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CONTENTS

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G. Thomas Watters

Megan L. Smith

David J. Sneddon

Anton E. Oleinik

Alexander B. Modys

Angelina M. Tetu

Shugian Zhang

Suping Zhang

Kazutaka Amano

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Unusual shaping: The defecation behavior in Cochlorina aurisleporis

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THE NAUTILUS 134(1):1-34, 2020

Page |

The subfamily Abbottellinae (Gastropoda: Annulariidae): origins,

associations, and a review of the Hispaniolan taxa

G. Thomas Watters

Megan L. Smith

David J. Sneddon

Museum of Biological Diversity

Department of Evolution, Ecology and Organismal Biology

Ohio State University

Columbus, OH 43212 USA

[email protected]

ABSTRACT

The subfamily Abbottellinae of the Annulariidae has been

recognized as a speciose group in northern Hispaniola. This

study, using phylogenetic (COI, 12S, H3), radular, and con-

chological characteristics, compared this group to other non-

Hispaniolan annulariids to determine their wider associations

and potential origin. Results indicate that the subfamily is related

to the eastern Cuban Annularisca sensu lato and that genus

should be moved to the Abbottellinae. The group dates from the

time when eastern Cuba and northern Hispaniola were in close

contact prior to tectonic movement. This is the first such zoo-

geographic connection reported for the family. The subfamily is

not related to any Jamaican taxa tested. The Abbottellinae also

do not occur on the southern Tiburon/Barahona peninsulas of

Hispaniola, which has a different geologic history from the north.

The eastern Cuban and Bahamian genus Opisthosiphon sensu

lato was also shown to be a member of the subfamily. The

nominal genus Abbottella is further divided into new genera

based on these results. The new genera Abbottipoma, Are-

nabbottella, Microabbottella, and Preclaripoma are described.

The new species Rolleia simonaikeni is described, and Petasi-

poma is synonomized with Rolleia.

Additional Keywords: Zoogeography, phylogenetics, Cuba,

Hispaniola, new taxa, terrestrial gastropods, land snails

INTRODUCTION

The land snail family Annulariidae is an exclusively

Caribbean/Central American group. For its limited geo-

graphic distribution it is extremely speciose with >700

recognized taxa (Watters, 2006). The major centers of di-

versity are Cuba, Hispaniola, and Jamaica. The snails are

strict calciphiles and many species are highly endemic to a

particular mountain range, mogote, or outcrop. Of these, the

genus Abbottella was phylogenetically identified as a discrete

In absentia

clade apart from other annulariids by Skomrock (2014) and

Watters and Larson (2017). Watters (2016b) formally raised

the clade to subfamily level (Abbottellinae) based on ad-

ditional radular and shell morphology characteristics.

However, the questions of the origin of the subfamily and its

relationships to other annulariids remained unanswered.

In all of the various schemes for higher annulariid

systematics (Watters, 2006) subfamilies are distributed

across the geographic range of the family — only the

Abbottellinae, as previously understood, were endemic to

a single portion of Hispaniola. Yet it may have been

derived from elsewhere. In the Paleogene, proto-Central

America was connected to Cuba and non-Tiburon His-

paniola, an association that may have lasted until the

Eocene (Chakrabarty, 2006) or Miocene (Pindell, 1994)

and provided ample time for faunal exchange. Since this

time, these islands drifted east to their present positions,

separating 20 to 25 million years ago (Pindell, 1994) and

taking their snail faunas with them to further evolve in

isolation. Buskirk (1985), reviewing the distributions of

numerous groups of terrestrial animals (including snails)

and pollen, also suggested a radiation from North and

proto-Central America to the Greater Antilles by the

Eocene. This would suggest a possible origin or con-

nection for the abbottellines in Cuba or Jamaica.

This study examines the Abbottellinae based on phylo-

genetic, radular, and shell morphological consider-

ations. In particular, membership to the subfamily is

sought outside of Hispaniola in an attempt to deter-

mine: 1) if the group is indeed endemic to Hispaniola;

2) where might the origin of the group reside; and 3) what

is its relation to the other subfamilies. Potential candidates

are drawn from Cuba, Jamaica, and elsewhere based on

similarities in shell morphology and radulae. Of particular

interest are the similar-appearing Annularia of Jamaica and

Annularisca of Cuba. In addition, the relationships between

the recognized abbottelline genera are also examined.

Abbreviations used in the text are: ANSP, Academy of

Natural Sciences, Philadelphia, Pennsylvania, USA; BMSM,

Page 2

Bailey-Matthews National Shell Museum, Sanibel, Flor-

ida, USA; GTW, collection of the author, Columbus,

Ohio, USA; MNHN, Muséum national d’Histoire

naturelle, Paris, France; NHMUK, Natural History

Museum, London, UK; OSUM, Ohio State University

Museum of Biological Diversity, Columbus, Ohio,

USA; UF, Florida Museum of Natural History,

Gainesville, Florida, USA; USNM, National Museum

of Natural History, Washington, DC, USA; ZMB,

Zoologisches Museum Berlin, Federal Republic of

Germany. Numbers in ( ) after catalog numbers refer

to the number of specimens.

MATERIALS AND METHODS

To evaluate the phylogenetic placement of the Abbottel-

linae and to understand relationships among genera within

Abbottellinae, we collected molecular sequence data

from 19 individuals from Abbottellinae and 28 individ-

uals from Annulariidae and Pomatiidae (Appendix1). We

also downloaded data from 53 individuals from GenBank

(NCBI nucleotide database) (Appendix1). As outgroups,

we used Pomatias elegans (Miiller, 1774) and Tudorella

ferruginea (Lamarck, 1822) from Pomatiidae, and, as a

more distant outgroup, Littorina littorea (Linnaeus,

1758) from Littorinidae. Samples collected for this study

were stored in ethanol after collection, and DNA was

extracted from foot tissues of preserved specimens using

Qiagen DNeasy Blood and Tissue Kits (Qiagen Hiden,

Germany) following the manufacturers standard proto-

col. A region of the mitochondrial gene 12S was se-

quenced using primers developed by Watters and Larson

(2017). A region of cytochrome oxidase 1 (COI) was

sequenced using universal invertebrate primers

LCOI490 and HCO2198 (Folmer et al. 1994). The

nuclear gene histone 3 (H3) was amplified using primers

H3aF and H3aR (Colgan et al., 1998). PCR products

were cleaned using ExoSAP-IT (Thremofisher Scien-

tific) and sent to the Ohio State University Biomedical

Research Tower for sequencing. Chromatograms were

assembled in Geneious v6.1.7 (Kearse et al., 2012) and

edited by eye when necessary. Sequences were aligned

using the MUSCLE (Edgar, 2004) algorithm imple-

mented in Geneious (Kearse et al., 2012) using default

parameters.

The best model of nucleotide substitution and par-

titioning scheme was determined using PartitionFinder

v2.1.1 (Lanfear et al., 2016). We used a greedy search

scheme, considered all models of nucleotide substitu-

tion, and performed model selection using corrected

AIC. A maximum-likelihood tree was inferred for COI

using RAxML v8.2.10 (Stamatakis, 2014) as imple-

mented on the Cipres Science Gateway (Miller et al.,

2015). We used the partitioning scheme determined by

PartitionFinder and the GTR model with Gamma rate

variation. We used 100 bootstrap replicates to assess

confidence in the inferred relationships. To infer a

posterior distribution of gene trees, we used Bayesian

THE NAUTILUS, Vol. 134, No. 1

Inference in MrBayes v3.2.2 (Ronquist et al, 2012). We

used the partitioning scheme and models determined by

PartitionFinder. Two independent analyses were run for

5,000,000 generations over four chains, sampling every

1,000 generations. We discarded 25% of runs as burn-in, and

we assessed convergence using Tracer v1.6.0 (Rambaut and

Drummond, 2007).

To infer a more robust hypothesis of relationships

between groups, we used a concatenated dataset in-

cluding the mitochondrial COI and 12S genes and the

nuclear gene H3. While this dataset included fewer

taxa than the COI dataset, it included a larger data

matrix. Given that many taxa within Abbottellinae and

those grouping with Abbottellinae in the COI gene

tree could be included in this dataset and that rela-

tionships between these groups were not well resolved

using COI (see Results), we analyzed this dataset in an

attempt to further resolve relationships between these

groups. While we acknowledge that a coalescent-based

analysis would have been ideal (Edwards et al., 2016),

given the limited sampling (2 independent loci, not

sampled in all specimens), only a concatenation ap-

proach was possible for this study. All samples for

which at least two of the three genes were sequenced

were retained for this analysis. As above, we used

PartitionFinder to select the best model and parti-

tioning scheme, RAxML to infer a maximum likelihood

tree, and MrBayes to infer the posterior distribution of

trees under Bayesian inference.

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 measured from the tip of the

protoconch (or teleoconch of decollate specimens) to

the opposite anterior-most extension 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 1 D method of Van Osselaer (1999).

Detailed redescriptions, accounts, and translated

original descriptions of species previously covered can

be found in Watters (2013, 2016b) and are not du-

plicated here. Distribution maps are Google’ ™ Earth

NOAA, US Navy, NGA, GEBCO.

RESULTS

Our final COI alignment included 82 individuals and

582 base pairs. The concatenated alignment of COI,

12S, and H3 included 50 individuals and 1,351 base

pairs. For COI, the best scheme determined using AIC

in PartitionFinder included three partitions (first,

second, and third codon positions). The best models of

nucleotide substitution were TVM+G, TRN+I1+G,

and GTR+I for the first, second, and third codon

positions, respectively. The best scheme identified by

G.T. Watters et al., 2020

PartitionFinder for the concatenated dataset included

seven partitions: One for each codon position of COI

and H3 (for a total of six) and a single partitioning

scheme for 12S. The best models inferred by Parti-

tionFinder were TRN, JC+I, K81UF+G, GTR+G,

HKY+G, TRN+G, and GTR+I for the first, second,

0.39/59

1194

6.96/61

1798

0.99/96 a .

0. 9 Gems Wt

(84 ploBostyids retro

1/100

USDA _Iber994_01 Tudorelia ferruginea

$436d Xenopomea spinosossimum

USDA_O7FRA03_01 Pomatias elegans

Page 3

and third codon positions of H3, 12S, and the first,

second, and third codon positions of COI, respectively.

Abbottellinae was recovered as a monophyletic group

(BS = 100, pp = 1.0) under both maximum likelihood and

Bayesian inference of the COI gene tree (Figure 1), but

there was not strong support for the relationship of this

7064d Diplopoma crenuiatum (Antigua)

-_ KX496717 Chondropoma pupitorme (Anguilla)

Figure 1. Maximum Likelihood tree based on COI. The numbers above the branches are posterior probability/ bootstrap support

values. Support values are only shown for branches with either > 70 bootstrap support or >0.90 posterior probabilities. Taxa from the

DR are highlighted in blue, taxa from Cuba are highlighted in yellow, taxa from the Bahamas are highlighted in green, taxa from Florida

are highlighted in orange, and taxa from Jamaica are highlighted in purple. The Abbottellinae are labelled.

Page 4

group to other sampled individuals. Within Abbot-

tellinae, there were three moderately-to-well sup-

ported groups. One included Abbottella moreletiana,

Abbottella crataegus, and Abbottella domingoensis

(BS= 100, pp = 1.0). A moderately supported group

(BS=73, pp = 0.91) included Abbottipoma abbotti,

Abbottipoma crossei, and Lagopoma lagopoma. The

third group included Arenabbottella sosuaensis, Are-

nabbottella rosaliae, Arenabbottella mellosa, and

Leiabbotella galaxius (BS = 67, pp = 0.92). The

placements of Meganiphe rhecta, Rolleia oberi, and

Preclaripoma thompsoni with respect to other

Abbottellinae were not well resolved in either analysis.

The Bayesian analysis supported a group including the

Hispaniolan abbottellines, Annularisca, Opisthosiphon,

and Parachondria dentatus (BS = 59, pp = 0.99).

The maximum likelihood and Bayesian analyses of

the concatenated dataset (Figure 2) also recovered the

Abbottellinae as a monophyletic group (BS = 100,

pp = 1.0). The same groups within Abbottellinae

supported by the COI dataset were recovered with the

concatenated dataset, and Meganiphe rhecta was re-

covered as sister to the group containing Leiabbottella

galaxius, Arenabbottella rosaliae, and Arenabbottella

sosuaensis (BS = 94, pp = 1.0). There was support

(BS=75, pp = 0.98) for a clade including the

Abbottellinae, Annularisca, Opisthosiphon, and Par-

achondria dentatus, but, as with the COI dataset, we

could not resolve the relationships between these

6.92/80

1/466

1/100

THE NAUTILUS, Vol. 134, No. 1

groups. There was also moderate support (BS = 62,

pp = 0.96) for the other Parachondria as sister to this

clade.

DISCUSSION

MOLECULAR PHYLOGENETICS

The results of this study indicate that the Abbottellinae

represents a distinct group well-supported by phylo-

genetic, radular, and conchological analyses. Though

results were originally limited to the Abbottella-like

taxa of Hispaniola, evidence presented here indicates

this subfamily also includes the eastern Cuban

Annularisca and Opisthosiphon. No abbottelline was

found to cluster with any Tiburon/Barahona penin-

sular (e.g., Chondropomella, Chondropomium, Cly-

donopoma, Superbipoma) or Jamaican taxa.

Phylogenetic analyses supported the monophyly of

Abbottellinae, based on the species included in this

study. Future work should evaluate additional Abbot-

tellinae to further test monophyly of the group. How-

ever, given that this is the first attempt at a molecular

phylogeny in this group and samples for DNA analyses

are limited for some species, these results are promising.

Results also supported the monophyly of the genera

Abbottella and Arenabbottella. The results presented

here do not support the monophyly of Abbottipoma with

respect to Lagopoma, and future work should include

1/700

USDA_O7FRAO3_01 Pomatias elegans

USDA_iberf9A_01 Tudorelia ferruginea

Figure 2. Maximum Likelihood tree based on the concatenated dataset. The numbers above the branches are posterior probability/

bootstrap support values. Taxa from the DR are highlighted in blue, taxa from Cuba are highlighted in yellow, taxa from the Bahamas are

highlighted in green, taxa from Florida are highlighted in orange, and taxa from Jamaica are highlighted in purple. The Abbottellinae are

labelled.

