type Material: Holotype: UF 446073; Paratype 1:

Nil M UK 201 10338, slightly juvenile; Paratype 2: OSUM

36520; both paratypes from the type locality.

Type Locality: “9.5 km ENE of Las Mercedes, Ped-

emales Province, Dominican Republic, on a mountain top

at 1,300 m elevation.

Type Figured: Watters 2012, figs. 45, 46.

Distribution: This is an upland species usually found

above 1000 m in the western Sierra de Baoruco.

Habitat: On limestone bills and in ravines under rocks in

scrub and pine s a van Mb.

Other Material Examined (83 Specimens): Domini¬

can Republic. Pedemales Province. GTW 14630d(l),

1285 m, abandoned quarry near end of road that runs

from Cabo Rojo N past Las Mercedes into Sierra de

Baoruco, 18.1203° N, -71.5725° W; UF 216467(7), 1000 m,

W rim of Hoya de Pelempito; UF 216463(29), 1410 m.

G.T. Watters and P. Larson, 2017

Page 1ST

Loma El Aceitillar; UF 249186 (10), 240 in, 19 km N of

Pedemales; UF 217958(46), 1510 in, 20 km N of Las

Mercedes.

Variation Among Specimens: Specimens are very

uniform in most characteristics but differ in tl it* degree of

coloration. The base color varies from pale tan to a greyish

cast.

Comparison with Other Species: Clydonopoma bart-

schi is very similar to C. peasei and C. poloense. All are

large, similarly colored species. Clydonopoma poloense

lacks the spiral grooves below the suture seen in C.

bartschi and C. peasei. Clydonopoma peasei is more

obviously turbinoid than the other two taxa. Clydonopoma

nobile is the largest of the genus and is usually almost

unicolored, often with a purplish tinge. Clydonopoma

pumilum is smaller, has a well-developed outer lip and

a pattern of minute dots and dashes rather than the

complex pattern of chevrons and bands seen in C. bart¬

schi, C. poloense, and C. peasei. Clydonopoma pumilum

differs from C. bahorucensis and C. hemmdezi in the lack

of spiral sculpture on the posterior half of the whorl.

Nevertheless, all of these species are closely related and

similar in appearance.

Remarks: This species was originally placed in Licina,

a genus requiring a thorough review. Licina has a pseu-

dolamellate operculum like Clydonopoma but possesses

distinct spiral sculpture over the entire shell. Species

assigned to Licina probably represent several genera. The

original images of this species in 2012 were unfortunately

vertically foreshortened by the printer. No material was

available for phylogenetic study.

Etymology: Paul Bartsch (1871-1960), malacologist at

the US National Museum and expert on the Annulariidae.

Clydonopoma bermudezi (Jaume, 1984)

(Figures 114—117, 184)

Chresonymy

Chondropoma ( Chondropomium ) clenchi Pilsbry, 1933:

126, pi. 9, figs. 2, 3 [non Chondropoma erne.sti clenchi

Aguayo, 1932b],

Kisslingia clenchi (Pilsbry, 1933): Bartsch, 1946: 1 12, 1 15,

pi. 17, fig. 1.

Licina? clenchi (Pilsbry, 1933): Baker, 1964: 169.

Kisslingia bermudezi Jaume, 1984: 3 [replacement name,

see Remarks].

Chondropoma (Chondropomium) compressa Boyko and

Cordeiro, 2001: 30 [replacement name, see Remarks].

Licina clenchi (Pilsbry, 1933): Watters, 2013: map 1.

Description: Shell medium for family (largest adult

specimen 18.0 mm maximum length, decollate, in¬

cluding peristome, smallest adult specimen 13.0 mm

maximum length, including peristome), solid, conic,

umbilicus minute. Protoconch whorls usually retained

in adults, consisting of 1.5 smooth whorls, well-

delimited from teleoconch. Teleoconch of 3.5-4

whorls, final whorl detached just before peristome.

Suture deeply channeled. Peristome single, thick, tear

drop-shaped. Outer lip narrow, barely expanded on

outer side, auricle weak or absent. Spiral sculpture

present as ca. 8 wide, undulating cords below suture,

gradually diminishing in strength, and as ca. 10 cords in

umbilicus. Axial sculpture of numerous narrow, fine

closely-set ribs, ca. 1 10 on final whorl, forming minute,

elongate cusps at suture. Suture serrate. Intersections

of spiral and axial sculpture finely beaded or scalloped.

Background color white or pale tan with complex

pattern of faint, brown, very fine, zigzag markings axially

aligned, sometimes forming interrupted spiral bands on

base; suture with pattern of alternating white and dark

brown patches; markings do not persist onto either side

of lip. Rarely almost entirely patternless, but sutural

markings are consistent. Operculum paucispiral with

pseudolamella that extends 3/5ths of way to outer

margin, corneous portion with fine granular deposit.

Anatomy and radula unknown.

Type Material: Holotype: ANSP 160995a; Paratypes:

81534(4); USNM 426036(2); AMNH 81534(3).

Type Locality: “Sr. Del Monte coffee plantation, Station

85, between the main bate and the top of Alies in the

coffee finca proper, and in a verdant gully near Salvation,

at about 3,000 feet.” As Pilsbry (1933: 121) pointed out.

Salvation was the name of a division of the Del Monte

coffee finca and “of course not to be found on any

published map.”

Type Figured: Pilsbry, 1933, pi. 9, figs. 2, 3.

Distribution: Known only from tbe type locality in tbe

eastern Sierra de Baoruco.

Habitat: Pilsbry (1933: 122) described the habitat at tbe

type locality: “a gully near Salvation; the walls are steep

and with luxuriant growth of trees and underbrush; the

sides of the nearly waterless stream bed veiy wet, mossy,

with deep humus, and lined with boulders.

Other Material Examined (2 Specimens): Dominican

Republic. USNM 426036(2), from type locality.

Variation Among Specimens: Only two specimens have

been examined by us.

Comparison with Other Species: See under C

bartschi.

Remarks: This species was renamed twice based on the

fact that Chondropoma clenchi Pilsbry, 1933, was a junior

primary homonym of Chondropoma erne.sti clenchi

Aguayo, 1932. Jaume (1984) created the replacement

name Kisslinaia bermudezi. Unaware of this action, Boyko

and Cordeiro (2001) also created a replacement name,

Chondropoma (Chondropomium) compressa. The name

Page 188

THE NAUTILUS, Vol. 131, No. 3

of this taxon thus becomes Cltjdonoponw bemmclezi

(aume, 1984. No material was available for phylogenetic

study.

Etymology: Chondropoma clenchi Pilsbry, 1933: William

James Clench (1897-1984), American malacologist,

M useum of Comparative Zoology, Harvard University.

Kisslingia bermudezi Jaume, 1984: Pedro Joaquin Ber¬

mudez y Hernandez (1905-1979), Cuban naturalist and

geologist. Chondropoma compressa Boyko and Cordeiro,

2001: L. compressa , “to clench,” apparently an un¬

fortunate pun on Clench.

Clydonopoma nob He (Pfeiffer, 1852)

(Figures 118-128, 183)

Chresonymy

Tudora nobilis Pfeiffer, 1852a: 252; Pfeiffer, 1853a: 176;

Pfeiffer, 1858b: 128; Bland, 1861: 355; Pfeiffer, 1865:

138; Pfeiffer, 1876: 185; Kobelt, 1880: 277; Crosse,

1891: 177.

Cyclostoma ( Tudora ) nobile (Pfeiffer, 1852): Pfeiffer,

1854a: 142, pi. 13, figs. 2a-e.

Cyclostoma nobile (Pfeiffer, 1852): Pfeiffer, 1854c: pi. 47,

figs. 27-28; Pfeiffer, 1854d: 366-367.

Cistula ( Tudora ) nobilis (Pfeiffer, 1852): Adams and

Adams, 1856: 294.

Chondropoma ( Chondropomium ) nobilis (Pfeiffer, 1852):

Henderson and Bartseh, 1920: 60.

Parachondria ( Clydonopoma ) nobilis (Pfeiffer, 1852):

Pilsbrv, 1933: 127-129, pi. 6, figs. 14, 16-19; Wenz,

1939: '546, fig. 1454.

Clydonopoma (Clydonopoma) nobile (Pfeiffer, 1852):

Bartseh, 1946: 135-136, pi. 21, fig. 3.

Chondropomium nobile (Pfeiffer, 1852): Watters, 2006:

63, 375; Watters, 2012: 13-14, fig. 48, 53 [animal], 58

[habitat]; Watters, 2013: 4, pi. 2, fig. 14, map 1.

Clydonopoma nobile (Pfeiffer, 1852): Watters, 2013:

map 1 .

Description: Shell very large for farnilv (largest adult

specimen 32.3 mm maximum length, decollate, including

peristome, smallest adult specimen 25 1 mm maximum

length, including peristome), very solid, high-spired,

umbilicus open but minute. Protoconch whorls often

retained in adults, consisting of 1.5 smooth, blunt white

whorls, well-delimited from teleoeonch. Teleoeonch of

4-5 whorls, final whorl detached ca. 14 length before

peristome. Suture deeply channeled. Peristome double,

thick, tear drop-shaped. Inner lip thickened, prominent,

erect, projecting well beyond outer lip. Outer lip thick,

narrowly expanded, lamellate, undulating anteriorly, often

fused with inner lip on umbilical side, with minute pos¬

terior auricle. Outer lip with concave or sinuous un¬

dulation on outer margin. Spiral sculpture present as ca.

1 2-20 wide, thick cords in umbilicus. Axial sculpture of ca.

1 90 verv narrow erect lamellae, widely spaced on center of

whorl, gathered together on base; lamellae grouped in

growth series, sometimes anastomosing where shell has

been damaged. Suture undulating and serrate, each axial

element slightly enlarged and minutely blade-like.