G.T. Watters et al., 2020

additional representatives from Lagopoma to evaluate

the monophyly of this group.

The Jamaican genera Annularia Schumacher, 1817,

Colobostylus Crosse and Fischer, 1888, and Adamsiella

Pfeiffer, 1851, were not found to be related to the

Abbottellinae in this study. Although Annularia taxa have

a similar shell form to abbottellines, the similarity is ap-

parently due to convergence.

Cuban Annularisca, Bajaman and Cuban Opisthosi-

phon, and Parachondria dominguensis from Key West

grouped with Abbottellinae in this study. However, we

were unable to resolve the relationships between these

groups and Abbottellinae, either with the COI dataset,

or with the concatenated dataset with more infor-

mation. Both trees include a polytomy with Annu-

larisca, Opisthosiphon, and Parachondria dentatus.

Future work with additional loci may be necessary to

resolve these relationships.

SYSTEMATICS

Torre and Bartsch (1941) finely divided a large group of

similar Cuban species into several subgenera, all of

which they placed under Annularia Schumacher, 1817.

Annularia is now considered a Jamaican endemic ge-

nus. Their subgeneric taxa were differentiated by minor

differences in the strength of the sculpture and form of

the lip. These taxa included: Annularella, Annularisca,

Annularosa, Chondropomatus, all Henderson and

Bartsch, 1920, and Annularex and Lugarenia, both

Torre and Bartsch, 1941. Watters (2006) consolidated

all of these under the genus Annularisca, although

Lugarenia is probably not related. They are all endemic

to eastern Cuba. In addition to being phylogenetically

sister to the Hispaniolan abbottellines, they share the

same “Mitten” type radula (see below), similar oper-

cular structure, and shell sculpture. Annularisca shell

forms are very similar to Abbottella but consistently

differ from those of other annulariids: low turbinoid,

non-decollate, openly umbilicate shells. Cuban Annu-

larisca species appear to be larger versions of Hispa-

niolan abbottellines (Figures 3-7). With the exception

of Lugarenia, all of these Cuban taxa are here moved to

the Abbottellinae.

Opisthosiphon was found to be part of the Abbot-

tellinae clade as well. This genus (and its various sub-

genera) are widespread in eastern Cuba and _ the

Bahamas, with numerous nominal species. Watters

(2006) reduced the ten subgenera of previous workers to

three. No species are known from Hispaniola, Puerto

Rico, or Jamaica. It is probable that the genus originated

in Cuba and has dispersed throughout the Bahamas by

rafting. Opisthosiphon have “breathing” devices—

structures that allow the animal to communicate with the

outside when the operculum is withdrawn. This device

varies from a simple pore to (more often) a short re-

flected siphon or snorkel. However, Opisthosiphon

shares the “Mitten” type radula with abbottellines. It

differs from the rest of the abbottellines in the following

Page 5

ways: the rhytidopomine operculum (pin-wheel exten-

sions across the operculum rather than the single spiral

lamellum of other abbottellines); the shells are decol-

lated in Opisthosiphon but entire in the remainder of

other abbottellines; the shell shape of Opisthosiphon is

pupoid in overall shape rather than depressed turbinoid.

Given the diversity of Opisthosiphon species in neigh-

boring eastern Cuba, it is peculiar that none exist in

Hispaniola. It is possible they did at one time but have

been extirpated.

Opisthosiphon has been included in the Rhytidopo-

minae, a heterogeneous group (as envisioned by Hen-

derson and Bartsch, 1920) of Cuban taxa that Watters

(2006) placed under the Chondropomatinae — a scheme

that can no longer be maintained based on these results.

The type genus is Rhytidopoma Sykes, 1901, based on

the Cuban Cyclostoma rugulosum Pfeiffer, 1839, which

resembles Opisthosiphon and has a breathing device as

well. No Rhytidopoma taxa were available for this study.

This presents a potential problem in the taxonomy of the

Abbottellinae. If Rhytidopoma is part of the Abbottella

clade, the nomen Rhytidopominae Henderson and

Bartsch, 1920, would have precedence over Abbottel-

linae Watters, 2016. Future studies involving additional

Cuban taxa may well show that not only Abbottella and

Opisthosiphon, but Rhytidopoma and similar groups

such as Ramsdenia Preston, 1913, and Cubadamsiella

Torre and Bartsch, 1941, are related in one clade under

Rhytidopominae that spans northern Hispaniola, eastern

Cuba, and the Bahamas. Representative members of

Opisthosiphon are shown in figures 8-12.

The Floridian Parachondria dentatum (Say, 1825)

grouped here with Opisthosiphon although it lacks a

breathing device. As noted by Watters (2016a), “Para-

chondria” is a wide-ranging catch-all group that un-

doubtedly contains numerous unrelated taxa in unnamed

genera; the type of the genus is Jamaican. Parachondria

dentatum is very similar to Opisthosiphon and is here

considered a relative of that group lacking a siphon; it

may require a new generic name.

ZOOGEOGRAPHY

The Hispaniolan Abbottellinae do not occur in the

Tiburon/Barahona peninsulas, which have a different

geological history and a unique annulariid fauna apart

from the remainder of northern Hispaniola, here

designated Hispaniola(N). Occupying two different

tectonic plates, the Tiburon Peninsula and Hispa-

niola(N) have been colliding since perhaps the Oli-

gocene (van Benthem et al., 2013) but have exchanged

very few annulariid genera. Based on the results

presented here, the abbottellines are more closely

related to taxa from eastern Cuba than they are with

those from the Tiburon Peninsula, with which they

share the island. This is the first such zoogeographic

connection reported for the family.

Central to the understanding of this zoogeographic

puzzle is the geologic history of the Caribbean. Eastern

Page 6 THE NAUTILUS, Vol. 134, No. 1

20 x 24

Figures 3-21. Annularisca, Opisthosiphon, radular types, distribution maps of genera. 3-7. Representative Cuban Annularisca. 3.

Annularisca mayariensis (Torre and Bartsch, 1941). GTW 14027a, Cantera de Mella, 11.7 mm diameter. 4. Annularisca auricoma putre

(Pfeiffer, 1863). GTW 7175a, Tortuguilla, 11.8 mm diameter. 5. Annularisca heynemanni (Pfeiffer, 1864). GTW 14026a, Maisi,

14.2 mm diameter. 6. Annularisca victoris (Torre and Bartsch, 1941). GTW 7163d, Maisi, 12.1 mm diameter. 7. Annularisca mackinlayi

(Pfeiffer, 1859). GTW 14458a, Yateras, 11.7 mm diameter. 8-12. Representative Opisthosiphon. 8. Opisthosiphon moreletianum (Petit,

1850). GTW 7141b, Isla de Juventud, Cuba, 14.8 mm length. 9. Opisthosiphon berryi Clapp, 1919. GTW 7076a, Paso Gunaja, Cuba,

8.4mm length. 10. Opisthosiphon paredonense Torre and Henderson, 1921. GTW 8050a, Paso de Los Paredones, Cuba, 10.3 mm length.

11. Opisthosiphon pupoides velazquexi Torre and Bartsch, 1941. GTW 7140c, Isla de Juventud, Cuba, 13.8 mm length. 12. Opisthosiphon

cucullatum Torre and Bartsch, 1941. GTW 16588a, Calvario, Cuba, 16.4 mm length. 13-17. Radular types. R — rachidian; L— lateral; IM —

inner marginal; OM — outer marginal. Bar = 100w. 13. Glove — Halotudora sumichrasti (Crosse and Fischer, 1874). Mexico. 14. Hook —

Chondropomium marmoreum (Watters and Duffy, 2010). Dominican Republic. 15. Mitten — Arenabbottella rosaliae (Pfeiffer, 1858).

Dominican Republic. 16. Mitten — Annularisca roemeri (Pfeiffer, 1864). Cuba. 17. Mitten — Opisthosiphon bahamense (Pfeiffer, 1865).

Bahamas. 18-21. Distribution maps of genera. 18. white — Abbottipoma; blue — Abbottella; red — Meganiphe. 19. white — Arenabbottella.

20. white — Rolleia; red — Microabbottella. 21. white — Leiabbottella; red — Lagopoma; blue — Preclaripoma.

G.T. Watters et al., 2020

Cuba and Hispaniola(N) were connected beginning with

the Oligocene (Rosen, 1976; Judd, 2001; Ottenwalder,

2001), with Puerto Rico connected with eastern His-

paniola(N) from the middle Eocene to lower Miocene

(Ottenwalder, 2001). When these were contiguous

during the middle Cenozoic it formed an emergent

pathway for dispersal from eastern Cuba (then a separate

island from the remainder of Cuba) to the Cordillera

Septentrional in northern Hispaniola and eastward to

Puerto Rico. As a result, most of the terrestrial snail

groups in Puerto Rico (van der Schalie, 1948) also occur

in eastern Cuba. Most Hispaniolan abbottellines are

endemic to the northern Cordillera Septentrional and its

extension into the Samana Peninsula. Similarly, the

Cuban Annularisca inhabits the Sierra Maestra, the

eastern mountain range, which is tectonically related to

the Cordillera Septentrional and was an avenue for

dispersal to Hispaniola(N)—but not to the Tiburon/

Barahona peninsulas.

Cuba has been a “hot-spot” for annulariids with much

of the diversity of this family apparently originating on

that island. Containing nearly 300 of the approximately

700 recognized species, the Cuban taxa are some of the

most diverse and probably encompass multiple unique

lineages. The central location of the island, its complex

geologic history of being at least three distinct islands,

and evidence of being at least partially emergent since

the middle Eocene (Graham, 2003) gives support to this

island as a major center for annulariid origination and

dispersal.

In Hispaniola the northern Cordillera Septentrional is

separated from the Cordillera Central to the south by

the Cibao Basin (Oligocene—Pliocene) (Edgar, 1991). The

Cordillera Central is largely metamorphic and supports

very few of these calciphilic snails. South of the Cordillera

Central is the Sierra de Neiba, which although composed

of limestone has no Abbottellinae. Clearly the granitic

Cordillera Central acts as a barrier to dispersal from the

northern Cordillera Septentrional to the southern Sierra

de Neiba, and the Sierra de Neiba is the gateway to the

Tiburon/Barahona peninsulas.

Some abbottelline groups are endemic to the Samana

Peninsula of eastern Hispaniola(N), which is of late-

Miocene or early-Pliocene origin (inundated until

Miocene (Joyce, 1991)). This modern day peninsula was

an island even in historic times (Ross, 1921). Gabb (1873:

161) noted “the Gran Estero which separates Samana

from the main-land was a century ago a navigable

channel, but is now entirely closed. It is said to have been

closed by drift and mud from the [Rio] Yuna.” Cochran

(1941: 2) commented: “Almost within the memory of the

oldest inhabitant, Samanad Peninsula was an island sep-

arated from the main body of Hispaniola by a rapidly

filling strait, in which pirate vessels were said to have lain

in wait for richly laden merchantmen sailing for Europe.”

It is likely that some of the Abbottellinae evolved on

that island rather than being invaders from the mainland

and have never left (e.g., Abbottipoma, Lagopoma,

Microabbottella).

Page 7

SYSTEMATICS

Annulariidae Henderson and Bartsch, 1920

Type Genus: Annularia Schumacher, 1817.

Description: Shells to 42 mm in length, planispiral to

high-spired, often with reflected lip at maturity. Spire

typically decollate. Many taxa wholly or partially solute.

Shell sculpture usually present, rarely produced into short

spines or serrations. Spiral sculpture usually as cords or

threads, axial sculpture as lamellae. In some Cuban and

Bahamian taxa breathing devices are developed, varying

from simple punctures to elaborate internal or external

siphons. Operculum present, varying from paucispiral to

multispiral. Operculum often with calcareous deposit that

may take the form of erect spiral lamellae, overlapping

plates, or pin-wheel-like extensions. Very few species with

obvious periostracum consisting of bristle-like projections.

Animals with bifid snout and foot longitudinally bisected

into lateral lobes. Locomotion ditaxic between lobes of foot.

Snout produced into two short accessory tentacles that

continuously sample substrate. Sexes separate.

Taenioglossate radula with rachidian, single pair of

laterals, and two pairs of marginals. Rachidian and lateral

teeth usually unicuspid, very rarely multicuspid. Inner

marginal multicuspid. Outer marginal pectinate.

Remarks: The taenioglossate radula of annulariids is very

conservative and only the inner marginal is of sufficient

variability to be systematically informative. This tooth is

categorized here as either a Glove, a Mitten, or a Hook

based on its overall shape. The Glove type has a broad

inner marginal tooth divided into blunt, shallowly

indented cusps of varying strength and number

(Figure 13). The Hook radula has a blunt, triangular tooth

with weak cusps along the distal margin and a pronounced

basal hook on the proximal side (Figure 14). The Mitten

type has an elongated triangular tooth characterized by a

single, small basal tooth on the distal side, rarely ac-

companied by weak serrations along the remainder of the

distal margin (Figure 15). The Glove type is by far the

most common radular type seen in >100 annulariid

species examined (Watters, unpubl.). In contrast the

Mitten type is the only radula found in the Abbot-

tellinae (as here defined), including Annularisca

(Figure 16) and Opisthosiphon (Figure 17). The Hook

type is unique to a few genera (e.g., Clydonopoma)

isolated on the Tiburon Peninsula of Hispaniola.

These snails spend an inordinate amount of time

making more snails. Pairs are often found in a mating

position even during dormant periods, with the male

attached to the female on her right side. There appears to

be a distinct sexual color dimorphism in some species

(e.g., Rolleia oberi).

Many annulariids are highly endemic (Watters, 2017).

The entire distribution of some species may consist of

only a few kilometers (or less) of suitable habitat.

Goodfriend and Mitterer (1988, 1993) found that

Page 8

terrestrial snails in Jamaica have had relatively long

spans of endemism, remaining in a single locality for

thousands of years. They suggested that many ter-

restrial snails, at least in Jamaica, have persisted in

place from the Late Pleistocene to the present

(Goodfriend and Mitterer, 1988, 1993). Similarly,

many of the main lineages within Annulariidae ap-

parently have persisted for long periods of time and are

the result of the separation and emergence of the

major landmasses followed by the fragmentation of

populations into discrete, isolated species.

Abbottellinae Watters, 2016

Description: Shells to 15 mm in diameter, pupoid to

turbinoid to planispiral, with reflected lip at maturity.

Spire decollate only in Opisthosiphon and relatives. Some

taxa partially or completely solute. Shell sculpture usually

present, varying from smooth to pustulose to short spines

or serrations. Spiral sculpture, when present, of cords or

threads, axial sculpture, when present, as lamellae.

Operculum present, as multispiral, single, erect, calcar-

eous plate or with pin-wheel ribs across surface in

Opisthosiphon and relatives. Very few species with per-

iostracum consisting of bristle-like projections. Radula

taenioglossate, of Mitten type.

Remarks: In contrast to most annulariids, most abbot-

telline species normally have a nondecollated spire.

But like other annulariids abbottellines may suspend

themselves from mucus strands when at rest. This

behavior has been found in numerous annulariid

groups from many localities. It is not found in the sister

group Pomatiidae and it must be a very ancient be-

havioral trait unique to the Annulariidae.

Key to Hispaniola Genera

1) Shell sculptureless or having only weak spiral cords in

patan loi lheiiliey Bart ea 5 Mania a eet call hhh ade ada eo 2

1) Sloe tl) aatiamar Gyettyl o el Ue ll ee e dae late ad oe 3

2) Outer lip narrowly expanded ......... Leiabbottella

2) Outer lip widely expanded ........... Preclaripoma

3) Axial sculpture of widely spaced, prominent, erect,

Re slices Lanes ey 1 iia t Pat eel Stok: Meganiphe

DB) Amar ectniire MOL asasiye LF) .< tx Adina. 28 19 4

4) SeuIplure spiny OF Serrated ost ih i... 408s ees bog eae 5

AONE IO Be BORE (2 a Pane. aa Bie ef. a's * ayes 6

5) Shell nearly planiepiral www, or) .haew Hs. ie Abbottella

Dt Shela bined: es ALB! Abbottipoma

6) Outer lip notched and enrolled ......... Lagopoma

SCN ORS Met [C\/ nh Oe, ae ee a ees

7) Axial sculpture tightly packed, forming almost

continuous layer, with prominent spiral keels

Wael. fae” SE eee, Microabbottella

THE NAUTILUS, Vol. 134, No. 1

FOTO OcIUNTe UCL \C2e 6: ON clare WC ay a a gn Gee eS 8

8) Spiral sculpture weaker than axial sculpture .......

Nice beet MEER rpc ler, neater asi AM, we « Rolleia

8) Spiral sculpture stronger than axial sculpture .......

Roe REE Se ARR ot RUT, Meine Pane iene Arenabbottella

Genus Abbottella Henderson and Bartsch, 1920

Type Species: Chondropoma moreletianum Crosse,

1873, by original designation.

Description: Shell small (ca. 8-15 mm in diameter),

depressed to planispiral, widely-umbilicate. Non-

decollate. Widely expanded, often fimbriated or un-

dulating outer lip, auriculate. Axial and spiral sculpture

forming sharp cusps or serrations at their intersec-

tions. Multispiral operculum with erect or reflected,

single calcified spiral lamella.

Distribution: Central Dominican Republic from the

south coast to the Majugual area and the southern shore of

Samana Bay (but apparently absent from the Samana

Peninsula); only a single species has been found in Haiti

(Figure 18).

Remarks: An undescribed species is known from a single

specimen from San Pedro de Mecoris.

Etymology: William Louis Abbott (1860-1936), Amer-

ican physician, ornithologist, field naturalist, and patron of

the United States National Museum, now National Mu-

seum of Natural History (USNM).

Key to Species

1) Sculpture of very fine spiral cords and axial lamellae,

forming minute denticies, Maiti... ..2...... haitensis

1) Sculpture coarse, forming denticles or thorny points,

Iowa meaneheptNG hemos nade’... ss: 2

2) Final whorl with 40-80 axial lamellae, sculpture mi-

VVUS) Waele 210 eee Meee ey ee domingoensis

2) Final whorl <50 axial lamellae, sculpture thorny .... 3

3) Final whorl! with 30-40 axial lamellae, spiral opercular

lamella reflected back over base ......... craetegus

3) Final whorl! with 40-50 axial lamellae, spiral opercular

lamellatoblique jit erect. j.4) 452 -. moreletiana

Abbottella crataegus Watters, 2016

(Figures 18, 29-28)

CCHRESONYMY

Abbottella crataegus Watters, 2016: 116-117, figs.

aaa 29.

Type Material: Holotype: UF 216138; Paratypes: UF

216138(5), from the type locality.

G.T. Watters et al., 2020

Type Locality: “5 km W of Majagual, Monte Plata

Province, Dominican Republic.”

Distribution and Habitat: Known only from the

limestone outcrops near Majugual and Guaragao. This is

the western part of the Cevicos Formation composed of

karsted, massive limestone of upper Miocene—lower

Pliocene age (Draper and Lewis, 1991). All specimens

seen were found between 100—260 m elevation. Entire

distribution may be <80 km”. Snails live on mesic

limestone outcrops.

Etymology: L. crataegus, a flowering thorny plant, a

masculine noun in apposition.

Abbottella domingoensis Bartsch, 1946

(Figures 18, 29-36)

CHRESONYMY

Abbottella moreletiana gabriella Bartsch, 1946: 146, 148,

pl. 25, figs. 4-6; Watters, 2006: 362.

Abbottella moreletiana wetmorei Bartsch, 1946: 146, 148,

pl. 26, figs. 8-10.

Abbottella moreletiana domingoensis Bartsch, 1946: 147,

pl. 25, figs. 1-3; Watters, 2006: 361-362.

Abbottella (Abbottella) moreletiana domingoensis (Crosse,

1873): Watters, 2006: 362.

Abbottella (Abbottella) moreletiana gabriella Bartsch,

1946: Watters, 2006: 83, 362; Watters, 2016b: 114-116

[in synonymy of A. domingoensis |.

Abbottella (Abbottella) moreletiana wetmorei Bartsch,

1946: Watters, 2006: 83, 362.

Abbottella (Abbottella) domingoensis (Crosse, 1873):

Watters, 2016b: 114-116, figs 9-16, 30.

Type Material: Abbottella moreletiana domingoensis

Bartsch, 1946: Bartsch (1946) listed USNM 504118 as

the holotype; this is now USNM 504117. Abbottella

moreletiana gabriella Bartsch, 1946: Holotype: USNM

504124. Abbottella moreletiana wetmorei Bartsch,

1946: Holotype: USNM 504119.

Type Locality: Abbottella moreletiana domingoensis

Bartsch, 1946: ““Santo Domingo,” probably Santo Domi-

ngo City.” Abbottella moreletiana gabriella Bartsch, 1946:

“San Gabriel Isle, Samana Bay.” Abbottella moreletiana

wetmorei Bartsch, 1946: “Pelican Keys, San Lorenzo Bay,

Samana Bay, Dominican Republic.”

Distribution and Habitat: This species is found in the

eastern end of the Cordillera Septentrional and the

northern Cordillera Oriental in the Rio Yuna valley and

its tributaries; also in the low hills of the Los Ranchos

Formation in the Sierra de El Seibo extending south to

Santo Domingo and Boca Chica. It has been found under

rocks and rubble on mesic limestone ridges where it is

Page 9

locally common. All specimens seen were found below

320 m elevation.

Remarks: Abbottella moreletiana gabriella Bartsch, 1946,

is known only from San Gabriel Isle, just east of Boca del

Infierno, possibly from the San Gabriel Cave (Angel Cave).

Abbottella moreletiana wetmorei Bartsch, 1946, is known

from the Pelican Keys. Both are here regarded as stunted

forms of A. domingoensis. The name Abbottella domingoensis

Bartsch, 1946, is here chosen as the valid name despite the

page precedence of these two forms.

Etymology: Abbottella moreletiana domingoensis

Bartsch, 1946: from Santo Domingo. Abbottella

moreletiana gabriella Bartsch, 1946: from San Gabriel

Isle. Abbottella moreletiana wetmorei Bartsch, 1946:

Frank Alexander Wetmore (1886-1978), American orni-

thologist at Smithsonian Institution.

Abbottella haitensis Bartsch, 1946

(Figures 18, 37-39)

CHRESONYMY

Abbottella haitensis Bartsch, 1946: 143, 145-146, pl. 24,

figs. 10-12.

Abbottella (Abbottella) haitensis Bartsch, 1946: Watters,

2006-783) 287-255.

Type Material: Holotype: USNM 504111.

Type Locality: “|Haiti] on the road to Grand Bois, 4

miles north of Thomazeau, at an elevation of 1,000 feet.”

We have not located this Grand-Bois.

Distribution and Habitat: Known only from the type

locality without habitat information.

Remarks: This species is a mystery. It occurs so far from

other Abbottella that it is tempting to believe it was

mislocalized. However it was collected by the eminent field

naturalist W.L. Abbott. The holotype label reads “1920” and

Abbott was known to have collected in Haiti in 1920.

It is also possible this species belongs to a different

genus. The pustulose sculpture is rather different from

the spinose sculpture of the other Abbottella. The species

also has some characteristics of Rolleia. Do not confuse

with the similarly named Rolleia haitensis Bartsch, 1946.

Preserved material was not available.

Etymology: William Louis Abbott (1860-1936),

American physician, ornithologist, field naturalist, and

patron of the United States National Museum, now

National Museum of Natural History (USNM).

Abbottella moreletiana moreletiana (Crosse, 1873)

(Figures 18, 40-43, 46)

Page 10 THE NAUTILUS, Vol. 134, No. 1

Figures 22-39. Abbottella. 22-28. Abbottella crataegus Watters, 2016. 22-24. Holotype, UF 216138 (12.0 mm diameter). 25.

GTW 16492b (11.0 mm diameter). 26. Live individual (photo S. Aiken ©). 27. Mating pair (photo ©S. Aiken). 28. Distribution map.

29-36. Abbottella domingoensis Bartsch, 1946. 29, 30. Abbottella moreletiana domingoensis Bartsch, 1946. Holotype, USNM 504117

(12.2 mm diameter). 31, 32. UF 216133 (15.2 mm diameter). 33. Abbottella moreletiana gabriella Bartsch, 1946: Holotype, USNM

504124 (8.6 mm diameter). 34. Abbottella moreletiana wetmorei Bartsch, 1946: Holotype, USNM 504119 (7.0 mm diameter). 35.

Mating pair (photo S. Aiken ©). 36. Distribution map. 37-39. Abbottella haitensis Bartsch, 1946. 37, 38. Holotype, USNM 504111

(12.7 mm diameter). 39. Distribution map.

G.T. Watters et al., 2020

CHRESONYMY

Choanopoma moreletiana Crosse, 1873: 354; Crosse,

1874: 85, pl. 3, figs. 3, 3a; Pilsbry, 1933: 130; Watters,

2006:361.

Choanopoma moreletianum Crosse, 1873: Pfeiffer, 1876:

160-161; Kobelt, 1880: 277; Crosse, 1891: 168.

Tudorae |sic| moreletianae |sic] (Crosse, 1873): Arango,

1884: 211.

Abbottella moreletiana (Crosse, 1873): Henderson and

Bartsch, 1920: 75; Thiele, 1931: 134; Abbott, 1989: 53;

Watters, 2016b: 112, 114, figs. 1-8, 29.

Choanopoma (Abbotella |sic]) moreletianum Crosse,

1873: Clench and Aguayo, 1937: 67.

Chondropoma moreletianum (Crosse, 1874): Bartsch,

1946: 143.

Abbottella moreletiana moreletiana (Crosse, 1873):

Bartsch, 1946: 147, pl. 25, figs. 10-12.

Abbottella (Abbottella) moreletiana moreletiana (Crosse,

1873): Watters, 2006: 361-362.

Type Material: Holotype: ANSP 14166, not figured in

online database; from the Bland collection, collected by

Gabb.

Type Locality: “In regione Dominicanum insulae Haiti,

Antillarum.” Restricted by Watters (2016b) to “Sabana de

la Mar, Hato Major Province, Dominican Republic.”

Distribution and Habitat: It occurs on the southern

side of Samana Bay, including the nearshore Lower and

Upper Orange keys, from approximately Boca del Infierno

to Sabana de la Mar in Samana and Hato Major provinces

in the Los Haitises Limestone. This corresponds to the

eastern part of the Cevicos Formation, composed of

Page 11

karsted, massive limestone blocks of Upper Miocene-

Lower Pliocene age (Lebron and Mann, 1991). It is found

in mesic forests in association with wet limestone cliffs,

where it may be locally abundant. Its range may be

<100 km”.

Etymology: Named after Pierre Marie Arthur Morelet

(1809-1892), French naturalist and conchologist working

in Central America, Africa, and Europe.

Abbottella moreletiana kriegeri Bartsch, 1946

(Figures 44-46)

CHRESONYMY

Abbottella moreletiana kriegeri Bartsch, 1946: 146, pl. 24,

figs. 4-6.

Abbottella (Abbottella) moreletiana kriegeri Bartsch,

1946: Watters, 2006: 83, 362; Watters, 2016b: 112.

Type Material: Holotype: USNM 504115.

Type Locality: “North side of the Samana Peninsula on

the center of the south side of San Juan Bay, Dominican

Republic.” Based on Gabb’s 1873 description and map,

this apparently is the bay at Playa de Limon north of El

Limén. However, Abbottella is not known from the

Samana Peninsula and this locality remains suspicious.

Distribution and Habitat: Known only from the pos-

sibly erroneus type locality without habitat infromation.

Remarks: We suspect this is a stunted form of A. mor-

eletiana and the actual provenance is the south side of

Figures 40-46. Abbottella. 40-43. Abbottella moreletiana moreletiana (Crosse, 1873). 40. Choanopoma moreletiana Crosse, 1873.