Background color white, brown, or purplish with 10-15

spiral bands of very minute spots alternating with verv

narrow interrupted spiral markings; markings do not

persist onto either side of lip. Operculum paucispiral with

pseudolamella that extends 3/5ths of way to outer margin,

but pseudolamella maybe completely worn away exposing

underlying pin-wheel erect ribs. Animal white with pale

grey foot and pale grey base of snout; large diffuse grey

patch between eves; tentacles orange. Radula as in genus.

Type Material: Three specimens glued to board at

NHMUK, unnumbered, none marked with an “X,” label

reads “Haiti;” these are possible syntypes but do not

match Pfeiffer, 1854a: pi. 13, figs. 2a-e.

Type Locality: “in insula Haiti. Restricted here to

Virgen de San Rafael, Barahona Province, Dominican

Republic.

Type Figured: Pfeiffer, 1854a, pi. 13, figs. 2a-c.

Distribution: This species occupies a restricted coastal

range along the eastern Barahona Peninsula from Parafso

north and then curves inland along the northeastern Si¬

erra de Baoruco to Barahona and Cabral. Locally

common.

Habitat: Under limestone rocks and in stone fence

crevices in verdant, mesic forest, found to ca. 450 m el¬

evation. Individuals were observed actively crawling about

and mating in the early morning but they had disappeared

by late morning. It occurs with uropygid arachnids, which

mav be predaceous on this snail; shells often show signs of

repaired breakage. This species can occupy lower eleva¬

tions than its congeners bv virtue of its coastal distribution

in the prevailing offshore winds that create a mesic habitat

at sealevel.

Other Material Examined (178 Specimens): Do¬

minican Republic. Barahona Province. UF 217802

(10), UF 217810(2), 6 km SSW of Cabral; UF 217809(2),

20 m, 4 km E of Cachon; UF 217803(16), UF 217805(13),

200 rn, 5 km SE of Barahona; UF 217811(6), UF 217812

(4), 300 m, 5 km SE of Barahona; UF 217815(25), 450 m,

5 km SF of Barahona; UF 155894(11), 15 km S of

Barahona; GTW 7089a(l), 16 km S of Barahona; UF

217813(4), 400 m, 2 km SW of La Guazara; UF 217808

(2). 180 m, 4 km NF of La Guazara; UF 217807(2), 150 m,

6 km NF of La Guazara; GTW 70S9e(2), 18 m, cleared

area off RD 44, near Juan Esteban, 18.1294° N, -71.0697°

W; UF 45691(25), 10 km NNF ofParaiso; GTW 7089b(10),

GTW 7089c(l), OSUM 42366(10), Villa Miriam, Virgen de

San Rafael, ca. 7 km NNF ofParaiso, 18.4601° N, -69.6096°

W; UF 217814(25), 30 m, 4 km SW of La Cienaga; UF

217804(10), UF 217806(10), 30 m, 6 km SW of La Cienaga.

Variation Among Specimens: The base color of the

shell varies from white to a pu rplish-brown color hut the

very fine spiral bands and dots are always present.

G.T. Watters and P. Larson, 2017

Page 189

Figures 129-150. CluMonopoma species. 129-133. Clydonopoma peasei (Pilsbry, 1933). 129, 130. ANSP 160976, holotype,

22.0 mm [photo courtesy of ANSP], 131, 132. USNM 426043, paratype, 21.4 mm. 133. USNM 426043, paratype, operculum, 7 mm

length. 134-142. Clydonoporna poloense (Bartsch, 1946). 134, 135. USNM 504040, holotype, 20.5 mm. 136. OSUM 42372, 20.4 mm.

137. UF 217969, 23.7 mm. 138. UF 217969, 21.5 mm. 139. UF 217969, 20.8 mm. 140. UF 217970, 23.0 mm. 141. Living individual,

note supemumery eye on left tentacle. 142. OSUM 42372, radula. Scale bar = 300 (jliu. 143-150. Clydonoponui pwnilwn (Watters and

Duffy, 2010). 143, 144. UF 420734, holotype, 15.0 mm. 145. GTW 71721), 19.9 mm. 146. GTW 7172c, 15.0 mm. 147. CTW 7172b,

17.3 mm. 148. UF 217798, 20.0 mm. 149. Living individual. 150. GTW 7172b, radula. Scale bar = 200 |xm.

Page 190

THE NAUTILUS, Vol 131, No. 3

Comparison with Other Species: See under Clydo-

nopoma hartschi. The large size, lamellate sculpture, thick

double lip, and peculiar coloration are unique in this

group. The lip of C. titanum is even more expanded and

hroadlv attached to the previous whorl.

Remarks: The outer lip is sinuate recalling the male shell

of some other annulariids such as Tudora. But it appears

that all individuals, male and female, have this

characteristic.

Original Description: Pfeiffer, 1852a: 252. From the

Latin. “Shell umbilieate, ovate-turret, solid, crowded longi¬

tudinal course plications, a little shining, brownish-violet; spire

elongated, conic, entire, obtuse; suture with crowded white

crenulations; remaining 7 whorls moderately convex, last

narrowly solute before, base concentrically striate; aperture

vertical, irregularly oval, inside 1 >rown; lip white, double: inner

narrowly erect, a little expanded, upper margin attached

angle, columella very si lortlv arched, outer narrowly expanded

everywhere. Operculum paucispiral. deep oblique sulcus.

Etymology: L. nobilis, noble.

Clydonopoma peasei (Pilsbry, 1933)

(Figures 1, 129-133, 182)

Chresonymy

Parachondria (. Eccritoponm ) peasei Pilsbry, 1933:

128-129, pi. 6, figs. 5-7; Wenz, 1939: 545, fig. 1453;

Richards and Old, 1969: 69; Watters, 2006: 396.

Clydonopoma (. Eccritoponm ) peasei (Pilsbry, 1933):

Bartsch, 1946: 136-137, pi. 21, fig. 2.

Licirui peasei (Pilsbry, 1933): Baker, 1964: 170.

Chondropomium peasei (Pilsbry, 1933): Watters, 2006:

63, 396.

Clydonopoma peasei (Pilsbry, 1933): Watters, 2013: 4,

map 1

Description: Shell large for family (largest adult specimen

22.0 mm maximum length, decollate, including peristome,

smallest adult specimen 21.4 mm maximum length, in¬

cluding peristome), thin, turbinoid, umbilicus wide and

open. Protoconch whorls may be retained in adults, con¬

sisting of 2 dark, microscopically granulate whorls, well-

delimited from teleoeonch. Teleoconch of 3.5 whorls,

final whorl detached just before peristome. Suture deeply

channeled. Peristome double, oval. Inner lip erect, pro¬

jecting slightly beyond the outer lip. Outer lip narrow, barely

expanded on outer side, lamellate, with weak posterior

auricle. Spiral sculpture present as ca. 6 wide, undulating

cords below suture, gradually diminishing in strength, and as

ca. 13 cords within umbilicus. Axial sculpture of numerous

narrow, fine, closely-set ribs, ca. 230 on final whorl, forming

minute, elongate cusps at suture. Suture serrate. In¬

tersections of spiral and axial sculpture finely beaded or

scalloped. Background color white or pale tan with complex

pattern of faint, brown, zigzag markings spirally aligned;

markings do not persist onto either side of lips. Early whorls

darker. Inside margin of aperture yellow. Operculum pau¬

cispiral with pseudolamella that extends Vi of way to outer

margin, remainder with pin-wheel arranged, erect, calcified

libs. Anatomy and radula unknown.

Type Material: Holotype: ANSP 160976a (as L in

Richardson et ah, 1991); Paratypes: ANSP 374376 (as PL

in Richardson et ah, 1991); USNM 426043; P: Pease coll.

Type Locality: “Sr. Del Monte’s plantation, 5 or 6 miles

west of Barahona, at Station 85, in a verdant gully near

Salvation, at about 3,000 ft." See "Type locality" under C.

clench i , above.

Type Figured: Pilsbry, 1933, pi. 6, figs. 5-7.

Distribution: Clydonopoma peasei is known only from

the type locality west of Barahona in the Sierra Baoruco.

Habitat: The holotype was collected in the same location

as C. bermudezi , see above.

Variation Among Specimens: Only type specimens

seen.

Comparison with Other Species: See under Clydono¬

poma bartsclii. If consistent, the yellow color inside the ap¬

erture is unique. It is very (similar to C. poloense found just to

the south in the same mountain range. It is less elongate and

has stronger spiral cords in the umbilicus thru does C. poloense.

Remarks: This species is known only from the type lots.

This is the type species of Eccritopoma Pilsbry, 1933,

which was differentiated primarily by the presence of

a few weak spiral grooves below the suture. We consider

this a species-level feature; Eccritopoma was synonymized

under Clydonopoma by Watters (2006). No material was

available for phylogenetic study.

Etymology: Daniel C. Pease, American Museum of

Natural History, collector of the types. Pease collected in

Hispaniola in 1932.

Clydonopoma poloense (Bartsch, 1946)

(Figures 2, 134-142, 182)

Chresonymy

Kisslingia poloensis Bartsch, 1946: 112, 114-115, pi. 17.

fig. 4; Watters, 2006: 415.

Licina poloensis (Bartsch, 1946): Watters, 2006: 75, 415;

Watters, 2012: 13; Watters, 2013: map 1.

Description: Shell large for family (largest adult

specimen 23.7 mm maximum length, decollate, in¬

cluding peristome, smallest adult specimen 18.9 mm

maximum width, including peristome), solid, conic,

umbilicus wide. Protoconch whorls often retained in

adults, consisting of 1.5 smooth tan whorls with dark

median spiral band, well-delimited from teleoconch.

Teleoconch of 4 whorls, final whorl detached just before

peristome. Suture channeled. Peristome single, thick.