Crosse, 1874: 85, pl. 3, figs. 3, 3a. 41-43. UF 216190 (12.3 mm diameter). 44, 45. Abbottella moreletiana kriegeri Bartsch, 1946.

Holotype, USNM 504115 (10.0 mm diameter). 46. Distribution map.

Page 12

Samana Bay. Further collections will be needed to verify

this. Preserved material was not available.

Etymology: Herbert William Krieger (1889-1970),

American anthropologist at the Smithsonian Institution,

collector of the type.

Genus Abbottipoma new genus

Type Species: Abbottella abbotti Bartsch, 1946.

Description: Shell small (ca. 6-10 mm in diameter),

turbinoid, with spiral keels and minutely serrated or spiny

sculpture, outer lip expanded and/or fimbriated or en-

rolled. Operculum as in family.

Distribution: Endemic to the eastern half of the Samana

Peninsula in mesic areas with limestone outcrops (Figure 18).

Etymology: Abbottella abbotti, the type species +

-poma, a standard suffix for many annulariids.

Key to Species

1) Sculpture of coarse spines, outer lip deeply fimbriated

Eeablet a, wlan rhe WAN A 2 eld dean ee hs tee abbotti

1) Sculpture of minute serrations, outer lip not deeply

EESTI ON CES (2 0 ML pie kee a ae COO Pere ey ERY erg ee

2) Anterior margin of outer lip enrolled ......... gabbi

2) Anterior margin of outer lip not enrolled .... crossei

Abbottipoma abbotti (Bartsch, 1946)

(Figures 18, 47-54)

CHRESONYMY

Abbottella abbotti Bartsch, 1946: 143, 154-155, pl. 27,

figs. 10-12.

Abbottella (Abbottella) abbotti Bartsch, 1946: Watters,

2006: 83, Watters, 2013: 11, figs. 3 A-E, 7 F.

Type Material: Holotype: USNM 504108. The type is a

worn specimen that does not show the characteristic spiny

sculpture.

Type Locality: “Near Laguna, Samana Bay” [? Punta

Laguna Salada, probable error for Samana Peninsula, see

Watters, 2013].

Distribution and Habitat: This narrowly endemic

species occurs in the Sierra de Samana of the Cordillera

Septentrional in the E] Valle and the Las Galeras areas on

the northeast coast of the peninsula. A record from Cayo

Levantado (GTW 10829a) seems unlikely and needs

confirmation. Snails live below 50 m elevation on and at

the base of limestone walls in cool, shaded areas with

THE NAUTILUS, Vol. 134, No. 1

abundant mosses and other vegetation. It may occur

in <10 km”. At El Valle it lives within a few hundred

meters of the beach in association with Lagopoma lago-

poma, Helicina salleana Pfeiffer, 1852, and the very large

pleurodontid Caracolus excellens (Pfeiffer, 1853).

Remarks: This species has been observed suspending

itself by a mucus thread(s) from limestone outcrops.

Etymology: William Louis Abbott (1860 —1936), Ameri-

can physician, ornithologist, field naturalist, and patron of

the United States National Museum, now National Mu-

seum of Natural History (USNM).

Abbottipoma crossei (Pilsbry, 1933)

(Figures 18, 55-62)

CCHRESONYMY

Choanopoma crossei Pilsbry, 1933: 130, pl. 7, figs. 5, 5a, 6.

Abbottella crossei (Pilsbry, 1933): Bartsch, 1946: 144, 153,

pl. 27, figs. 13-15.

Abbottella gabbi pilsbryi Bartsch, 1946: 152-153, pl. 27,

figs. 7-9.

Hales crossei (Pilsbry, 1933): Baker, 1964: 169.

Abbottella (Abbottella) crossei (Pilsbry, 1933): Watters,

2006: 83, 222; Watters and Duffy, 2010a: 2; Watters,

2013: 14, figs. 3 S-W, 7 G.

Abbottella (Abbottella) gabbi pilsbryi Bartsch, 1946:

Watters, 2006: 83, 269.

? Abbottella (Abbottella) harpeza Watters and Duffy,

2010a: 1-2, figs. 1-3.

Type Material: Choanopoma crossei Pilsbry, 1933:

Holotype: ANSP 7951; Paratype: ANSP 373769(1). The

type lot of A. gabbi contained two species, gabbi and

crossei. Pilsbry (1933) designated Crosse’s figured spec-

imen of the lot as the type of A. gabbi. Abbottella gabbi

pilsbryi Bartsch, 1946: Holotype: USNM 504099.

Abbottella harpeza Watters and Duffy, 2010: Holotype:

UF 420731.

Type Locality: Choanopoma crossei Pilsbry, 1933:

“Santo Domingo.” Restricted by Watters (2013) to “4 km

E of Samana, Samanaé Province, Dominican Republic.”

Abbottella gabbi pilsbryi Bartsch, 1946: “in a cave on the

Rio Seco near Samana, Samana Bay, Dominican Re-

public.” Abbottella harpeza Watters and Duffy, 2010: “Isla

Beata,” probably in error for Samana Peninsula.

Distribution and Habitat: This species inhabits the

coastal limestone bluffs of the eastern Samana Peninsula

in the Sierra de Samana including Cayo Levantado.

Specimens have been found on limestone outcrops in

fields and on cliffs and stone walls, often abundant; oc-

casionally arboreal. All specimens seen were found below

50 m elevation.

G.T. Watters et al., 2020 Page 13

Figures 47-70. Abbottipoma. 47-54. Abbottipoma abbotti (Bartsch, 1946). 47, 48. Holotype, USNM 504108 (6.0 mm diameter).

49-51. GTW 10829b (7.1 mm diameter). 52. Mating pair. 53. Live individual. 54. Distribution map. 55-62. Abbottipoma crossei (Pilsbry,

1933). 55, 56. Choanopoma crossei Pilsbry, 1933. Holotype, ANSP 7951 (7.5 mm diameter, photo courtesy of ANSP). 57, 58. Abbottella

gabbi pilsbryi Bartsch, 1946. Holotype, USNM 504099 (7.5 mm diameter). 59. Abbottella (Abbottella) harpeza Watters and Dutty, 2010.

Holotype, UF 420731 (7.1 mm diameter). 60, 61. GTW 9432a (7.3 mm diameter). 62. Distribution map. 63-70. Abbottipoma gabbi

(Crosse, 1873). 63. Choanopoma gabbi Crosse, 1873. Crosse, 1874: 84, pl. 3, fig. 2. 64-66. Syntype, MNHN 2000-5452 (7.4 mm diameter,

photo courtesy of Manuel Caballer, MNHN). 67-69. Julian Joseph coll. (6.3 mm diameter, photos ©S. Aiken). 70. Distribution map.

Page 14

Remarks: Abbottella harpeza Watters and Duffy,

2010, was described from Isla Beata. This now seems

very unlikely, being far out of the range of all other

abbottellines, and probably represents a mislocalized

specimen. Additional study at Isla Beata is needed to

confirm this.

Etymology: Choanopoma crossei Pilsbry, 1933: Joseph

Charles Hippolyte Crosse (1826-1898), French con-

chologist, editor of the Journal de Conchyliologie.

Abbottella gabbi pilsbryi Bartsch, 1946: Henry Augustus

Pilsbry (1862-1957), American conchologist, Academy of

Natural Sciences of Philadelphia. Abbottella harpeza

Watters and Duffy, 2010: L. harpeza, a thorny bush, a

thicket.

Abbottipoma gabbi (Crosse, 1873)

(Figures 18, 63-70)

CHRESONYMY

Choanopoma gabbi Crosse, 1873: 353-354; Crosse, 1874:

84, pl. 3, fig. 2; Pfeiffer, 1876: 159; Kobelt, 1880: 277;

Crosse, 1891: 168; Henderson and Bartsch, 1920: 75:

Pilsbry, 1933: 129-130, pl. 7, figs. 1-4; Fischer-Piette,

1950: 80.

Abbottella gabbi (Crosse, 1873): Henderson and Bartsch,

1920; 75:

Abbottella gabbi gabbi (Crosse, 1873): Bartsch, 1946: 144,

152, pl. 26, figs. 5-7.

Abbottella (Abbottella) gabbi gabbi (Crosse, 1873): Wat-

ters, 2006: 83, 269.

Type Material: Syntype: MNHN 2000-5452 (Only

one specimen mentioned in database, number of ad-

ditional specimens unknown). This specimen is a worn

example.

Type Locality: “in regione Dominicanum insulae Haiti,

Antillarum.” Restricted here to “near E] Salto del Limén,

central Samana Peninsula, Samana Province, Dominican

Republic.”

Distribution and Habitat: This unlocalized species was

recently rediscovered after nearly 150 years at El Salto del

Lim6n in the central Samana Peninsula. It was found in

“soil/leaf litter around coral/limestone boulders along the

edge of the forest path” to the waterfall (J. Joseph, 2017,

pers. comm. ).

Remarks: The combination of the enrolled anterior

portion of the outer lip, purple apex, and strong, spiral,

fimbriated carinations is unique. Preserved material was

not available.

Etymology: William More Gabb (1839-1878), American

geologist and paleontologist who surveyed in Hispaniola.

THE NAUTILUS, Vol. 134, No. 1

Genus Arenabbottella new genus

Type Species: Abbottella sosuaensis Bartsch, 1946.

Description: Shells small (6-11 mm in diameter), tur-

binoid, sculpture of minute pustules or weak serrations on

more-or-less well-developed spiral cords often forming

weak keels, cords often prominent in umbilicus, suture

channeled, outer lip expanded, often auriculate. Lip may

be folded or rolled abaperturely to varying degrees.

Sculpture is often arranged in axial “blocks” probably

representing growth lines.

Distribution: This is the most widely distributed abbot-

telline group, occurring around the eastern half of the

island on the northern slopes of the Cordillera Septen-

trional from Luperon to the Samana Peninsula and around

the southeastern lowlands from Las Lagunas de Nisibon to

San Cristobal, including Isla Saona (Figure 19). Snails live

under leaf litter and limestone rubble, from open fields to

mesic forests.

Remarks: This is the most speciose of the Hispaniolan

abbottelline genera. Based on conchological features and

distributional patterns, there seems to be a split below

Samana Bay between a northern and a southern group of

species but none of the northern species were available for

phylogenetic study. These two groups may yet be found to

constitute different genera.

Etymology: L. arenosus, full of sand + Abbottella; in

reference to the sandy sculpture of the shells.

Key to Species

The questionable taxa Choanopoma adolfi Pfeiffer,

1852, and Cyclostoma tentorium Pfeiffer, 1850, are not

included.

1) Columellar side of outer lip enrolled... samanaensis

1) Columellar side of outer lip not enrolled ........ 2

2 Outer No Memomated "tamer can akan-a 8: rosaliae

2) vine mle Net nermio ten mies. cence 2 s,m. 3

3) Shell with a prominent central, serrated keel .........

LG se tbe’ 5 ee ealted Se ene a he RR NR ce Wah calliotropis

S) sliell wathout-a prominent keel | iaee (tue - 4

4) ‘Outer lip narrowly expanded: <i.4i2:.5. 82 asian. - 5

Mistery. wide Wexpanded: aqt.cek eee ee. 8

5) Sculpture nearly obsolete, shell with a metallic sheen

a ae eae a sales Vataicon ecb rata aot GRAS co Cis EE ad eo es nitens

Dec OMe WM as OVE 5 nan pak cet cen + ayaa ates Sat 6

6) Final whorl with ca. 120 axial lamellae, shell usually

‘00? Ae eh eae ang al Gicgte hue R Lets ERea wk es cae milleacantha

6) Final whorl with <100 axial lamellae, shell usually

BOMOOMREGIOVE 5... van wuts veer e oe kaha f

G.T. Watters et al., 2020

8) Sculpture nearly obsolete except for umbilicus, shells

rather smooth: | bo) sapye ue ene =e Ree 9

9) Strong spiral cords present in umbilicus ........ 10

10) Shell pale yellow with brown spots, suture with white,

TANCOMOCeme les). a) eae ect ek een oe mellosa

10) Shell brown without spots, suture lacking white,

TAMCOUM COMICS Ta 4. ennui hey Wai ee newcombi

Strong spiral cords absent from umbilicus ...... 12

Strong spiral cords present in umbilicus ........ 13

11)

12) Shell pale brown or yellow, sculpture fine .... wilhelmi

12)

Shell dark brown or white, sculpture coarse .......

13) Spiral cords not all of uniform strength, outer lip

RAC Gee Goro g gir as eo aman, 2 anchezi

13) Spiral cords approximately the same strength, outer

lo marowlemmatded: 2n aun Gee sosuaensis

Arenabbottella adolfi (Pfeiffer, 1852)

(Figures 19, 71, 72)

CHRESONYMY

Choanopoma adolfi Pfeiffer, 1852a: 167; Pfeiffer, 1853:

117-118; Adams and Adams, 1856: 296; Hjalmarson

and Pfeiffer, 1858: 139: Pfeiffer, 185Sb: 102; Kobelt,

1880: 277; Crosse, 1891: 166.

Cyclostoma (Choanopoma) adolfi (Pfeiffer,

Pfeiffer, 1854a: 142.

Cyclostoma adolfi (Pfeiffer, 1852): Pfeiffer, 1854c: pl. 48,

figs. 5-8; Pfeiffer, 1854d: 371.

Choanopoma adolphi |sic| Pfeiffer, 1852: Bland, 1861:

355; Pfeiffer, 1865: 111; Pfeiffer, 1876: 161.

Cyclostoma adolphi |sic| (Pfeiffer, 1852): Reeve, 1862: pl.

22, fig. 146.

Abbottella adolphi |sic| (Pfeiffer, 1852): Henderson and

Bartsch, 1920: 75.

Abbottella adolphi adolphi |sic| (Pfeiffer, 1852): Bartsch,

1946: 143, 149-150, pl. 26, fig. 4.

Abbottella (Abbottella) adolft adolfi (Pfeiffer, 1852):

Watters, 2006: 83, 130.

Abbottella adolft (Pfeiffer, 1852): Watters and Duffy,

201.0a:-2.

non Abbottella (Abbottella) cf. adolft adolfi (Pfeiffer, 1852):

Watters, 2013: 13, figs. 3 F-L 9 D [= A. mellosa].

1852):

Type Material: The type has apparently been lost. Many

of Pfeiffer’s types were probably destroyed by Allied

Page 15

bombing of the Museum Naradone in Szczecin, Poland, in

WW ILI.

Type Locality: “Habitat in insula Haiti.”

Distribution and Habitat: Known only from the unlo-

calized type locality without habitat information.

Remarks: Pfeiffer (1854: pl. 48, figs. 5-8) and Reeve

(1862: pl. 22, fig. 146) illustrated it but it cannot be

confidently identified from these figures. It is here

considered a nomen dubium.

Etymology: Probably Emil Adolf Rossmassler (1805?-1867),

German conchologist.

Arenabbottella aenea (Watters, 2010)

(Figures 19, 76—79)

CHRESONYMY

Abbottella (Abbottella) aenea Watters, 2010: 16-17, text fig.,

pl. 1, figs. 1-4; Watters, 2013: 13, figs. 3 N-R, 8 C.

Type Material: Holotype: UF 434777; Paratype: UF

434778(1); Paratypes: BMSM_ 17971(2); Paratypes:

OSUM 35490(2), all from the type locality.

Type Locality: “Dominican Republic, La Altagracia

Province, Punta Cana.”

Distribution and Habitat: Known only from the type

locality. Specimens were found under moldy leaf litter.

Remarks: Preserved material was not available.

Etymology: L. aeneus, copper.

Arenabbottella calliotropis (Watters, 2013)

(Figures 19, 80-83)

CHRESONYMY

Abbottella (Abbottella) calliotropis Watters, 2013: 3, figs. 1

Rate BO 7 ea.

Type Material: Holotype: UF 456810; Paratypes: UF

216131(59), from the type locality.

Type Locality: “Dominican Republic, La Altagracia

Province, along Rio Yuna, 2 km S of La Guana. ca. 18.79°

IN GGT” We”

Distribution and Habitat: Under limestone rubble

in fields on the easternmost outcrops of the cordillera

in La Altagracia Province in the area of Higiiey and Las

Lagunas de Nisibon.

Page 16 THE NAUTILUS, Vol. 134, No. 1

Figures 71-91. Arenabbottella. 71, 72. Arenabbottella adolft (Pfeiffer, 1852).'71. Choanopoma adolfi Pfeiffer, 1852. Pfeiffer, 1854c:

pl. 48, fig. 8. 72. Cyclostoma adolphi [sic] (Pfeiffer, 1852): Reeve, 1862: pl. 22, fig. 146. 73. Arenabbottella newcombi (Crosse, 1873).

Syntype, MNHN 5436 (photo courtesy of MNHN). 74, 75. Arenabbottella tentorium (Pfeiffer, 1850). 74. Cyclostoma tentorium

Pfeiffer, 1850. Pfeiffer, 1854b: pl. 38, fig. 18. 75. Cyclostoma tentorium Pfeiffer, 1850. Reeve, 1862: pl. 22, fig. 145. 76-79. Are-

nabbottella aenea (Watters, 2010). 76-78. GTW 14181a (6.5 mm diameter). 79. Distribution map. 80-83. Arenabbottella calliotropis

(Watters, 2013). 80-82. Holotype, UF 456810 (7.5 mm diameter). 83. Distribution map. 84-87. Arenabbottella dichroa (Watters,

2013). 84-86. Holotype, UF 456801 (9.9 mm diameter). 87. Distribution map. 88-91. Arenabbottella mellosa (Watters and Dutfty,

2010). 88-90. GTW 7020c (5.0 mm diameter). 91. Distribution map.

G.T. Watters et al., 2020

Remarks: Preserved material was not available.

Etymology: Gr. kallos, beauty + Gr. trope, a turning.

Arenabbottella dichroa (Watters, 2013)

(Figures 19, 84-87)

CHRESONYMY

Abbottella (Abbottella) dichroa Watters, 2013: 6-7, figs. 1

K-@75-@. 771.

Type Material: Holotype: UF 456801; Paratypes: UF

216112(31), from the type locality.

Type Locality: “Dominican Republic, Samana Province,

just E of El Limon, 10 km E of Las Terrenas, at sea level.

ca. 19.29° N, -69.44° W.”

Distribution and Habitat: Known from the north coast

of the Sierra de Samana of the Cordillera Septentrional

from the western Samana Peninsula to Nagua on lime-

stone hills in mesic, shaded forests.

Remarks: This species is peculiar in having two color

morphs in the same population: white or brown. This may

be due to sexual dimorphism. Preserved material was not

available.

Etymology: Gr. di-, two + Gr. chroa, color of the skin; in

reference to the two color morphs.

Arenabbottella mellosa (Watters and Duffy, 2010)

(Figures 19, 88-91)

CHRESONYMY

Abbottella (Abbottella) mellosa Watters and Duffy, 2010a:

2, figs. 4-6; Watters, 2013: 8, fig. 1 T.

Abbottella (Abbottella) cf. adolft adolfi (Pfeiffer, 1852):

Watters, 2013: 13, figs. 3 F-I, 9 D.

Type Material: Holotype: UF 420729; Paratype: OSUM

32477(1), from the type locality.

Type Locality: “Dominican Republic, Los Brazos, near

Sostia.”

Distribution and Habitat: Known only from three adjacent

outcrops: Loma Catalina and Loma Blanca near Sostia and

the outcrop near Rio San Juan on the northern coast. All

specimens seen were found below 70 m elevation.

Remarks: Rare, well-preserved specimens have perios-

tracal hairs.

Etymology: L. mellosa, honey-colored.

Page 17

Arenabbottella milleacantha (Watters and

Duffy, 2010)

(Figures 19, 92-95)

CCHRESONYMY

Abbottella (Abbottella) milleacantha Watters and Duffy,

2010a: 2-3, figs. 7-9; Watters, 2013: 4, 7, 13, 14, figs. 3

XB BR 77H:

Type Material: Holotype: UF 420728; Paratype: OSUM

32478(1), from the type locality.

Type Locality: “Dominican Republic, northeast Isla

SaAoOna

Distribution and Habitat: This species occurs on Isla

Saona and the adjacent coast from San Pedro de Macoris

to Bavaro in sub-mesic to mesic forests along limestone

ridges under blocks and in leaf litter; also in caves.

Remarks: Preserved material was not available.

Etymology: L. mille, thousand + Gr. akantha, thorn,

prickle.

Arenabbottella newcombi (Crosse, 1873)

(Figure 73)

CHRESONYMY

Choanopoma newcombi Crosse, 1873: 352-353; Crosse,

1874: 82, pl. 3, figs. 1, la; Pfeitter, 1876: 160; Kobelt,

1880: 277: Crosse, 1891: 167-168; Fischer-Piette,

1950: 79-80.

Abbottella newcombi (Crosse, 1873): Henderson and

Bartsch, 1920: 75; Bartsch, 1946: 143, 144-145, pl. 24,

figs. 7-9.

Abbottella (Abbottella) newcombi (Crosse, 1873): Watters,

YO0G. 83, 31 ls Watters, 2013) 13.

Type Material: Syntype: MNHN 5436.

Type Locality: “in regione Dominicanum insulae Haiti,

Antillarum.”

Distribution and Habitat: Known only from the unlo-

calized type locality.

Comparison with Other Species: Based on two spec-

imens, this is a relatively smooth, brown shell with a deep

suture and lacking keels.

Remarks: Bartsch’s (1946) specimens were received

from Newcomb but with only “Santo Domingo” as their

provenance, undoubtedly referring to the country rather

than the city. Although clearly a member of this group it

has not been collected since its description.

THE NAUTILUS, Vol. 134, No. 1

Figures 92-113. Arenabbottella. 92-95. Arenabbottella milleacantha (Watters and Duffy, 2010). 92-94. Holotype, UF 420728

(7.5 mm diameter). 95. Distribution map. 96-99. Arenabbottella nitens (Watters, 2013). 96-98. Holotype, UF 456806 (7.8 mm

diameter). 99. Distribution map. 100-103. Arenabbottella rosaliae (Pfeiffer, 1858). 100-102. UF 216191 (9.2 mm diameter). 103.

Distribution map. 104-107. Arenabbottella samanensis (Bartsch, 1956). 104, 105. Holotype, USNM 504090 (10.6 mm diameter).

106. GTW 7085c (8.7 mm diameter). 107. Distribution map. 108-113. Arenabbottella sanchezi (Bartsch, 1946). 108. Abbottella

anchezi Bartsch, 1946. Holotype, USNM 504092 (9.1 mm diameter). 109. Abbottella adolphi peninsularis Bartsch, 1946. Holotype,

USNM 504095 (9.8 mm diameter). 110-112. UF 216154 (9.1 mm diameter). 113. Distribution map.

G.T. Watters et al., 2020

Etymology: Wesley Newcomb (1808-1892), American

physician, conchologist, Hawaiian temperance leader.

Arenabbottella nitens (Watters, 2013)

(Figures 19, 96-99)

CHRESONYMY

Abbottella (Abbottella) nitens Watters, 2013: 7, figs. 1 P-S,

5 Ry Seok

Type Material: Holotype: UF 456806; Paratypes: UF

456808(15), from the type locality.

Type Locality: “Dominican Republic, La Altagracia

Province, 16 km S of Higiiey, at 100 m. ca. 18.46° N,

— 68.715 We

Distribution and Habitat: Known only from the type

locality under rocks among boulders, on a limestone bluff

west of Highway 4.

Remarks: Preserved material was not available.

Etymology: L. niteo, shine, glitter, in reference to the

metallic sheen of most specimens.

Arenabbottella rosaliae (Pfeiffer, 1858)

(Figures 15, 19, 100-103)

CHRESONYMY

Choanopoma rosaliae Pfeiffer in Hjalmarson and Pfeiffer,

1858: 139-140; pl. a figs. 4-6: Bland, 1861: 355;

Pfeiffer, 1865: 111; Pfeiffer, 1876: 161; Kobelt, 1880: 277;

Crosse, 1891: 166; Pilsbry, 1933: 130.

Abbottella rosaliae (Pfeiffer, 1858): Henderson and

Bartsch, 1920: 75; Bartsch, 1946: 144, 155, pl. 27,

figs. 4-6.

Abbottella (Abbottella) rosaliae (Pfeiffer, 1858): Watters,

2006: 83, 447; Watters, 2013: 14-16, figs. A A-OK.

Type Material: Not located, presumed lost.

Type Locality: “In insulae Haiti.” Restricted by Watters

(2013) to “Dominican Republic, Puerto Plata Province,

Loma Catalina.”

Distribution and Habitat: This species is known only

from Loma Catalina, an outcrop of the Cordillera Sep-

tentrional southeast of Sostia, occurring in ca. 100 km”.

Much of this area is contained in the Parque Nacional E]

Choco. All specimens seen were found between 30-280 m

elevation on limestone bluffs.

Remarks: This species possesses a periostracum although

it is lost in most specimens. On fresh examples this

periostracum forms short bristles at the junctures of the

Page 19

axial and spiral sculptures. In general, a periostracum

appears to be very rare in the Annulariidae.

Etymology: Unknown, the identity of Rosalie has been

lost to time.

Arenabbottella samanensis (Bartsch, 1946)

(Figures 19, 104-107)

CHRESONYMY

Abbottella samanensis Bartsch, 1946: 143, 148-149, pl.

25, figs. 7-9.

Abbottella (Abbottella) samanensis Bartsch, 1946: Watters,

2006: 83, 460; Watters, 2013: 16, figs. 4 F-I, 9 B.

Type Material: Holotype: USNM 504090.

Type Locality: “Cape Samana, Dominican Republic.”

Distribution and Habitat: Samand Peninsula in the

Sierra de Samana of the Cordillera Septentrional. Live

snails have been found in leaf litter.

Remarks: Preserved material was not available.

Etymology: From the Samana Peninsula.

Arenabbottella sanchezi (Bartsch, 1946)

(Figures 19, 108-113)

CHRESONYMY

Abbottella sanchezi Bartsch, 1946: 143, 149, pl. 26,

figs. 1-3.

Abbottella adolphi |sic| peninsularis Bartsch, 1946:

150-151, pl. 26, figs. 14-16.

Abbottella (Abbottella) adolft peninsularis Bartsch, 1946:

Watters, 2006: 83, 130; Watters, 2013: 6, 13, figs, 3 J-L,

8 H.

Abbottella (Abbottella) sanchezi Bartsch, 1946: Watters,

2006: 83, 460; Watters, 2013: 6, 8, 13, 16, 17, figs. 4

EN. oC.

Type Material: Abbottella sanchexi Bartsch, 1946: Ho-

lotype: USNM 504092. Abbottella adolphi peninsularis

Bartsch, 1946: Holotype: USNM 504095.

Type Locality: Abbottella sanchexi Bartsch, 1946: “2

miles northwest of SAnchez, Dominican Republic.”

Abbottella adolphi peninsularis Bartsch, 1946: “On the

trail from Samana to Rio San Juan, Samana Peninsula.”

Distribution and Habitat: The species is found in the

Sierra de Samana from at least SAnchez to Los Cacaos in

the south and Las Terrenas in the north. Found in as-

sociation with limestone boulders, leaf litter, and talus in

mesic forests and near caves. All specimens seen were

found below 320 m elevation.

Page 20

Remarks: Bartsch (1946) differentiated A. sanchezi

and A. a. peninsularis based on differences in spire

height. Both fall within the range of variation of a

single species. This species may exhibit color dimor-

phism, probably sexual. Abbottella sanchezi is here

chosen as the valid name for the taxon.

Etymology: Abbottella sanchezxi Bartsch, 1946: from

Sanchez, Dominican Republic. Abbottella adolphi pen-

insularis Bartsch, 1946: from the Samana Peninsula.

Arenabbottella sosuaensis (Bartsch, 1946)

(Figures Os; Lt4—aehk7)

CHRESONYMY

Abbottella sosuaensis Bartsch, 1946: 143, 151, pl. 26, figs.