Figures 151-160. Chjdonopoma species. 151-155. Chjdonopoma? subglobosum (Bartsch, 1946). 151, 152. USNM 504141.

paratvpe, 18.0 nun. 153. USNM 380218. 18.3 nun. 154, 155. USNM 380245, 19.9 nun. 156-160. Clydonopoma titanum new

species. 156, 157. OSUM 42351 , holotype, 29.1 nun. 158. GTW 16816a, 27.1 nun 159. UF 492090, paratvpe 3. 28.1 in. 160. BMSM

121709. paratvpe 4, 27.9 nun.

tear drop-shaped. Outer lip narrow, barely expanded on

outer side, auricle weak or absent. Spiral sculpture

absent except for 4-6 cords in umbilicus. Axial sculp¬

ture of numerous narrow, fine, closely-set rounded ribs,

ca. 140 on final whorl, often arranged in growth series,

forming minute, blade-like cusps at suture. Suture

serrate. Background color white or pale tan with

complex pattern of brown zigzag or chevron markings

axially aligned, sometimes forming interrupted spiral

bands on base; markings tlo not persist onto either side

of lip; umbilicus without markings. Rarely almost en¬

tirely patternless. Early teleoconcb whorls may be

colored darker brown or purplish. Operculum with

thick pseudolamella divided into two regions: outer half

of spiral with very' coarse erect lamellae, inner half

similar hut at lower level. Radula as in genus. Animal

white with pale grey foot and head patch between eyes,

tentacles pale orange.

Type Material: Holotype: USNM 504040.

Type Locality: “Near Polo, Bahoruco Mountains, Do¬

minican Republic.”

Type Figured: Bartsch, 1946, pi. 17, fig. 4.

Distribution: This species is known from the Sierra

Baoruco in the vicinity of Polo and La Cueva.

Habitat: Thi s is an upland species known only above

700 m elevation. It lives under fossil limestone blocks

in mesic, well-vegetated or forested areas and ravines.

Portions of its range are now in coffee and banana

groves.

Other Material Examined (60 Specimens): Domini¬

can Republic. Barahona Province. GTW7 16226a(2),

996 m, quarried area along Carr Los I arios el Gharco (dirt

road), 18.0965° N, -71.2516° W; OSUM 42372(9), 1 120

m, along Carr Los Linos el Gharco (dirt road) near village

of Carbon de Polla, 18.1175° N, -71 .2622° W; UF217973

(1), 910 m, 14 km S of Cabral; UF 217976(3), 1 km N of

Los Auyamas; UF 217972(1), 4 km NE of Los Auyamas;

UF 217975(2), 765 m, 2 km NNE of Polo; UF 217970(9),

990 m, 5 km NNE of Polo; UF 45604(2), 1020 m, 6 km

SSE of Polo; UF 217971(4), 910 m, 7 km NNE of Polo;

UF 217969(22), 1040 m, 5 km SE of Polo; UF 217967(3),

940 m, 9 km SF of Polo; UF 45616(2), 980 m, 7 km SSE of

Polo.

Variation Among Specimens: Specimens vary in the

strength of the background color (white to tan) and the

pattern (dark chevrons to absent).

Comparison with Other Species: See under Clydo¬

nopoma bartschi.

Page 192

THE NAUTILUS, Vol. 131, No. 3

Remarks: The holotype is a broken shell collected by

William L. Abbott between 1916-1923. The species was

not collected again until the 1970s.

Etymology: Polo, Dominican Republic.

Clydonopoma pumilum (Watters and Duffy, 2010)

(Figures 143-150, 182)

Chresonymy

Chondropomium pumilum Watters and Duffy, 2010:

8-10, figs. 27-30; Watters, 2012: 14, figs. 49, 54 [ani¬

mal]; Watters, 2013: 4, pi. 2, fig. 15, map 1.

Description: Shell medium for family (largest adult

specimen 20.0 nun length, decollate, including peristome;

smallest adult specimen 14.2 nun maximum length, de¬

collate, including peristome), solid, conic, umbilicus wide.

Protoconch whorls unknown, decollate in adults. Tele-

oconch of 4.25-4.5 whorls, final whorl adnate except for

immediately behind peristome. Suture channeled. Peri¬

stome double, thick, tear drop-shaped. Outer lip widely

expanded but narrowest facing umbilicus, scalloped on

anterior margin, produced into prominent, concave au¬

ricle posteriorly. Inner lip very short and erect. Spiral

sculpture present only as few feeble cords in umbilicus.

Axial sculpture of numerous closely spaced, thin, low

lamellae, occasionally anastomosing. Axial lamellae

slightly elongated at suture into blade-like cusps. Back¬

ground color tan or brownish purple with diffuse narrow,

spiral bands brown bands. Outer lip white. Operculum

paucispiral with pseudolamella. Animal with grev foot,

snout, and base of tentacles; bead white with dark grey

mottling between tentacles; distal portion of tentacles

orange. Radula with broad, triangular inner marginal

teeth with obsolete, minute denticles; rachidian and lat¬

eral teeth unicuspid and very elongate.

Type Material: Holotype: UF 420734; Paratypes:

OSUM 32484(1); NHMUK 1996350(1).

Type Locality: “Dominican Republic, Barahona, Ped-

ernales Province, 19-32 km N of Cabo Rojo,

-500-900 m.”

Type Figured: Watters and Duffy, 2010, figs. 28, 29.

Distribution: Known only from ca. 400-1500 m eleva¬

tion on the western face of the Sierra de Baoruco.

Habitat: Under limestone rocks in upland coniferous

forests; some areas burnt over.

Other Material Examined (98 Specimens): Domini¬

can Republic. Pedemales Province. GTW 7172a(l),

760 m, 21-24 km N of Cabo Rojo; GTW 7172b(9), 1300

in, on road to Las M ereedes, 21 km NE of Pedemales; UF

217794(12), 1080 m, 12 km N of Las Mercedes; UF

217793(12), 1510 m, 20 km N of Las Mercedes; GTW

7 1 72e(l ). 1 285 m, abandoned quarry near end of road that

runs from Cabo Rojo N past Las Mercedes into Sierra de

Baoruco, 18.1203° N, -71.5727° W; UF 217798(18), 1410 m.

Dana El Aceitillar, 12 km FNF of Las Mercedes; UF 217799

(12), UF 217801(8), 1300 m, Loma El Aceitillar, 19 km FNF

of Las Mercedes; UF 21 7797(6), 39 road km N of Cabo Rojo;

UF 217800(3), 1000 m, W rim of Hoyo de Pelempito; UF

217795(14), UF 217796(2), 380 m, 3 km N of Nigueron [not

found].

Variation Among Specimens: Specimens differ pri¬

marily in the background color, varying from nearly

uniform white to dark purplish brown.

Comparison with Other Species: See under Cli/do-

nopoma hartschi.

Etymology: L. pumilum , dwarfish, a miniature C. nobile.

Clydonopoma? subglobosum (Bartsch, 1946)

(Figures 151-155).

Chresonymy

I ncertipoma subglobosum Bartsch, 1946: 1 71-172, pi. 29,

fig. 6; Watters, 2006: 495.

Chondropomium subglobosum (Bartsch, 1946): Watters,

2006: 63, 495.

Description: Shell large for family (largest adult speci¬

men 22.0 mm length, decollate, including peristome;

smallest adult specimen 18.3 mm maximum length, de¬

collate, including peristome), solid, turbinoid, umbilicus

wade, polished. Protoconch whorls of 2 brown, smooth

whorls, usually decollate in adults. Teleoconch of 4-4.5

whorls, final whorl adnate except for immediately behind

peristome. Suture indented but not channeled. Peristome

single, teardrop-shaped. Lip scarcely expanded, posterior

auricle lacking. Spiral sculpture present only as single

thread in umbilicus. Axial sculpture of numerous closely

spaced, flat cords on early whorls, obsolete by final whorl.

Suture smooth on final whorl. Background color white

patterned with brown spiral bands of dots alternating with

3 spiral bands of triangular browm markings. Lip white.

Operculum, anatomy, and radula unknown.

Type Material: H: USNM 504141.

Type Locality: “Trou Louise, Gonave Island. ' This is the

place now known as Platon Trou Louis.

Type Figured: Bartsch, 1946, pi. 29, fig. 6.

Distribution: Originally recorded from lie de la Gonave

and Petit-Goave. Bartsch (1946) listed Trou Louise, South

Abrieots, and Point Fantasque on lie de la Gonave but

only the first locality has been located by us on that island.

There is a Point Fantasque at the entrance to Cavernites

Bav cm tin' mainland on the north shore of the Tiburon

Peninsula across from lie de la Gonave (Wyman, 1874).

It is probable that Bartsch misloeated Point Fantasque

on lie de la Gonave. Petit-Goave is a town also on the

north shore of the peninsula. If our localities are correct

then this species lives on lie de la Gonave and on the

G.T. Watters and P. Larson, 2017

Page 193

Figures 161-181. Superbipoma species. 161-171. Superbipoina asijmmetricum (Pilsbry, 1933). 161. ANSP 46656, lectotype,

18.2 mm [photo courtesy of ANSP]. 162. Chondroporna enriquillense Pilsbry, 1933. ANSP 146715, holotype, 23.8 mm [photo courtesy

of ANSP]. 163, 164. USNM 471939, 18.6 mm. 165. UF 216851, 25.8 mm. 166. UF 216851, 25.7 mm. 167, 168. OSUM 42375, 26.9

mm. 169. USNM 471937, 25.7 mm. 170. Living individual. 171. OSUM 42364, radula. Scale bar = 200 pan. 172-181. Superbipoma

superburn (Henderson and Simpson, 1902). 172, 173. USNM 168798, holotype, 27.6 mm. 174, 175. UF 119165, 24.8 mm. 176. UF

216852, 28,5 mm 177. UF 216854, 22.6 mm. 178. CTW 7086b, 23.8 mm. 179, 180. GTW 7086b, 25.5 mm. 181. GTW 7086d, radula.

Scale bar = 100 pm.