112.

Abbottella (Abbottella) sosuaensis Bartsch, 1946: Watters,

2006: 83, 489-490; Watters, 2010: 17; Watters, 2013: 6,

17, 18, figs. 4 O-S, 9 E.

Type Material: Holotype: USNM 336768.

Type Locality: “Sostia, 16 miles east of Puerto Plata,

Dominican Republic.”

Distribution and Habitat: This species is found on the

northern slopes of the Cordillera Septentrional from

Luperon to Rfo San Juan under rocks along rolling

limestone hills, pastures, and under limestone rubble at

the base of cliffs; it may be locally common. It does not

occur at El] Valle as reported by Watters (2013). All

specimens seen were found below 100 m elevation.

Etymology: From Sostia, Dominican Republic.

Arenabbottella tenebrosa (Watters, 2013)

(Figures 16113)

CHRESONYMY

Abbottella (Abbottella) tenebrosa Watters, 2013: 6, 8, figs.

1 U-Y, 8 B.

Type Material: Holotype: UF 456796; Paratypes: UF

456797(15); Paratypes: OSUM 37271(12); Paratypes:

BMSM 17937(16), all from the type locality.

Type Locality: “Dominican Republic, Puerto Plata

Province, El] Choco, near Cabarete. ca. 19.74° N,

—{),42° W”

Distribution and Habitat: This species is found from

the limestone outcrops of El Choco near Cabarete to just

west of Rio San Juan. All specimens seen were found

below 20 m elevation.

THE NAUTILUS, Vol. 134, No. 1

Etymology: L. tenebrosus, dark, gloomy, in reference to

the shell color.

PArenabbottella tentorium (Pfeiffer, 1850)

(Figures 74, 75)

CHRESONYMY

Cyclostoma tentorium Pfeiffer, 1850: 77; Pfeifter,

1854b: 284, pl. 38, figs. 16-18; Reeve, 1862: pl. ines

fig. 145.

Choanopoma tentorium (Pfeiffer, 1850): Pfeiffer, 1851:

155; Pfeiffer, 1852a: 168; Pfeiffer, 1852b: 27; Pfeiffer,

1853: 118; Adams and Adams, 1856: 296; Pfeiffer,

1858b: 102; Bland, 1861: 355; Pfeiffer, 1865: 111;

Pfeiffer, 1876: 161; Kobelt, 1880: 277: Crosse,

1891: 167.

Abbottella (Abbottella) tentorium (Pfeiffer, 1850): Watters,

2006: 83, 505.

non Abbottella tentorium (Pfeiffer, 1850): Henderson and

Bartsch, 1920: 75; Bartsch, 1946: 144, 153, pl. 27, figs.

1l— 3 [= PA. wetmorei|.

Type Material: Not located, presumed lost.

Type Locality: “in insula Haiti.”

Remarks: Pfeiffer’s 1850 description of “Interdum

rubro-punctata” does not conform to any known

Abbottella. Reeve’s 1862 figure seems to show this feature

more so than does Pfeiffer’s 1854 figure. At 6.5 mm in

diameter only M. diadema and A. urbana are smaller.

Bartsch’s 1946 specimen (his pl. 27, figs. 1-3) from an

unknown locality does not seem to be the same species

and lacks any red spots; it appears to be a pale specimen

of A. wetmorei. This may possibly be an earlier name for

A. urbana.

Etymology: L. tentorium, a tent, a noun in apposition.

Arenabbottella urbana (Watters, 2012)

(Figures 19, 122-128)

CHRESONYMY

Abbottella (Abbottella) urbana Watters, 2012: 1, 3, figs.

1-3; Watters, 2016: 115-166.

Type Material: Holotype: UF 446061; Paratype: OSUM

36509(1), from the tdype locality.

Type Locality: “Parque Central, off José Contreras

Blvd., N of the Loteria barrio of western Santo Domingo,

Distrito Nacional, Dominican Republic, 18.44 N

—69.97 W.”

Distribution and Habitat: Known from Parque Central/

Parque Mirador del Sur in downtown Santo Domingo

G.T. Watters et al., 2020 Page 21

Figures 114-135. Arenabbottella. 114-117. Arenabbottella sosuaensis (Bartsch, 1946). 114, 115. Holotype, USNM 336768

(7.0 mm diameter). 116. UF 218136 (6.9 mm diameter). 117. Distribution map. 118-121. Arenabbottella tenebrosa (Watters, 2013).

118-120. Holotype, UF 456796 (8.9 mm diameter). 121. Distribution map. 122-128. Arenabbottella urbana (Watters, 2012).

122-124. Holotype, UF 446061 (4.9 mm diameter). 125. Distribution map. 126. Live individual (photos ©S. Aiken). 127. Mating pair

(photos ©S. Aiken). 128. Type locality in Parque Central in Santo Domingo. 129-135. Arenabbottella wilhelmi (Pteitter, 1858). 129.

Choanopoma wilhelmi Pfeiffer in Hjalmarson and Pfeiffer, 1858: pl. 2, fig. 3. 130. UF 216150 (8.0 mm diameter). 131. Live individual

suspended from mucus thread. 132-134. UF 216150 (8.5 mm diameter). 135, Distribution map.

THE NAUTILUS, Vol. 134, No. 1

Figures 136-157. Lagopoma, Leiabbottella, Meganiphe, Microabbottella. 136-141. Lagopoma lagopoma Bartsch, 1946. 136-137.

Holotype, USNM 356198 (5.0 mm diameter). 138. GTW 10081a (9.7 mm diameter). 139. Live individual. 140. Radula (bar=100,).

141. Distribution map. 142-145. Leiabbottella galaxius Watters, 2010. 142-144. Paratype, OSUM 35491 (8.4 mm diameter). 145.

Distribution map. 146-153. Meganiphe rhecta Thompson, 1978. 146-148. Paratype, GTW 8227a (10.6 mm diameter). 149-150.

GTW 11371b (8.0 mm diameter). 151. Distribution map. 152. Live individual (photo ©S. Aiken). 153. Habitat (photo ©S. Aiken).

154-157. Microabbottella diadema (Watters, 2013). 154-156. Holotype, UF 456814 (6.0 mm diameter). 157. Distribution map.

G.T. Watters et al., 2020

(Figure 128), among vacant lots in Boca Chica to the

east, and almost to Peravia Province to the west. It is

likely that this species historically occurred along the

southern extent of Santo Domingo Province, which is

now heavily developed. It still exists in remaining pockets

of suitable habitat scattered throughout the region, in-

cluding local land fills. It has been found under limestone

rubble.

Etymology: L. urbana, of a city.

Arenabbottella wilhelmi (Pfeiffer, 1858)

(Figures 19, 129-135)

CHRESONYMY

Choanopoma wilhelmi Pfeiffer in Hjalmarson and

Pfeiffer, 1858: 139, pl. 2, figs. 1-3; Bland, 1861: 355;

Pfeiffer, 1865: 110-111: Pfeiffer, 1876: 161; Kobelt,

1880: 277; Crosse, 1891: 165-166.

Abbottella wilhelmi (Pfeiffer, 1858): Henderson and

Bartsch, 1920: 75; Bartsch, 1946: 143, 144, pl. 24,

figs. 1-3.

Abbottella (Abbottella) wilhelmi (Pfeiffer, 1858): Watters,

2006: 83, 549; Watters, 2013: 13, 16, 17, figs. ATX. 6 WD.

Type Material: Not located, presumed lost.

Type Locality: “In insulae Haiti.” Pfeiffer continued that

it “was collected under stones in moist places near Puerto

Plata” [German translation].

Distribution and Habitat: This species occurs on the

northern slopes of the Cordillera Septentrional from

Puerto Plata to Nagua, including Pico Isabel de Torres,

Loma Blanca at El Choco, and the north side of the

outcrop between Rio San Juan and Cabrera. Individuals

are found from cloud forests to open, disturbed areas

under limestone rubble.

Remarks: This species has been found suspending itself

from mucus threads (Figure 131).

Etymology: Probably Rudolph Wilhelm Dunker

(1809-1885), German conchologist.

Genus Lagopoma Bartsch, 1946

Type species: Lagopoma lagopoma Bartsch, 1946, by

original designation.

Description: Shells small (ca. 9 mm in diameter),

depressed turbinoid, sculpture of minute pustules on

more-or-less weak spiral cords, cords more prominent

in umbilicus, suture channeled, outer lip expanded,

auriculate. Lip folded posteriorly and conspicuously

notched. Radula as in subfamily (Figure 140).

Page 23

Distribution: Samana Peninsula of Dominican Republic

(Figure 21). Snails live under leaf litter, limestone rubble,

and on wet outcrops in mesic forests.

Remarks: Bartsch (1946) established Lagopoma as a

monotypic genus characterized by a folded and notched

outer lip. The enrolled lip also occurs in Arenabbottella

samanaensis and Abbottipoma gabbi. In this study

Lagopoma was found to be sister to Abbottipoma, and like

that genus is endemic to the Samana Peninsula.

Lagopoma lagopoma Bartsch, 1946

(Figures 21, 136-141)

CHRESONYMY

Lagopoma lagopoma Bartsch, 1946: 142-143, pl. 23, figs.

1-3; Watters, 2006: 92, 324; Watters, 2013: 18, figs. 4

¥—-CC. SE.

Type Material: Holotype: USNM 356198 (2 specimens

in lot, holotype listed as 5.0 mm long specimen).

Type Locality: “Laguna, 4 miles N of Samana, Samana

Province, Dominican Republic.”

Distribution and Habitat: Occurs on the limestone

outcrop of the Sierra de Samana that occupies the north-

eastern-most prominence, from El Valle to Loma Trav-

esada. This species is found in association with A. abbotti.

Comparison with Other Species: This species is

characterized by the peculiar folded auricle and notched

outer lip.

Arenabbottella samanaensis and Abbottipoma gabbi oc-

casionally have a similar auricle but lack the notch.

Etymology: Sp. lago, lake, lagoon + -poma, a standard

ending for annulariid genera. The type was collected at

Laguna.

Leiabbottella Watters, 2010

Type species: Leiabbottella galaxius Watters, 2010, by

original designation.

Description: Shells small (ca. 10 mm in diameter), nearly

planispiral, sculptureless except for microscopic axial

threads and faint spiral ridges in umbilicus, outer lip

narrowly expanded.

Distribution: Endemic to an outcrop of the Villa Trina

Formation of the Rio San Juan complex (Figure 21).

Remarks: In this phylogenetic study Leiabbottella gal-

axius, the type species, was a weakly supported member

of the Arenabbottella clade. Conchologically it is very

dissimilar from Arenabbotella species. At the risk of

Page 24

perpetuating a possibly paraphyletic genus, we maintain

Leiabbottella as a distinct genus pending further study.

However, other smooth- shelled taxa previously assigned

to Leiabbottella (soluta and thompsoni) are unrelated and

are reassigned to the new genus Preclaripoma. Con-

chologically they differ primarily in the greatly expanded

outer lip, which is very narrow in L. galaxius. The lack of

shell sculpture appears to be convergent between the two

groups.

Etymology: G. leios, smooth + Abbottella.

Leiabbottella galaxius Watters, 2010

(Figures 21, 142-145)

CHRESONYMY

Leiabbottella galaxius Watters, 2010: 17-19, pl. 1, figs. 5,

6; Watters, 2013: 10, 18-19, figs. 2 J—L, 8 F.

Type Material: Holotype: UF 434779; Paratype: UF

434780(1); Paratypes: BMSM 17972(2); Paratype: OSUM

35491(1), all from the type locality.

Type Locality: “Dominican Republic, Samana Province,

Samand Peninsula, along Rt. 5 between Santa Barbara

de Samana and Sanchez, in the southern foothills of the

Sierra de Samana.” However, subsequent extensive col-

lections in the area of the original type locality have failed

to find this species. All other records place it exclusively

on an outcrop of the Villa Trina Formation between Rio

San Juan and Cabrera. It seems certain that the original

type locality was a mislabled lot and in error. The type

locality is here corrected to “outcrop 0.5 km NNE of Caro

Clara, Maria Trinidad Sdénchez Province, Dominican

Republic.”

Distribution and Habitat: Endemic to an outcrop of the

Villa Trina Formation of the Rio San Juan complex, an

area < 150 km7. This is undifferentiated limestone, silt-

stone, and marl of Upper Miocene—Lower Pliocene age

(Draper et al., 1991). All specimens seen were found from

sea level to 30 m elevation. At Abreu, individuals were

found within several meters of the sea at <1 m elevation.

Etymology: Gr. galaxius, an allusion to having the ap-

pearance of the Milky Way Galaxy.

Genus Meganiphe Thompson, 1978

Type Species: Meganiphe rhecta Thompson, 1978, by

original designation.

Description: Shell small (8-10 mm in diameter), tur-

binate, whorls adnate, nondecollate. Moderately wide

umbilicus. Axial sculpture of distantly spaced, greatly

expanded, erect and undulating lamellae, ca. 12-14 on

THE NAUTILUS, Vol. 134, No. 1

final whorl. Spiral sculpture absent. Aperture circular, lip

greatly expanded rather evenly but less so facing body

whorl, barely separated from body whorl, composed of

numerous greatly expanded lamellae, gently reflected

abaperturally, undulating, weakly auriculate. Operculum

multispiral with erect, calcareous lamella that is reflected

to form almost flat plate. Radula as in subfamily.

Distribution: Endemic to Loma del Puerto (Figures 18,

153fig10).

Remarks: Meganiphe is apparently a narrowly endemic,

monotypic genus. It is most closely related to Rolleia, which

occurs in the same high elevation habitats. This appears to be

an oftshoot of Rolleia with exaggerated sculpture.

Etymology: L. mega, giant + G. nipha, snowtlake.

Meganiphe rhecta Thompson, 1978

(Figures 19, 146-153)

CHRESONYMY

Meganiphe rhecta Thompson, 1978: 43-48, figs. 1-4;

Abbott, 1989: 53; Watters, 2006: 92, 441-442: Aiken,

2018: 4-9.

Type Material: Holotype: UF 22745; Paratypes: UF

22746(1), 22747(1); Paratype: USNM 711132(6); Para-

type: GTW 8227a(1).

Type Locality: “Dominican Republic, Puerto Plata

Province, Loma del Puerto, Yaroa, 700 m elevation.” This

site appears to be on the border between Santiago and

Puerto Plata provinces.

Distribution and Habitat: This beautiful species is

apparently endemic to Loma del Puerto. This is a

portion of the La Piedra Member of the Villa Trina

Formation. It is characterized by massive, cliff-forming

reefal limestone of Upper Miocene—Lower Pliocene age

(Zoeten et al., 1991). The habitat is a 700-800 m ele-

vation, nearly inaccessible ridge in a cloud forest, on

limestone blocks with lichens, moss, and ferns (Aiken,

2018, and figures).

Remarks: Shells may be banded or not. This species co-

occurs with the lizard Anolis distichus ignigularis Mertens,

1939, which may be predaceous on them (Aiken, 2018).

Etymology: Gr. rhecta, brittle. Meganiphe rhecta —

Brittle Giant Snowflake.

Genus Microabbottella new genus

Type species: Abbottella (Abbottella) diadema Watters,

2013.

G.T. Watters et al., 2020

Description: Shell minute (4-6 mm in diameter), tur-

binate, whorls adnate except just before lip. Umbilicus

wide. Protoconch of ca. 1.5 minutely pustulose whorls but

demarcation between protoconch and teleoconch not

well- defined. Teleoconch of 2.5—2.75 whorls. Axial

sculpture of numerous, crowded, low lamellae. Spiral

sculpture present only as keels. The axial sculpture on

keels forms very dense, finely serrate edge. Aperture

double, circular, solute from final whorl. Inner lip smooth,

narrowly exserted. Outer lip slightly expanded, consisting

of numerous fused lamellae, with two low auricles 180°

apart. Operculum multispiral with oblique, erect, cal-

careous lamella. Radula and anatomy unknown.

Distribution: Endemic to Loma Travesada (Figure 20).

Remarks: This genus has the smallest shells of any

abbottellines and has the fewest number of teleoconch

whorls (<3); the shells are also among the smallest known

annulariids. The curious combination of tightly packed

axial sculpture and prominent spiral cords is unlike any

other annulariid. Currently this is a monotypic genus.

Although preserved material was not available for phy-

logenetic work, the unique conchological features indicate

a genus apart from other abbottellines. Its apparently very

limited range on a single hill in the eastern—most Samana

Peninsula suggests it may have evolved in isolation there

when the peninsula was still an island.

Etymology: L. micro, very small + Abbottella.

Microabbottella diadema (Watters, 2013)

(Figures 20, 154-157)

CHRESONYMY

Abbottella (Abbottella) diadema Watters, 2013: 4, 6, figs. 1

F-J,5 P, 7B.

Type Material: Holotype, UF 456814; Paratypes: UF

456815(5), from the type locality.

Type Locality: “Dominican Republic, Samana Province,

Cabo Cabron, at 280 m. ca. 19.34° N, -69.25° W.”

Distribution and Habitat: Known only from the type

locality at the isolated Loma Travesada of the Sierra de

Samana under limestone boulders on a slope in virgin rain

forest; abundant. It may occur in <10 km”.

Etymology: Gr. diadema, headband, crown.

Genus Preclaripoma new genus

Type Species: Leiabbottella thompsoni Watters, 2013.

Description: Shells large for subfamily (9-15 mm in

diameter), nearly planispiral, sculptureless except for

Page 25

microscopic axial threads and faint spiral ridges in um-

bilicus. Outer lip greatly expanded. Operculum as in

subfamily.

Distribution: The few records indicate isolated localities

in the Cordillera Septentrionale and the Los Haitises of

the Majugual area (Figure 21).

Remarks: Originally considered members of Leiabbot-

tella, results here indicate these taxa are unrelated to that

genus and constitute a new taxon. The lack of sculpture

seen in Leiabbottella and Preclaripoma is convergent.

Preclaripoma is weakly supported as a sister group of

Abbottella. These are very rare species in collections.

Etymology: L. preclarus, very beautiful, splendid +

-poma.

Key to Species

1) Whorls ca. 3X as high as wide, outer lip barely adnate

to preceding whorls Sas. ee ah See soluta

1) Whorls ca. 4X as high as wide, outer lip broadly adnate

£0: ECE CIS AVTIOM pines ogee aah aaa thompsoni

Preclaripoma soluta (Pfeiffer, 1852)

(Figures 21, 158-161)

CHRESONYMY

Choanopoma solutum “Richard” Pfeiffer, 1851: 155 [no-

men nudum]; Adams and Adams, 1856: 296; Hjalmarson

and Pfeiffer, 1858: 138-139; Bland, 1861: 355; Pfeiffer,

1865: 110; Pfeiffer, 1876: 160; Crosse, 1891: 169.

Choanopoma? solutum “Richard:” Pfeiffer, 1852a: 167;

Pfeiffer, 1853: 117; Pfeiffer, 1858b: 102.

Cyclostoma solutum (“Richard”): Pfeiffer, 1854b: 295, pl.

39, figs. 8-10; Pfeiffer, 1858a: 188.

Cyclostoma (Choanopoma) solutum “Richard” Pfeiffer,

1852: Shuttleworth, 1856: 261-263, 270.

Cyclostoma solutum “Richard” Pfeiffer, 1852: Reeve,

1862: pl. 23, figs. 160a,b.

Choanopoma solutum “Richard” Pfeiffer, 1852: Kobelt,

1880: 277.

Abbottella solutum (Pfeiffer, 1852): Henderson and

Bartsch, 1920: 75.

Choanopoma (Abbotella |sic]) solutum “Richard” Pfeiffer,

1852: Clench and Aguayo, 1937: 67.

Incertipoma solutum (Pfeiffer, 1852): Bartsch, 1946: 171,

173-174, pl. 30, figs. 2-4.

Rolleia ? soluta (Pfeiffer, 1852): Watters, 2006: 93,

487-488.

Leiabbottella soluta (Pfeiffer, 1852): Watters, 2010: 18, 19:

Watters, 2013: 9, 11, 17, figs. 2 F-I, 7 D; Aiken, 2018: 7.

Type Material: > NHMUK, not located, but in Cuming

collection, fide Shuttleworth, 1856: 262.

THE NAUTILUS, Vol. 134, No. 1

Figures 158-176. Preclaripoma, Rolleia. 158-161. Preclaripoma soluta (Pfeiffer, 1852). 158-160. GTW 16924a (13.5 mm di-

ameter). 161. Distribution map. 162-169. Preclaripoma thompsoni (Watters, 2013). 162-164. Holotype, UF 456799 (13.7 mm

diameter). 165-166. Aiken coll. (13.4 mm diameter, photo ©S. Aiken). 167. Aiken coll. (12.9 mm diameter, photo ©S. Aiken). 168.

Live individual (photo ©S. Aiken). 169. Distribution map. 170-172. Rolleia bombardopolensis (Bartsch, 1946). 170-171. Holotype,

USNM 504085 (7.1 mm diameter). 172. Distribution map. 173-176. Rolleia haitensis Bartsch, 1946. 173-175. Holotype, USNM

504088 (12.0 mm diameter, photo courtesy of USNM). 176. Distribution map.

G.T. Watters et al., 2020 Page 27

Figures 177-199. Rolleia. 177-179. Rolleia martensi (Maltzan, 1888). 177-178. Possible syntype: ZMB 40725 (15 mm diameter,

photo courtesy Malacological Collection, Museum fiir Naturkunde Berlin, Leibniz Institute for Research in Evolution and Biodiversity

at the Humboldt University, photography L. Maitas). 179. Distribution map. 180-187. Rolleia oberi Watters and Duffy, 2010.

180-182. Holotype UF 434775, 10.3 mm diameter. 183. GTW 14180b (7.3 mm diameter). 184. GTW 14180b (8.6 mm diameter). 185.

Live individual (photo ©S. Aiken). 186. Mating pair suspended on mucous threads (photo ©S. Aiken). 187. Distribution map. 188-191.

Rolleia paradoxa (Watters, 2013). 188-190. Holotype, UF 456812 (7.8 mm diameter). 191. Distribution map. 192-199. Rolleia

simonaikeni new species. 192-194. Holotype, OSUM 46107 (9.6 mm diameter). 195. Paratype, UF 525639 (9.0 mm diameter). 196.

Paratype, BMSM 128507 (10.6 mm diameter). 197. Hyperstrophic individual (8.2 mm length, photo ©S. Aiken). 198. Mating pair

(photo ©S. Aiken). 199. Distribution.

Page 28

Type Locality: “Island of Santo Domingo.” Originally

restricted by Watters (2012) to “the southern edge of Los

Haitises Mountains, N of Majagual, ca. 12 km NW of

Sabana Grande de Boya, Monte Plata Province, Do-

minican Republic.” However, this was found to be a

different species described as Leiabbottella thompsoni

Watters, 2013. The type locality was corrected by Watters

(2013) to “Ydsica Abajo, Puerto Plata Province, Domin-

ican Republic.”

Distribution and Habitat: Known only from the region

of Loma del Puerto near Lajas de Yaroa and Gurabito de

Yaroa in the Cordillera Septentrional, between Puerto

Plata and Santiago de Los Caballeros; ca. 500-900 m

elevation. It lives in the soil under rocks (Aiken, 2018).

Remarks: This species occurs at higher elevations than P.

thompsoni, up to at least 900 m.

Etymology: L. solutus, dissolved; possibly used here to

refer to the openly coiled whorls.

Preclaripoma thompsoni (Watters, 2013)

(Figures 21, 162-169)

CHRESONYMY

Leiabbottella thompsoni Watters, 2013: 9-11, 17, 18, figs.

DM ag

Type Material: Holotype: UF 456799; Paratypes: UF

2,36225(91), from the type locality.

Type Locality: “Dominican Republic, Monte Plata

Province, 5 km N of Majagual [Majugual], at 150 m. ca.

19,09° N, —69.83° W.”

Distribution and Habitat: Found in the karstic region of

Los Haitises between the Cordillera Central and the

Cordillera Oriental around Guaragao and Majugual on

and at the base of limestone knolls and in ravines in mesic

forests under debris and leaf litter. The Majugual region is

the western part of the Cevicos Formation, composed of

karsted, massive limestone blocks of Upper Miocene—

Lower Pliocene age (Draper and Lewis, 1991). All

specimens seen were found between 100-320 m eleva-

tion. Locally common.

Etymology: Named for the late Dr. Fred Thompson,

Curator, UF, who collected the types.

Genus Rolleia Crosse, 1891

Type Species: Cyclotus martensi Maltzan, 1888, by

original designation.

Petasipoma Bartsch, 1946, is a synonym.

THE NAUTILUS, Vol. 134, No. 1

Description: Shell medium-sized for family (8-15 mm in

diameter), depressed to turbinoid, terminal portion of final

whorl solute from previous whorl and may be deflected

anteriorly. Axial sculpture of numerous close-set, very fine

threads or low lamellae. Spiral sculpture as weak cords that

may be limited to umbilicus or may be entirely absent.

Outer lip widely expanded in most species.

Distribution: Specimens are very rare in collections

and the ranges of the species are not well-known. All

appear to be upland species in the Massif du Nord and

Montagnes Noires of Haiti and the Cordillera Sep-

tentrional of the Dominican Republic (Figure 20).

These highlands are largely unexplored and _ nearly

impenetrable. Given the amount of endemism in this

group, it is to be expected that other species un-

doubtedly await discovery.

Remarks: The Haitian species may be unrelated to those

from the Dominican Republic. Paradoxa and bom-

bardopolensis also may constitute a separate genus but

require additional material.

Etymology: Hermann Rolle (f. 1887-1911), German

shell dealer.

Key to Species

1) Operculum fits within aperture, spiral sculpture absent

Ol Preset Om ly Ai) UIMMOHICHG, Quint 0.8 a. eee ae 2

1) Operculum too large to fit in aperture, spiral sculpture

Gl WeAKICOUGS OVEL WHOTIS: oo). per ne sarees oon oe

2) Shell nearly smooth, final portion of last whorl widely

CCIE Sone ale A a gee et Re a a martensi

2) Shell with coarser axial sculpture, final portion of last

Seal LAO Ay? SOM e aga bay hayes, ah) Seay ae ee 3

3) Outerdip Marrow expanded <i. wk ed es haitensis

3) Outer lig witelycxman@e@ yc ke. eld los wins 4

4) Axial sculpture close-set and barely separated by width

ORIEL 2 (lpn ane a ee eR as Se ee oberi

4) Axial sculpture separated by gaps several times wider

‘eld VOOR TIGUo 0 Mae el aee eee e ae simonaikeni

5) Outer lip widely expanded and notched .... paradoxa

5) Outer lip narrowly expanded and complete ........

Pe ta RIND fo OR Soh nS bombardopolensis

Rolleia bompardopolensis (Bartsch, 1946)

(Figures 20, 170-172)

CHRESONYMY

Petasipoma bombardopolense Bartsch, 1946: 138-139, pl.

21, figs. 4-6.

Abbottella (Petasipoma) bombardopolensis (Bartsch,

1946): Watters, 2006: 84, 179.

Abbottella (Gundlachtudora) bompardopolensis (Bartsch,

1946): Watters, 2013: 17-18, figs. 1 DD, $81.

G.T. Watters et al., 2020

Type Material: Holotype: USNM 504085 (2 specimens

in lot, holotype listed as 6.5 mm long specimen).

Type Locality: “Crevices in rocks in a ravine a little west

of Bombardopolis, Haiti.”

Distribution and Habitat: Rolleia bombardopolensis is

known from Bombardopolis and Terre Neuve in the

western-most extent of the Cordillera Centrale in Haiti.

Its habitat is unknown.

Remarks: The structure of the operculum, which is

nearly flat and cannot be withdrawn into the aperture,

separates this from all other abbottellines except R.

paradoxa. The shell sculpture of both is consistent with

Rolleia and the species occupy the known range of Rolleia

but their assignment here is highly conjectural. No suit-

able preserved material was available. The opercular

features are here considered a species-level characteristic.