Page 194

THE NAUTILUS, Vol. 131, No. 3

Figures 182-185. Distribution maps. All maps Google™ Earth Pro. Image Landsat. © 2017 Google. Data: SIO, NOAA, US Navy,

NGA, GEBCO. Inset - general area covered. Not shown - Chnndropomium ignotum, ClydonopomM subglobosum, see text. 182.

Chondropomium blaineorum - yellow; Chondropomium lynx - purple; Cltjdonopoma peasei - blue; Clydonopoma poloense - green:

Clijdonopoma pumilum - white; Clydonopoma titanum - red square; Chondropomium sardonyx - red. 183. Clydonopoma bahorucense -

purple; Clydonopoma bartschi - red; Chondropomium beatense - blue; Chondropomella magnified - white; Clydonopoma nobile -

yellow; Chondropomella plat y chilli m - green. 184. Chondropomium asymmetricum - green; Chondropomium caelicum - blue;

Clydonopoma bennudezi — yellow; Chondropomium gimhiense - purple; Chondropomium marmoreum - white; Chondropomium

superbum - red. 1 85. Chondropomium weinlandi . “ azuense form” - white; barahonense form - red; “ weinlandi form” - green. Some

dots overlap.

northern shore of the Tiburon Peninsula opposite that

island.

Habitat: Not reported but lie de la Gonave is known to be

xerie (an ISIS map labeled it as "sterile et inhabitee”). No

live specimens have been recorded. The label to USNM

499359 reads "Point Fantasque... top of first ridge from

shore.”

Other Material Examined (23 Specimens): Haiti.

USNM 380245(14), Trou Louise, lie de la Gonave; USNM

380218(1), USNM 504142(1), lie de la Gonave; USNM

499359(1), Point Fantasque; USNM 504143(6), Petit-

Goave.

Variation Among Specimens: Specimens differ in the

strength of the brown patterns with some specimens

being almost uniformly white.

Comparison with Other Species: The large, polished,

turbinoid shell, usually distinctly marked with brown on a

white background, is unique in all Hispaniolan annulariids.

Remarks: This species is placed in Clydonopoma with

reservations. No live specimens have been collected and

the operculum is unknown. It closely resembles other

Clydonopoma and Chondropomium in sculpture and

color patterns. The addition of this species would greatly

increase the range of either genus to the west. Only the

discovery of the operculum or live material for phylo¬

genetic study will determine its proper place.

Etymology: L. sub- somewhat + L. globosum, round.

Clydonopoma titanum new species

(Figures 156-160, 182)

G.T. Watters and P. Larson, 2017

Page 195

Description: Shell very large for family (largest adult

specimen 29. 1 mm length, decollate, including peristome;

smallest adult specimen 27.5 nun maximum length, de-

collate, including peristome), solid, conic, umbilicus wide,

surface silky. Protoconch whorls unknown, decollate in

adults. Teleoconch of 3. 5-4.0 whorls, final whorl adnate

except for immediately behind peristome. Suture nar¬

rowly but deeply channeled. Peristome single, thick,

somewhat reflected, tear drop-shaped, widely expanded

but narrowest facing umbilicus, composed of numerous

irregular fused lamellae, with or without low auricle

posteriorly. Spiral sculpture present as numerous (ea. 45)

minute, low feeble threads, most evident ;is scallops on axial

sculpture, more widely spaced in umbilicus. Axial sculpture

of numerous (ea. 250) closely spaced, minute threads,

gathered in groups of 10-20 between growth lines. Axial

lamellae slightly elongated at suture into minute, blade-like

cusps, rendering suture finely serrate, no fused tufts.

Background color pale tan, orangish, or brownish purple,

base paler. Pattern indistinct or bold, of spiral brown bands

of "> ’’-shaped markings and minute dots. Lip usually white,

unpattemed. Operculum paueispiral with pseudolamella;

pseudolamella covers proximal 75% of opercular whorls

exposing underlying comeus base on outer edge. Animal

pale cream colored; head with dark grey mottling on top of

snout; distill portion of tentacles orange. Radula unknown.

Type Material: Holotype: OSUM 42351. 29.1 nun;

Paratype 1: OSUM 42352, 27.7 nun; Paratvpe 2: OSUM

42353, 27.5 mm; Paratype 3: UF 492090, 28. 1 mm;

Paratype 4: BMSM 121709, 27.9 mm.

Type Locality: 775 m elevation, along Rte. 204, ea. 1 km

N of Decouze, Departement d’Ouest, Haiti, 18.3414° N,

-72.5836° W.

Distribution: Known only from the type locality.

Habitat: Specimens were found on rock piles by the side

of the road at ea. 775 m elevation.

Other Material Examined (5 Specimens): Haiti.

Departement d’Ouest. GTW 16816a(5), from the type

locality.

Variation Among Specimens: Specimens vary in degree

of color pattern and background color, which ranges from

pale tan to orangish and dark purple-brown.

Comparison with Other Species: This species differs

from the similar C. nobile and C. poloense in lacking the

channeled suture and in having a much more broadly

expanded outer lip.

Remarks: This species shares as much in common with

taxa such as Licina habichi (Weinland, 1862) from

western-most Haiti as it does with other Clydonopoma .

Licina species are rare and collections currently lack

preserved material for phylogenetic work, but future

studies may reassign many of the species now placed in

these two genera. We are indebted to Jose Coltro (Brazil)

for calling our attention to this species.

Etymology: G. Titan , in reference to its size.

Genus Superbipoma new genus

Type Species: Chondroponui snperbwn Henderson

and Simpson, 1902.

Description: Shells large to very large for family (to ea.

35 mm length), elongate-conic, usually decollate. Proto¬

conch of 1 .5 smooth, minute whorls. Final whorl detached

from previous whorl for V4-V2 turn, forming sharp pos¬

terior keel. Axial sculpture of wide, flattened, close-set ribs

separated by incised grooves. Spiral sculpture absent on

final whorl except present in umbilicus. Overall sculpture

appears veiy smooth, often polished. Suture minutely

beaded but lacks fused tufts, deeply channeled. Adult with

reflected lip. Lip single or double, but if double inner lip is

usually fused to outer lip. Color pattern of bands and

blotches, continuous or interrupted; rarely lacking any

pattern. Operculum paueispiral with uniform, thin granular

deposit. Taenioglossate radula with rachidian tooth, single

pair of lateral teeth, and two pairs of marginal teeth.

Rachidian and lateral teeth usually unicuspid. Inner marginal

deeply multicuspid on outer side. Outer marginal pectinate.

Animal with foot longitudinally bisected into lobes. Ixico-

motion ditaxie between lobes of foot. Eyes at base of ten¬

tacles; bifid snout produced into short secondary tentacles.

Distribution: Rift valley of Hoya de Enriquillo in the

Dominican Republic and the Plain du Cul-de-Sae in Haiti.

Remarks: This genus differs from Clydonopoma and

Chondropomella in lacking a pseudolamellate operculum and

in having deeply incised inner marginal radular teeth. It

differs from Chondropomium in the expanded lip and greater

adult size. The genus is primarily recognized as distinct based

on phylogenetic evidence. The included Superbipoma spe¬

cies form a group sister to the clade consisting of Clt/do-

noponui and Chond ropom in in. Although this position is not

strongly supported (Figure 5), it was recovered in all twenty

of the independentlv run analyses and the monophyly of

these three genera is strongly supported.

Superbipoma asymmetricum (Pilsbry, 1933)

(Figures 161-171, 184)

Chbesonvmy

Chondropoma (Chondropomella) asymmetricum Pilsbry,

1933: 126-127, pi. 9, figs. 4-5.

Chondropoma ( Chond mpomella ) enriquillense Pilsbry,

1933: 127, pi. 9, fig. 1; Bartsch, 1946: 31 [in synon¬

ymy of Chondropoma asymmetricum Pilsbry, 1933].

Chondropoma (Chondropomium) asymmetricum Pilsbry,

1933: Bartsch, 1946: 31-32, pi. 6, fig. 1.

Chondropoma (Chond ropomium) inaequilabnim

Bartsch, 1946: 32-33, pi. 5, fig. 6.

Chondropoma asymmetricum Pilsbry, 1933: Baker,

1964: 169.

Chondroponui enriquillense Pilsbry, 1933: Baker, 1964: 169.

Page 196

THE NAUTILUS, Vol. 131, No. 3

Chondropomium asymmetricum (Pilsbry, 1933): Watters,

2006: 62, 152.

Chondropomium inaequilahrum (Bartsch, 1946):

Watters, 2006: 63, 304-305.

Chondropomium enriquillense Pilsbry, 1933: Watters,

2006: 250.

Chondropomella asymmetricum (Pilsbry, 1933): Watters

and Duffy, 2010: 1 1 .

Chondropomella inaequilatnim (Bartsch, 1946): Watters

and Duffy, 2010: 11.

Description: Shell very large for family (largest adult

specimen 33.5 mm maximum length, decollate, including

peristome; smallest adult specimen 20.8 mm length, de¬

collate'. including peristome), solid, polished, conic, umbilicus

open, spire ca. Vi total length. Protoconch of 1.5 minute,

smooth whorls, white with wide tan, central hand; protoconch

rarely retained in adults. Decollate specimens have jagged,

irregular break from earlier whorls, which seems to involve

more than just protoconch whorls. Teleoeonch ot 3.5-3.75

whorls, final whorl distinctly detached for last I4tli of whorl

and deflected anteriorly. Suture narrow, deeply channeled.

Peristome double, tear drop-shaped. Inner lip not erect, fused

to outer lip. Outer lip expanded, narrowest facing umbilicus,

non-auriculate, distinctly detached from previous whorl,

forming sharp posterior keel. Spiral sculpture absent except

for few very weak, almost obsolete cords within umbilicus.

Axial sculpture of flattened, low ribs, separated by incised

grooves, ca. 200-250 ribs on final whorl, each forming minute

bead at suture; no fused tufts present. Background color very

variable, from white to tan, with axial zig-zag brown markings

arranged in vague spiral bands; umbilicus white. Outer lip

usually white, rarely banded. Shell rarely uniformly white.