The Cuban species decolorata Pfeiffer, 1859, is main-

tained in Bartsch’s subgenus Gundlachtudora but prob-

ably belongs to the Abbottellinae as well.

Etymology: From Bombardopolis, Haiti.

Rolleia haitensis Bartsch, 1946

(Figures 20, 173-176)

CHRESONYMY

Rolleia haitensis Bartsch, 1946: 140, 141-142, pl. 23, figs.

7-9; Watters, 2006: 93, 288; Watters and Dufty, 2010b:

187, figs. To: Watters, 2013: 20. 22: figs. es ee

Type Material: Holotype: USNM 504088.

Type Locality: “Ennery, Haiti.”

Distribution and Habitat: It is known from the Mon-

tagnes Noires range of Haiti. Its habitat is unknown.

Remarks: It is possible that more than one species has

been treated under this name.

Etymology: From Haiti.

Rolleia martensi (Maltzan, 1888)

(Figures 20, 177-179)

CHRESONYMY

Cyclotus martensi Maltzan, 1888: 179.

Rolleia martensi (Maltzan, 1888): Crosse, 1891:

163-164; pl. a fig. 4. Wenz, 1939: 550, fig. 1471;

Bartsch, 1946: 140-141, pl. 23, figs. 4-6; Watters,

2006: 93, 349; Watters and Duffy, 2010b: 187, figs.

10, 11; Watters, 2013: 20.

Rolleia martensii |sic| (Maltzan, 1888): Thiele, 1929: 134.

Page 29

Type Material: Possible syntype: ZMB 40725.

Type Locality: “Sanssouci in parte meridionali insulae

Haiti.” Bartsch (1946: 141) gave “Plaisance, the type lo-

cality.” This is incorrect; the Sans-Souci Palace is located

in Milot, 27 km east of Plaisance.

Distribution and Habitat: This rarely seen species is

found in the Massif du Nord and Montagnes Noires of

Haiti. Habitat information is lacking.

Etymology: Karl Eduard von Martens (1831-1904),

German conchologist at Museum fiir Naturkunde, Berlin.

Rolleia oberi Watters and Duffy, 2010

(Figures 20, 180-187)

CHRESONYMY

Rolleia oberi Watters and Duffy, 2010b: 185, 187, figs. 1-6;

Watters, 2013: 20, 22, figs. 5 G-N, 8 G; Aiken, 2018: 7.

Type Material: Holotype: UF 434775; Paratypes: UF

434776(3); Paratypes: BMSM_ 17970(2); Paratypes:

OSUM 35489(2), all from the type locality.

Type Locality: “On road half way between Santiago de

los Caballeros and Puerto Plata, E] Puerto, La Has, Puerto

Plata Province, Dominican Republic, at 830-1000 m in

the Cordillera Septentrional.” This road is Carretera

Turistica Gregorio Luperon.

Distribution and Habitat: This species is probably

limited to the area of Loma del Puerto between Puerto

Plata and Santiago de Los Caballeros in the Cordillera

Septentrional. All specimens seen were found between

700-800 m elevation on limestone outcrops in cloud forest.

Remarks: As with a few other abbottellines (and heli-

cinids), this species is found in two different colors —

brown and yellow/white. This may be a sexual charac-

teristic with males being the yellow/white morph. This

species has been observed suspending itself from mucus

thread(s) (Figure 186).

Etymology: Jim Ober, collector of the types.

Rolleia paradoxa (Watters, 2013)

(Figures 20, 188-191)

CHRESONYMY

Abbottella (Gundlachtudora) paradoxa Watters, 2013: 8,

9, 18, figs. 1 Z-CC, 9 F.

Type Material: Holotype: UF 456812; Paratypes: UF

456813(22), from the type locality.

Page 30

THE NAUTILUS, Vol. 134, No. 1

Type Locality: “Dominican Republic, Santiago Province,

Loma Diego de Ocampo, at 1200 m. ca. 19.62° N, -'70.76° W.”

Distribution and Habitat: Known only from Loma

Diego de Ocampo of the Cordillera Septentrional, on

limestone rocks on the mountain crest in dense forest with

thick vegetative debris, mosses, and epiphytes. Loma

Diego de Ocampo is located north of Santiago de Los

Caballeros and is the highest point of the Cordillera

Septentrional at 1,200 m.

Remarks: Watters (2013) placed this species under

Gundlachtudora based on opercular features. However,

the shell sculpture is unlike that genus but more similar to

Rolleia. See Remarks under Rolleia bombardopolensis.

Etymology. L. paradoxus, strange, contrary to expectation.

Rolleia simonaikeni new species

(Figures 20, 192-199)

CHRESONYMY

Rolleia sp. Aiken, 2018: figs. top of p. 8.

Description: Shell small (largest specimen seen,

11.0 mm in diameter including outer lip; smallest seen,

8.0 mm in diameter including outer lip; holotype 9.6 mm in

diameter including outer lip), depressed turbinoid, widely

umbilicate. Whorls adnate except for just behind outer lip.

Nuclear whorls 1.5, smooth, minute, white or brown,

weakly demarcated from teleoconch whorls. Teleoconch

whorls 3. Axial sculpture of minute, low, erect lamellae

(90-100 on final whorl), evenly produced all around, more

or less evenly distributed but closer set near lip, with mi-

croscopic threads between some lamellae. Spiral sculpture

absent. Aperture deflected anteriorly. Outer lip very

broadly expanded, narrower facing umbilicus, just touching

previous whorl, somewhat concave posteriorly, composed

of numerous fused lamellae, with wide posterior auricle.

Inner lip round, exserted, projecting beyond outer lip. Shell

white or pale tan with ca. 9 spiral bands of smudged tan

spots or dashes that show through into aperture. Oper-

culum not preserved with specimens but based on pho-

tographs of live animals (Aiken, 2018), composed of single,

reflected spiral lamella. Radula unknown. Preserved ma-

terial was not available for phylogenetic analysis.

Type Material: Holotype: OSUM 46107 (9.6 mm in

diameter); Paratypes: UF 525639(1), 9.0 mm in diam-

eter; BMSM 128507 (1), 10.6 mm in diameter.

Additional Material: GTW 17281a(1); Aiken coll. (29),

all from the type locality.

Type Locality: Dominican Republic, Santiago Province,

Loma del Puerto, 1.4 km NNW of Lajas de Yaroa.

Distribution and Habitat: Known only from a crevice

in a single house-sized limestone block on Loma del

Puerto at 685 m elevation in a cloud forest. This is a

portion of the La Piedra Member of the Villa Trina

Formation. It is characterized by massive, cliff-forming,

reefal limestone of Upper Miocene—Lower Pliocene age

(Zoeten et al., 1991).

Remarks: This species may occupy the narrowest range

of any annulariid, having been found only in a single

fissure on a single small outcrop (Aiken, 2018). It is also

unique in having the only known dextral hyperstrophic

example recorded for the family (Figure 197).

Etymology: Named for Simon Aiken (UK), collector of the

types and generous contributor to this and other studies.

ACKNOWLEDGMENTS

The authors thank J. Slapcinsky (UF) and R. Herschler

(retired, USNM) for access to their invaluable collections.

Z. Feher (Hungary), S. Aiken (UK), J. Joseph (UK), A.

Gettleman (USA), H. Rhode (USA), G. Duffy (USA), M.

Coltro (Brazil), A.F. Velazquez (Cuba), R. Fernandez-

Garcés (Cuba), L. and J. Kremer (USA), and R. Hooks

(Dominican Republic) generously donated valuable ma-

terial used in this review. S. Aiken graciously allowed us to

use many of his excellent photographs. B. Kemminoe

(OSU) assisted with SEM preparations. Molecular work

was carried out on the equipment and in the laboratory

space of B.C. Carstens (Department of Evolution,

Ecology and Organismal Biology, OSU). Funding for

portions of the project were generously donated by L. and

J. Kremer.

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G.T. Watters et al., 2020 Page 33

Appendix 1. Specimens used in phylogenetic study. DR = Dominican Republic.

Appendix 1

Taxon GenBank (COIJ/ 12S/ H3) Source Accession Locale

Abbottella crataegus (Watters, 2016)

Abbottella moreletiana (Crosse, 1873)

Abbottella domingoensis (Bartsch, 1946)

Abbottipoma abbotti (Bartsch, 1946)

Abbottipoma crossei (Bartsch, 1946)

Arenabbottella mellosa (Watters and Dufty, 2010)

Arenabbottella rosaliae (Pfeiffer, 1858)

Arenabbottella sosuaensis (Bartsch, 1946)

Adamsiella ignilabris (Adams, 1849)

Annularia anomala (Adams, 1850)

Annularia chittyi (Adams, 1849)

Annularia fimbriatula (Sowerby, 1825)

Annularia fumbriatula (Sowerby, 1825)

Annularia fimbriatula (Sowerby, 1825)

Annularia hilliana hilliana (Adams, 1845)

Annularia hilliana leporilabre (Adams, 1851)

Annularia lima (Adams, 1845)

Annularia lincina (Linnaeus, 1758)

Annularia lincinella (Lamarck, 1822)

Annularia mitus (Pfeiffer, 1852)

Annularia pisum (Adams, 1849)

Annularia scabricula (Sowerby, 1843)

Annularia triplopoma (Baker, 1934)

Annularisca hendersoni (Torre and Bartsch, 1941)

Annularisca victoris (Torre and Bartsch, 1941)

Chondropoma laetum (Poey, 1857)

Chondropoma oculeum (Watters and Dutty,

2010)

Chondropoma pupiforme (Sowerby, 1843)

Chondropoma sp.

Chondropomella magnifica (Pfeiffer, 1852)

Chondropomium weinlandi (Pfeiffer, 1862)

MT119707/ NA/ NA

MT119681/ MT119719/

MT119733

MT119706/ MT119717/

MT119734

MT119705/ MT119718/ NA

MT119695/ MT119723/ NA

MT119696/ MT119724/ NA

GTW 16492c

GTW 707l1¢g

GTW 1649Ic

GTW 16491b

GTW 10829b

GTW 10829c

MT119691/ MT119726/

MT119730

MT119692/ NA/ NA

MT119679/ NA/ NA

MT119713/ NA/ NA

MT119711/ NA/ NA

MT119698/ MT119721/

MT119731

MT119699/ NA/ NA

MT119687/ NA/ NA

]Q964718/ JQ990572/ N

]Q964706/ ]Q990560/ N

]Q964705/ JQ990559/ N

]Q964707/ JQ990561/ NA

1Q964708/ ]Q990562/ NA

1Q964709/ ]Q990563/ NA

1Q964710/ ]Q990564/ NA

]Q964711/ J]Q990565/ N

]Q964712/ ]Q990566/ N

MT119690/ NA/ NA

]Q964713/ ]Q990567/ N

]Q964714/ J]Q990568/ N

]Q964716/ ]Q990570/ N

ae ae ge

]Q964717/ ]Q990571/ N

1Q964763/ ]Q990616/ N

MT119683/ NA/ NA

KX496697/ NA/ NA

KX496694/ NA/ NA

MT119684/ NA/ NA

KX496717/ NA/ NA

KX496702/ NA/ NA

KX496704/ NA/ NA

KX863609/ NA/ NA

KX863610/ KX863590/ NA

KX863598/ KX863577/ NA

KX863599/ KX863578/ NA

KX863600/ KX863579/ NA

KX863601/ NA/ NA

KX863602/ KX863581/ NA

KX863611/ KX863591/ NA

MT119712/ MT119722/ NA

GTW 9432b

GTW 9432c

GTW 7020f

GTW 16507b

GTW 16507c

GTW 16507a

GTW 13728c

GTW 13728f

GTW 7244¢

ANSP:]BS263-01

USDA:97]25-01

ANSP-]JBS148-01

USDA:97J1-01

ANSP-JBS378-01

ANSP-]BS253A-01

ANSP:JBS133-01

ANSP:JBS1-01

ANSP:]BS398-01

GTW 9209d

ANSP:]BS99-01

ANSP:]BS188-02

ANSP:]BS595-01

ANSP: ]JBS69B-01

ANSP:CO35B-01

GTW 7163e

GTW 15040a

GTW 7166d

UF 48724

GTW 15050a

GTW 15046a

OSUM 42368

OSUM 42358

OSUM 42362

OSUM 42363

OSUM 42378

OSUM 42361

OSUM 42377

OSUM 42359

Majugual, DR

Boca del Infierno, DR

Rio Naranjo, DR

Puerto Bonito, DR

El Valle, DR

Anadel, DR

Talanquera, DR

Rio San Juan, DR

Cabarete, DR

Monkey Jungle, DR

Sostia, DR

Cabarete, DR

Sostia, DR

Windsor Cave, Jamaica

Manchester Parish,

Jamaica

St. Elizabeth Parish,

Jamaica

St. Elizabeth Parish,

Jamaica

Hanover Parish, Jamaica

St. Ann Parish, Jamacica

Clarendon Parish, Jamaica

Manchester Parish,

Jamaica

St. Elizabeth Parish,

Jamaica

St. Catyherine Parish,

Jamaica

Rio Cobre, Jamaica

St. Catyherine Parish,

Jamaica

Portland Parish, Jamaica

St. Elizabeth Parish,

Jamaica

Trelawny Parish, Jamaica

Holguin, Cuba

Playa El] Baga, Cuba

Playa El Baga, Cuba

Velasco, Cuba

Pedernales, DR

Anguilla

Rafael Freyre, Cuba

Guardalavaca, Cuba

Puerto Escondido, DR

Bartolomé, DR

Bombita, DR

Bartolomé, DR

Galindo Adrento, DR

El Limén, DR

Bartolomé, DR

(Continued)

Page 34

Appendix 1. (Continued)

Appendix 1

Taxon

Chondropomium weinlandi (Pfeiffer, 1862)

Clydonopoma bartschi (Watters, 2012)

Clydonopoma nobile (Pfeiffer, 1852)

Clydonopoma poloense (Bartsch, 1946)

Clydonopoma pumilum (Watters and Duffy,

2010)

Colobostylus retrorsa ( Adams, 1850)

Crossepoma marmoreum (Watters and Dufty,

2010)