Operculum paucispiral with thin calcareous deposit. Radula as

in genus. Animal uniformly p:de tan or white with dark band at

base of snout.

Type Material: Chondropoma asymmetricum Pilsbry,

1933: Lectotype: ANSP 46656; Paralectotypes: ANSP

373761(4); USNM 426041(1). Chondropoma enri-

(piillense Pilsbry, 1933: Holotype: ANSP 146715. Chon¬

dropomium inaequilahrum (Bartsch, 1946): Holotype:

USNM 471939.

Type Locality: Chondropoma asymmetricum Pilsbry,

1933: “Fond Parisien, on the south shore of Etang

Saumatre, Haiti." Chondropoma enriquillense Pilsbry,

1933: “Lake Enriquillo, Santo Domingo." Collected by

Bond... James Bond in 1928. Chondropomium inaequi¬

lahrum (Bartsch, 1946): “Mount Petitchemin, De¬

partment de l’Ouest, Haiti.

T\pe Figured: Chondropoma asymmetricum Pilsbry,

1933: Pilsbry, 1933, pi. 9, figs. 4, 5. Chondropoma enri¬

quillense Pilsbry, 1933: Pilsbry, 1933, pi. 9, fig. I

Chondropomium inaequilahrum (Bartsch, 1946): Bartsch,

1946, pi. 5, fig. 6.

Distribution: Found below 500 m elevation in the

northern foothills of the Sierra Baoruco in the rift valley

of Hoya de Enriquillo/Plain du Cul-de-Sae facing the

south shores of Lago Enriquillo and Etang Saumatre.

Not recorded from the northern portion of the valley,

where it is replaced by S. superhum. Chondropoma

inaequilahrum Bartsch, 1946, was described from the

Haitian Department de l’Ouest from Mount Petitch¬

emin. This location is not listed among the 651 peaks in

Haiti by www.peakerv.com and we have not been able

to locate it. The Department de l’Ouest includes the

Plain tin Cul-de-Sac and, like most species in the genus,

this taxon probably originated from that rift valley as

well.

Habitat: Under fossilized coral nibble in xeric areas with

agave and cacti. Often common.

Other Material Examined (265 Specimens): Haiti.

Departement d'Ouest. USNM 471937(1), Fond Parisien;

UF 32148(20), UF 32149(25), 4 km S of Fond Parisien; UF

32162(21), 10km SSE of Fond Parisien; USNM 426041(1), S

shore Etang Saumatre; Dominican Republic. Inde-

pendeneia Province. OSUM 42369(15), 466 m. 1.3 km E of

center of Puerto Escondido, 18.3290° N, -71.5593° W;

OSUM 42357(9), avocado grove, just outside Puerto

Escondido; UF 216858b(24), 430 m, 8 km WML of Puerto

Escondido; UF 216859(13), 490 m, 11 km WNW of Puerto

Escondido; UF 216651(3), 275 m, 12 km WNW of Puerto

Escondido; OSUM 42370(42), 473 m, along RD 541, 2,3 km

NNW of El Naranjo, 18.3514° N, -71.6279° W; OSUM

42375(20), 224 m, hill top between Jimanf and El Limon, off

RD 46, 18.4504° N, -71.8115° W; UF 216851a(12), 210 m,

5km SW of El Limon; UF 216857(15), 40 m, 2 km NWof'El

Limon; UF 216652(1), 6.2 km SE of Jimanf; UF 216855a(29),

UF 216862b(25), 7 km SE of Jimanf; GTW 16218d(9), ridge

NW of Jimanf; GTW 16023a(5), 1.2-2 km E of Jimanf.

Variation Among Specimens: Specimens vary in the

degree of the outer lip expansion and the depth of the

color pattern, which ranges from nearly all white to heavily

mottled with brown.

Comparison with Other Species: This species differs

from the similar S. superhum in being less elongate and in

having the color pattern consist of axially arranged zig-zags,

whereas in S. superhum the pattern is of interrupted spiral

bands.

Remarks: Watters and Duffy (2010) placed this species in

Chondropomella. However, it lacks the highly modified,

calcareous operculum of that genus. Chondropomium

inaequilahrum (Bartsch, 1946) was differentiated from

C. asymmetricum only by size but it falls well within the

variation of this species.

Etymology: Chondropoma asymmetricum Pilsbry, 1933:

L. asymmetricus , asymmetrical [outer lip]. Chon¬

dropoma enriquillense Pilsbry, 1933: Lago Enriquillo,

Dominican Republic. Chondropomium inaequilahrum

(Bartsch, 1946): L. inaequi-, unequal + L. lahrum , lip.

The outer lip is unequally expanded around the

aperture.

G.T. Watters and P. Larson, 2017

Page 197

Superbipoma superbum (Henderson and Simpson,

1902)

(Figures 172-181, 184)

Chresowmy

Chondroponui superbum Henderson and Simpson, 1902:

88-89, text, fig.; Parodizand Tripp, 1988: 149; MeChie,

2008: 31.

Chondroponui ( Chondropomium ) weinlandi superha

Henderson and Simpson, 1902. Henderson and

Bartseh, 1920: 60.

Chondroponui ( Chondropomium ) superhum Henderson

and Simpson, 1902: Clench and Aguayo, 1937: 64;

Bartseh, 1946: 23-24, pi 5 fig. 1.

Chondropomium superhum (Henderson and Simpson,

1902): Watters, 2006: 63, 498.

Description: Shell large for family (largest adult

specimen 29.3 mm maximum length, decollate, in¬

cluding peristome; smallest adult specimen 20.1 mm

length, decollate, including peristome), polished, al¬

most translucent, conic to fairly high-spired, umbilicus

open, spire 14—1/3 total length. Protoconch of 1.5

minute, smooth, white whorls, decollated in adults.

Decollate specimens have jagged, irregular break from

earlier whorls, which seems to involve more than just

protoconch whorls. Teleoconch of 3.25-4.25 whorls,

final whorl distinctly detached for last 1 /4th of whorl

and deflected anteriorly. Suture narrow, deeplv

channeled. Peristome single, tear drop-shaped. Lip

expanded, narrowest facing umbilicus, with minute

triangular posterior auricle, distinctly detached from

previous whorl, forming sharp posterior keel. Spiral

sculpture absent except for few very weak, almost ob¬

solete cords within umbilicus. Axial sculpture of flat¬

tened, low ribs, separated by incised grooves, often

grouped in growth series, ea. 160 ribs on final whorl,

each forming minute bead at suture; no fused tufts

present. Background color very variable, from white to

tan to purplish brown, early whorls often darker; axial

spots and dashes axially aligned on trailing edge of

growth series, rarely without marks, rarely with contin¬

uous or nearly so spiral bands; lip and umbilicus white.

Operculum paucispiral with thin calcareous deposit.

Radula as in genus.

Type Material: Holotype: USNM 168798; Paratypes:

ANSP 386772(44); UF 119090(2); UF 119165(4); UF

119169(6); UF 22229(33), UF 22230(4); Manchester

Museum (UK); EE. 3688(4), listed as syntypes, see

McGhie (2008); CM Ml 62.38661(6), listed as syntypes.

Type Locality: “On a high limestone hill hack of Tho-

mazeau, Haiti.”

Type Figured: Henderson and Simpson, 1902, text fig.

Distribution: Th roughout the Hoya de Enriquillo/Plain

du Cul-de-Sac from north of Port-au-Prince to the NW

shores of Lago Enriquillo. It occurs at and below sea level

(Lago Enriquillo is below sea level), rarely to ca. 600 m. It

has not been found along the southern shore of Lago

Enriquillo where it is replaced by S', asymmetricum.

Locally common.

Habitat: Under fossilized coralline debris and in rock

piles in xeric areas with agave and cacti. Locally common.

Co-occurs with C. weinlandi , C. nuignificnm (Pfeiffer,

1854), and Crosseponui vermiculatum sd. (Bartseh,

1946).

Other Material Examined (406 Specimens): Haiti.

Departement de I’Ouest. UF 33813(16), 14 km SF, of

Duvalierville [Cabaret]; UF 31983(2), 18 km SE of

Duvalierville [Cabaret]; UF 32090(6), 350 m. I 1 km NNE

of Beudet; UF 1 19090(2), UF 1 19165(4); UF 119169(6),

UF 183694(44), UF 22229(33), UF 22230(4), Thoma-

zeau; UF 32559(7), 2 km E of Thomazeau; UF 32565(25),

UF 32726(1 ), 50 m, 5 km W of Thomazeau; UF 32041 ( 1 ),

24 km SSW of Mirebalais; UF 32052(5), 25 km SSW of

Mirebalais. Departement du Centre. GTW 7086f(7),

632 m. Fond Cavalier, 18.7164° N, -72.1658° W. Do¬

minican Republic. Independencia Province. UF

216860(5), 2 km E of La Lajas; UF 216852(44), 6 km SE

of La Lajas; GTW 70S6a(l),OSUM 42360(9), Bartolome,

N shore of Lake Enriquillo; GTW 7087i(l), Boca de

Cachdn; GTW 70S6b(34), old quarry, off HD 48, 1.8 km

ESE of Boca de Cachdn, 18.5514° N, -71.8232° W; UF

216853(35), UF 216856(13), 7 km NW of Boca de

Cachdn; GTW 7086c(7), 4.7 km S of Boca de Cachdn!

18.5129° N, -71.8440° W; GTW 7086d(16), Boca de

Cachdn; UF 216854(7), 2 km NNW of La Deseubierta;

UF 216849(37), 250 m, 6 km NE of La Deseubierta; UF

216826a(19), 7 km E of La Deseubierta; UF 216825(18),

6 km NW of Postrer Rio.

Variation Among Specimens: This species is much

more varied in color than described by either Henderson

and Simpson (1922) or Bartseh (1946). The degree of

background color ranges from nearly all white to nearly all

brown or purplish-brown; the strength of the axial

markings varies from small spots or dashes to nearly

complete spiral bands.

Comparison with Other Species: See under S.

asymmetricum.

Etymology: L. superhum, superb.

DISCUSSION

The relationships within and among annulariid species

groups are more fully understood when mutually in¬

formed by morphology and phylogeny. While Chon¬

dropomium and Superhipoma share features that have

been historically used to define genera, the well-

supported position of the interloping Clydonoponui

species group suggest that sharing these features does not

necessarily indicate a close relationship. Likewise, while

Page 198

THE NAUTILUS, Vol. 131, No. 3

Table 1. Specimens used in phylogenetic analyses and associated GenBank accession numbers for each sequence. New sequences

generated for this study are in boldface. DR = Dominican Republic.

branch lengths between genera (e.g., Clydonopoma and

Chondropomium ) are similar to or shorter than some of

those within a genus (i.e. Superbipoma), these same

characters provide justification for not combining all three

genera into a single large taxon. At the species level, our

phylogenetic results have implications for putative

subspecies or morphotypes. In particular, the two in¬

dividuals of Chondropomium weinlandi “azuense form”

highly supported relationships with two individuals of

different types (i.e., with C. weinlandi with 99% bootstrap

support and with C. weinlandi “barahonen.se" with 94%

bootstrap support). Similarly, C. weinlandi “ barahonen.se ”

G.T. Watters and P. Larson, 2017

Page 199

individuals are not most closely related, with high support.

These results are consistent with a single polymorphic

species rather than multiple separate species or subspecies.

Although Clydonopoma , Chondropomella , and other

Tiburon genera have a heavily calcified and complex

pseudolamellate operculum, this has been reduced to a fine

granular deposit in Chondropomium and Superbipoma.

This follows the findings of Skomrock (2014), which sug¬

gested that a calcified operculum was the ancestral con¬

dition of the Annulariidae. Taxa lacking calcification have

secondarily lost this feature and are representative of

a derived condition. It appears that the pseudolamella was

lost in Chondropomium and Superbipoma from a pseudo-

lamellate ancestor. Both of these genera are lowland forms

but it is not clear how (or if) the loss of the pseudolamella is

associated with that change in habitat.

Crossepoma venniculatum (Bartseh, 1946) is con-

cbologicallv verv similar to Chondropomium and is often

sympatric with C. weinlandi. Described as a Chon¬

dropomium, Watters (2012) moved it to Crossepoma

Bartseh, 1946, based on the structure of the operculum.

Although Crossepoma is not reviewed here, C. vennic¬

ulatum is confirmed as belonging to a genus apart from

Chondropomium. However, C. venniculatum is not the

tvpe species of the genus Crossepoma (= Chondropoma

emilianum Weinland, 1862) and additional work is needed

to determine the relationship of Crossepoma to

Chondropomium.

It is of interest to note which groups have “broken out "

of the Tiburon Peninsula to begin colonization of the

remainder of Hispaniola since the Miocene. Clydono-

poma is apparently limited to the mesie uplands of the

Sierra de Baoruco and may be ecologically restricted from

establishing itself in other mountain ranges by virtue of

the intervening xeric lowlands. Chondropomium, Chon¬

dropomella, and Superbipoma. on the other hand, prefer

these xeric lowland habitats and have spread up the rift

valley and connecting river valleys (in Chondropomium )

into adjacent Hispaniola, all the while remaining lowland

forms. Only C. caelicum , which has advanced the furthest

into Hispaniola, has colonized a more montane habitat

This study confirms the narrow endemism of some

Tiburon genera and probably is representative of annu-

lariid taxa in general. It also demonstrates the utility of the

phylogenetic study of this group to determine not only

genus-level patterns but also patterns within a species.

Unfortunately, the great majority of specimens in collections

consist of material unsuitable for phylogenetic work, in¬

cluding this study. Coupled with the endemism of most taxa,

future phylogenetic work on the group would require ex¬

tensive resampling on a fine scale that may not be possible

due to time, financial, and political constraints, as well ;is the

possibility of taxa having become extirpated or even extinct.

ACKNOWLEDCM ENTS

The author thanks G. Paulay, J. Slapcinsky, and the late F.

Thompson (UF) and B. Hersehler (USNM) for access to

their invaluable collections. E. Neubert at the Natur-

historisehe Museum, Bern, shared insights on the swifti/

weinlandi problem. Z. Feher (Hungary), S. Aiken (UK),

A. Gettleman (FL), H. Rhode (FL), G. Duffy (FL), L. and

J. Kremer (IF), R Hooks (Dominican Republic), M. and

D. Blaine (DE), and |. Coltro (Brazil) generously donated

valuable material used in this review. |. Ablett (NHMUK)

and C. Zorn (ZMB) spent considerable time attempting to

locate type specimens in those museums. B. Kemminoe

(OSU) assisted with SEM preparations. Molecular work

was carried out on the equipment and in the lab space of

M. Daly (Department of Evolution, Ecology and Or-

ganismal Biology, OSU). The manuscript benefited from

comments of two reviewers.

LITERATURE CITED

Adams, 11 and A. Adams. 1854 1858. The genera of Recent

Mollusca, arranged according to their organization, 2. J. Van

Voorst, London. 661 pp. Parts 16-18: pp. 1-92, pis. 61-72

[1854]; Parts 19-24: pp. 93-284, pis. 73-96 [1855]; Parts

25-28: pp. 285-412, pis. 97-112 [1856]; Parts 29-32, pp.

413-540, pis. 113-128 [1857]; Parts 33-36: pp. 541-661,

pis. 129-138 [1858],

Altschul, S.F., W. Gish, W. Miller, E.W. Myers and D.J. Lipman.

1990. Basic local alignment search tool Journal of Mo¬

lecular Biology 215: 403^410.

Arango y Molina, R. 1878. Contribution a la fauna malaeologica

Cubana. G. Montiel y Comp., Habana. 280+35 pp.

Baker, H.B. 1941. New Puerto Rican land snails. Notulae

Naturae (88): 1-6.

Baker, H.B. 1964. Type land snails in the Academy of Natural

Sciences of Philadelphia. Part III. Limnophile and tha-

lassophile Pulmonata. Part IN'. Land and (resh-water Pro-

sobranehia. Proceedings of the Academy of Natural

Sciences 116: 149-193.

Bartseh, P. 1932. A newly discovered West Indian mollusk

faunula. Proceedings of the U.S. National Museum 81(6):

1-12, pis. 1-3.

Bartseh, P. 1946. The operculate land mollusks of the family

Annulariidae of the island ol Hispaniola and the Bahama

Archipelago. Bulletin of the U.S. National Museum 192:

264 pp., 38 pis.

Bland, T. 1861. On the geographical distribution of the genera

and species of land shells ol the West India Islands; with

a catalogue of the species of each island. Annals ol the

Lyceum of Natural History of New York 7: 335-361 .

Boyko, C.B. and J.R. Gordeiro. 2001. Catalogue of Recent t\pe

specimens in the Division of Invertebrate Zoology. V.

Mollusca, Part 2. (Class Gastropoda [exclusive of Opis-

thobranchia and Pulmonata], with supplements to Gas¬

tropoda [Opisthobranchia], and Bivalvia). Bulletin of the

American Museum ol Natural History (262): 170 pp.

Clench, W.J. 1932. Some land mollusks from Beata Island, Santo

Domingo. Proceedings of the New England Zoological

Club 12: 103-107.

Clench, W.J. 1938. Robert Swift. The Nautilus 51: 142 143.

Clench, W.J. and C.G. Aguayo. 1948. Un nuevo Chondropoma

de la provincia de Oriente, Cuba. Revista de la Sociedad

Malaeologica “Carlos de la Torre” 6(2): 53-54.

Crosse, H. 1890. Faune malacologique terrestre et fluviatile de

file de Cuba. Journal de Conchyliologie 38: 173-335.

Page 200

THE NAUTILUS, Vol 131, No, 3

Crosse, H. 1891. Faune malacologique terrestre et fluviatile de

Pile de Saint-Domingue. Journal de Conehyliologie 39:

73-210.

Dali, W.H. and C.T. Simpson. 1901. The Mollusca of Porto Rico.

Bulletin ol the U.S. Fish Commission 20: 351—524.

Edgar, R.C. 2004. MUSCLE: multiple sequence alignment with

high accuracy and high throughput. Nucleic Acids Research

32: 1792-1797.

Folmer, O.. M. Black. W. lloeh, R. Lutz and R. Vrijenhoek.

1994. DNA primers for amplification of mitochondria]

cytochrome e oxidase subunit I from diverse metazoan

invertebrates. Molecular Marine Biology and Bio-

teclmolog)' 3: 294-299.

Cray. |.E. 1850. Nomenclature of molluscous animals and shells

in the collection of the British Museum. Part I Cyclo-

phoridiE. British Museum, London. 69 pp. + index.

Henderson, |.B. and P. Bartsch, 1920. A classification of the

American operculate land mollusks of the familv Annu-

lariidae. Proceedings ol the U.S. National Museum 58:

49-82.

Henderson, J.B. and C.T. Simpson. 1902. A new Haitian

Chondrnponui. The Nautilus 16: 88-89.

Henderson, R.W. and R. Powell. 2004. Thomas Barbour and the

Utowana voyages (1929-1934) in the West Indies. Bonner

zoologische Beitrage 52: 297-309.

Jaume, M L 1984. Rectificaciones a nombres de molluscos

terrestres. Miscelanea Zoologica, Academia de Ciencias de

Cuba (20): 1-4.

Kobelt, W. 1880. Die geographische Verbreitung der Mol-

lusken. Jahrbiicher der Deutschen Malakozoologischen

Gesellschaft nebst Nachrichtsblatt 7: 241-286.

Machida, R.J., M. Kweskin and N. Knowlton. 2012. PCR

primers for metazoan mitochondria] 12S ribosomal DNA

sequences. PLoS ONE 7, e35887.

M cGhie, H.A. 2008. Catalogue of the type specimens of mol¬

luscs in the collection of the Manchester Museum, the

University’ of Manchester, UK. ZooKeys 4: 1 — 46.

Mermod, G. 1952. Les hpes de la collection Lamarck au

M useum de Geneve. Mollusques vivants, III. Revue Suisse

de Zoologie 59(2): 24-97.

Miller, J.Y. and L.D. Miller. 2001. The biogeography of the West

Indian butterflies (Lepidoptera): an application of a vicari-

ance/dispersalist model. In: Woods, C.A. and F.E. Sergile

(eds.). Biogeography of the West Indies. 2nd ed. CRC Press,

Boca Raton, EL: 127-155.

Neubert, E. and M. Gosteli. 2005. The moUuscan species de¬

scribed by Robert James Slmttleworth 11. Polyplacophora,

Gastropoda (Caenogastropoda), Bivahia. Contributions to

Natural History, Scientific Papers from the Natural History

Museum Bern (5): 1-79, 20 pis.

Parodiz, J.J. and | J. Tripp. 1988. Types of Mollusca in the

collection of the Carnegie Museum of Natural History.

Part 1. Bivahia and Gastropoda (Prosobranchia and

Opisthobranchia). Annals of the Carnegie Museum 57(5):

111-154.

Pfeiffer, L. 1846. Die gedeckelten Lungenschnecken. (Heli-

cinacea et Cyclostomacea). Systematisches Conchylien-

Cabinet v'on Martini und Chemnitz (Krister ed.). Part 19.

Installment 61 : pis. 1-3, 5-7. pp. 1-24.

Pfeiffer, L. 1847. Die gedeckelten Lungenschnecken. (Heli-

cinaeea et Cyclostomacea). Systematisches Conchylien-

Cabinet von Martini und Chemnitz (Krister ed.). Part 19.

Installment 64: pis. 8, 10, 12-14, 17, pp. 25-40.

Pfeiffer, L. 1850. Beschreibungen neuer Landschnecken.

Zeitschrift frir Malakozoologie 7(5): 65-80.

Pfeiffer, L. 1851. Conspectus emendatus generum et specierum

Cyclostomaceoru i n . Zeitschrift fur Malakozoologie 8(9):

129-144: 8(10): 145-160; 8(11): 161-176; 8(12): 177-178.

Pfeiffer, L. 1852a. Monographia pneumonoporum viventum. T.

Fischer, Cassellis. 435 pp.

Pfeiffer, L. 18521). Conspectus cyclostomaceoru m emendatus et

auctus. Pneumonopomorum monographie prodromus. T.

Fischer, Cassellis. 50 pp., addenda.

Pfeiffer, L. 1853a. Catalogue of Phaneropneumona, or terres¬

trial operculated Mollusca, in the British Museum. British

M useum, London. 324 pp.

Pfeiffer. L. 1853b. Die gedeckelten Lungenschnecken. (Heli-

cinacea et Cyclostomacea). Systematisches Conclwlien-

Cabinet von Martini und Chemnitz (Kiister ed.). Part 19

Installment 128: pis. 31-36, pp. 229-268.

Pfeiffer, L. 1854a. Descriptions of twenty-four new species of

land shells, collected by M. Salle on the island of St.

Domingo, from Mr. Cuming’s collection. Proceedings of

the Zoological Society for 1852 (20): 138-144, pi. 13.

Pfeiffer, L. 1854b. Die gedeckelten Lungenschnecken. (Heli-

cinacea et Cyclostomacea). Systematisches Conchylien-

Cabinet von Martini und Chemnitz (Kiister ed.). Part 19.

Installment 133: pis. 37-42, pp. 269-308.

Pfeiffer, L. 1854c. Die gedeckelten Lungenschnecken. (Helicinaeea

et Cyclostomacea). Systematisches Conehylien-Cabinet von

Martini und Chemnitz (Kiister ed.). Part 19. Installment 136:

pis. 43-48, pp. 309-356.

Pfeiffer, L 1854d. Die gedeckelten Lungenschnecken. (Heli-

cinacea et Cyclostomacea). Systematisches Conehylien-

Cabinet von Martini und Chemnitz (Kiister ed.). Part 19.

Installment 137: pis. 49, 50, pp. 357—400.

Pfeiffer, L. 1858a. Zur Molluskenfauna der Insel Cuba. Mala-

kozoologische Blatter 5: 37-49.

Pfeiffer, L. 1858b. Monographia pneumonopomorum viventium,

Supplementum Primum. T. Fischer, Cassellis. 249 pp.

Pfeiffer, 1,. 1862. Bemerkungen zu den beschribenen Arten.

Malakozoologische Blatter 9: 91-97.

Pfeiffer, L. 1865. Monographia pneumonopomorum viventium,

Supplementum Secundum. T. Fischer, Cassellis. 284 pp.

Pfeiffer, L. 1876. Monographia pneumonopomorum viventium.

accedente fossilium enumeratione. Supplementum ter-

tium, monographiae auriculaceorum. Parte secunda auc-

tum. T. Fischer, Cassellis. x + 479 pp.

Pfenninger, M., M. Hrabakova, D. Steinke and A. Depraz. 2005.

Why do snails have hairs? A Bayesian inference ol character

evolution. BMC Evolutionary Biology 5: 111.

Pilsbry, H.A. 1933. Santo Domingo land mollusks collected by

Samuel C. Pease, 1932, and by A. A. Olsson, 1916. Proceedings

of the Academy of Natural Sciences 85: 121 162, pis. 6-11

Reeve, L.A. 1863. Monograph of the genus Chondropoma.

Conchologica Iconiea 14. Reeve and Co., London, pis.

1-8 + text.

Richardson, C.L , R. Robertson, G.M. Davis and E.E. Spamer.

1991. Catalog of the tvpes of neontologieal Mollusca in the

Academy of Natural Sciences of Philadelphia. Pt. 6. Gas¬

tropoda: Mesogastropoda: Viviparacea, Valvatacea, Littor-

inacea, Rissoacea (Pt. 1: Adeorbidae, Amnicolidae,

Anabathridae, Assimineidae, Barleeidae, Bithyniidae,

Caecidae, Cingulopsidae, Elashisinidae, Falsicingulidae).

Tryonia (23). 243 pp.

Richards, M.C. and W.E. Olds. 1969. A catalogue of moUuscan

type specimens in the Department of Living Invertebrates,

the American Museum of Natural History. Department of

Living Invertebrates, American M useum of Natural His¬

tory, New York. 147 pp.

G.T. Watters and P. Larson, 2017

Page 201

Ronquist, F. and | P. Huelsenbeck. 2003. MRBAYES 3:

Bayesian phylogenetic inference under mixed models.

Bioinformatics 19: 1572-1574.

Shuttleworth, R.J. 1854. Beitrage zur naheren Kenntniss der

Land- und Siisswasser-Mollusken der Insel Portorico.

Mittheilungen der naturforschenden Gesellschaft in Bern

(321-322): 89-103.

Skomrock, N. 2014. The biogeography of the Caribbean land

snail family Annulariidae. Master of Science Thesis, De¬

partment of Evolution, Ecology and Organismal Biology,

Ohio State University.

Sowerby 1 1. G.B. 1843. Thesaurus conchyliorum, or monographs

of genera of shells, 1 . Monograph of the genus Cyclostoma.

Sowerby, London, pp. 89-156.

Stamatakis, A. 2014. RAxML Version 8: A tool for phylogenetic

analysis and post-analysis of large phylogenies. Bio¬

informatics 30: 1312-1313.

Thiele, |. 1929. Handbuch der Systematischen Weichtierkunde.

Gustav Fischer, Jena. pt. 1: pp. 1-376.

Trvon, G.W. 1883. Structural and systematic conchology. Pri¬

vately printed. Philadelphia. 2: 430 pp., pis. 23-91.

van der Sehalie, H. 1948. The land and fresh-water mollusks of

Puerto Rico. Miscellaneous Publications, Museum of Zo¬

olog)', University of Michigan (70): 1-134.

Van Osselaer. C 1999. Counting shell whorls. Remarks. Apex 14:

33-42.

Watters. G.T. 2006. The Caribbean landsnail family Annular¬

iidae. A revision of the higher taxa and catalog of the

species. Backhuys Publ.. Leiden. 584 pp.

Watters, G.T. 2012. Hispaniolan Annulariidae (Gastropoda),

primarily from the Barahona Peninsula: New taxa and

notes. 'Phe Nautilus 126: I 14.

Watters, G.T. 2013. Description of a new species of Licina

(Annulariidae) from Haiti. Visaya 4: 4-9.

Watters, G.T. 2016a. Identification and redescription of the

Haitian Chondropoina semilabre ol Bartsch, 1946, non

Lamarck, 1822 (Gastropoda: Annulariidae). The Nautilus

30: 1-3.

Watters, G.T. 2016b. The Abbott eUa nwreletiana complex in

Hispaniola: distributional notes, a new species, and the

recognition of a new subfamily (Annulariidae: Abbottelli-

nae). Journal ol Conchology 42: 1 1 1-1 17.

Watters, G.T. and G. Duffy. 2010. New species of Annulariidae

(Gastropoda) from the Bahamas and Dominican Republic.

Novapex 11: 1-12.

Watters, G.T. and C. Frank-Fellner. 2017. A new species

of Annulariidae from eastern-most Hispaniola (Gastro¬

poda, Littorinoidea). |ournal ol Conchology 42:

531-534.

Weinland, D.F. 1880. Zur M oil usken fauna von Haiti.

Jahrbiicher der Deutschen Malakozoologischen Gesell¬

schaft nebst Nachrichtsblatt 7: 338-378. pi. 12

Wenz, W. 1939. Gastropoda. Handbuch der Palaozoologie. 6(3).

AlWmeiner Teil und Prosobranchia. Gebriider Born-

O

traeger, Berlin. Pp. 484-720.

Wyman, R.H. 1874. West Indies - West coast ol Hayti -

Cayemites Bay. U.S. Hydrographic Office, Hydrographic

Notice (41): 1-2.

Designation of a lectotype for Parvaspina

collina (Conrad, 1836) (Bivalvia:

Unionidae)

Perkins et al. (2017) erected a new genus, Parvaspina ,

for two species with spines formerly placed in Elliptic

and Pleurohema . Based on genetic information. Elliptic

steinstansana (Johnson and Clarke, 1983) and Pleurohema

collina (Conrad, 1836) were transferred to Parvaspina.

Unio collinns Conrad, 1837[sic] was designated the type

species of the new genus (Perkins et ah, 2017). They noted

the lectotype was ANSP 41007. They did not cite the

revisionary work of Johnson (1970) nor mention the

lectotype designation as presented bv Johnson and Baker

(1973:151) or Graf and Cummings (2017).

In his Monographs', Conrad (1835-1840), figured what

appears to be three specimens of Unio collinns before the

tex't description was published (Conrad, 1836f (No. 7)): pi.

36, fig. 2). This plate is dated 1836, so the taxon takes this

i.T/nio occidental is, C. Z. ZXcollinus, Co v .

Figure 1. Original illustrations of Unio collinns in Conrad (1836: pi. 36 fig. 2).

A.E. Bogan, 2017

Page 203

date of publication. This is an indication and is the first use

of the name Unio collinus . Conrad’s pi. 36, fig. 2 (Figure 1 )

is interpreted as representing all three of his original

syntype specimens: the two figures in the top row are one

specimen; the dorsal view in the middle row shows an

eroded umbo area without any evidence of spines is a

second specimen and the bottom figure of the outside of

the right valve is the third specimen.

Lectotype Designation: Conrad (1837a (No. 8 ) : 65 ) in

the Monography described Unio collinus and mentioned

he had three specimens and referred to pi. 36, fig. 2, citing

the catalog number “Cab. ANSP No. 20408". Sub¬

sequently, he remarked “since the publication of this

species in a former number of this work, 1 have received

a few specimens from the same locality with the first

described, which present the very remarkable character of

spines” (Conrad, 1840a (No. 12): 1 09 — 1 10, pi. 60, fig. 3).

No catalog number was listed for these new specimens.

One of these new specimens was figured in pi. 60, fig. 3

(Conrad 1840a) (Figure 5). The problem is that the

specimens later received and figured are topotypes

identified by the author of the species, not part of the

original type series (See Articles 74.1; 74.2). Johnson

(1970: 300) noted the type lot, ANSP 20408, was lost and

observed “this subsequently figured metatype, here selected,

lectotype ANSP 41007.” The IUCN Code does not recognize

die term metatype. This topotypic lot was not part of the

original type series and as such cannot be used as a source for

designating a lectotype. Therefore, Johnson’s (1970) lecto¬

type designation is invalid. The original figure of Conrad

(1836f: pi. 36, fig. 2) can be used to designated the lectotype

even if the specimens are lost (see Code Articles 74.1. 74.2,

74.4). Fig. 2 of pi. 36, assigned museum number ANSP

20408, represents the valid syntype series, even if the

specimens are lost The top two valves in Conrad’s fig. 2

(see Figure 1) are here designated the lectotype for

Unio collinus Conrad, 1836 (see Code Article 74.4).

The catalog number ANSP 20408 was listed for the

original type series by Conrad (1837a:65). This number

is not a Malacology Department number. The ANSP

Malacology Department did not begin assigning catalog

numbers until 1889. The invalid lectotype ANSP 41007 is

in the collection and identified as Unio collinus. Rosenberg

remarked “I suspect that there was an early ledger where

catalogue numbers were assigned to items across the

Academy, since in the 1830’s we didn’t have formal de¬

partments. If there was such a ledger, however, it no

longer exists ’ (Gary Rosenberg, Pers. Comm.).

Parvaspina collina. ANSP 41007, was designated as a

lectotype (Figures 2-4), but does not match any of the

original figures by Conrad (Figure 1), more closely re¬

sembling the subsequent figure by Conrad (see Figures 4

and 5). The ANSP 41007 specimen hits Rockbridge County

written in pencil on the inside of the left valve, and has some

dried tissue on the inside of the shell (Figure 3). but lacks

a catalog number written in the shell. Rockbridge Count}' is

Figures 2-5. Parvaspina collina (Conrad) ANSP 41007 invalid lectotype. 2. External surface of right valve. 3. Internal surface of left

valve. 4. Dorsal view of paired valves. 5. Original illustration of Unio collinus in Conrad (1840: pi. 56, fig. 3).

Page 204

THE NAUTILUS, Vol. 131, No. 3

listed in the type locality. It has been suggested that the

specimen catalogued today as ANSP 41007 was part of

tin1 original type series. However, comparison of the

valves of the ANSP 41007 specimen (Figures 2-4) with

the illustration of Conrad's type figures (Figure 1 ), makes

it clear that the catalogued specimen was not one

originally illustrated in Conrad’s pi. 36 fig. 2 (Figure 1). It

does resemble Conrad's later pi. 56, fig. 3 (Figure 5).

This being the case, then ANSP 41007 is a topotype and

not part of the original type series.

ACKOWLEDGM FNTS

Ms. }amie M. Smith and Mrs. Cindy M. Bogan both

reviewed an earlier draft of this note. Ms. Ellen Wildner

is thanked for her help with the ANSP Collection and

Ms. Krasimira Seizova took the pictures of the invalid

type specimen, both from the Malacology Department,

Academy of Natural Sciences of Drexel University,

Philadelphia. Dr. Gary Rosenberg, Malacology Depart¬

ment, Academy of Natural Sciences of Drexel Univer¬

sity, Philadelphia, was a reviewer and provided valuable

direction and suggestions to improve and clarify this

manuscript.

LITERATURE CITED

Conrad, T.A. 1835-1840. Monography of the family Unionidae,

or naiades of Lamarck, (fresh water bivalve shells) of North

America, illustrated by figures drawn on stone from nature.

J. Dobson, Philadelphia. Part 1(1835)4-12 [pages 13-16

not published], pis. 1-5; Part 2(1836a)4 7-24, pis. 6-10:

Part 3(1836b):25-32, pis. 1 1-15; Part 4( 1836c): 33-40, pis.

16-20; Part 5( 1 836d):4 1 — 48, pis. 21-25; Part 6(1836e);

49-56, pis. 26-30; Part 7(1836f):57-64, pis. 32-3.6; Part 8

(1837a);65-72, pis. 36-40; Part 9(1837b);73-80, pis. 41-45;

Part 10(1838a):81-94, pis. 46-51; Part 1 1 ( 1838b):95- 1 02,

pis. 52-57; Part 12( 1840a): 103-1 10, pis. 58-60; Part 13

[1840b, Part 13 not dated]:! 1 1-1 18, pis. 61-65.

Graf, D. and k Cummings. Mussel-project webpage: http://

musscl-in-oject.iiwsp.edu/ [accessed 18 April 2017],

International Commission on Zoological Nomenclature. 2017

International code of zoological nomenclature. Fourth

edition, http://wunc.iczn.org/iczn/injdex.jsp [accessed 25

April 2017],

Johnson, R.l. 1970. The systematics and zoogeography of the

Unionidae (Mollusca: Bivalvia) of the southern Atlantic

Slope Region. Bulletin ol the Museum of Comparative

Zoology 140; 263-450.

Johnson, R.l and H.B. Baker. 1973. The types of Unionacea

(Mollusca: Bivalvia) in the Academy of Natural Sciences

of Philadelphia. Proceedings of the Academy of

Natural Sciences of Philadelphia 125(9); 145-186, pis.

1-10.

Johnson, R.l and A. II. Clarke. 1983. A new spiny mussel,

Elliptio (Canthyria) steinstcmsana (Bivalvia; Unionidae),

from the Tar River, North Carolina. Occasional Papers on

Mollusks, Harvard University, Museum of Comparative

Zoology 4(61): 289-298.

Perkins, M.A., N.A. Johnson, and M.M. Gangloff. 2017. Mo¬

lecular systematics of the critically-endangered North

American spinvmussels (Unionidae: Elliptio and Pleuro-

hema) and description of Parvnspina gen. nov. Conserva¬

tion Genetics DOI 10.1007/1 1-592-0 17-924-z.

Arthur E. Bogan

North Carolina Museum of Natural Sciences

1 1 West Jones Street

Raleigh, NC 27601 USA

[email protected]

O O

m

PDo.

a United Malacologists

January 27, 2018

FLORIDA UNITED MALACOLOGISTS 2018

The ninth meeting of Florida United Malacologists (FUM 2018) wall take place on Saturday, January 27, 2018, at the

Bailey-Matthews National Shell Museum on Sanibel Island, Florida. The one-day gathering brings together researchers,

citizen scientists, and students interested in a broad swath of mollusk-related topics. FUM follows the pattern of similar

informal gatherings such as BAM (Bay Area Malacologists), SCUM (Southern California United Malacologists), MAM

(Mid-Atlantic Malacologists), and OVUM (Ohio Valley United Malacologists). The event circulates among different

Florida organizations, hut usually takes place at the Shell Museum every other year. Presentations are limited to

15 minutes plus 5 minutes for questions. Presenters are required to submit a brief abstract limited to 1 50 words or less. The

gathering will be free to presenters and pre-registered participants. Box lunches and dinner at a local restaurant (to he

arranged) will be available to participants and presenters. The deadline for abstract submission is December 31. 2017.

For registration and further information, visit http://shellmuseum.org/about/news/florida-united-malacologists-2018, or

email [email protected].

Sponsored in part by the State of

Florida, Department of State,

Division of Cultural Affairs and the

Florida Council on Arts and Culture

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