Zoosyst. Evol. 100 (1) 2024, 9-14 | DOI 10.3897/zse.100.110105

ae
BERLIN

Pilsbrylia, a dextral-shelled door snail from South America
(Gastropoda, Clausiliidae)

Rodrigo B. Salvador’, Abraham S. H. Breure*>:4

The Arctic University Museum of Norway, UiT — The Arctic University of Norway, Lars Thorings veg 10, 9006, Tromso, Norway

Royal Belgian Institute of Natural Sciences, Rue Vautier 29, 1000, Brussels, Belgium

Invertebrate Division, Department of Life Sciences, Natural History Museum, Cromwell Road, South Kensington, SW7 5BD, London, UK
Naturalis Biodiversity Center, Darwinweg 2, 2333 CR, Leiden, Netherlands

FW NM fF

https://zoobank. org/98538B32-8A69-40E9-85CE-1421B8CO60E4E

Corresponding author: Rodrigo B. Salvador (salvador.rodrigo.b@gmail.com)

Academic editor: Frank Kohler # Received 26 July 2023 Accepted 3 September 2023 @ Published 26 January 2024

Abstract

The land snail genus Pilsbrylia Hylton Scott, 1952 has been recently shown to not belong to the superfamily to which it was orig-
inally assigned (1.e., the Orthalicoidea), instead pointing out a relationship with the Clausilioidea. In this study, we included the
type species of the genus in a multi-marker molecular phylogenetic framework to reassess its family-level classification. Our results
show that Pilsbrylia belongs to family Clausiliidae (known as ‘door snails’) and more specifically, to subfamily Peruiniinae. This
family is unique among stylommatophorans for consisting almost exclusively of animals with sinistral (left-handed) shells, whilst

Pilsbrylia has a “typical” dextral shell.

Key Words

chirality, Eupulmonata, Orthalicoidea, Peruiniinae, Stylommatophora

Introduction

The South American genus Pilsbrylia Hylton Scott,
1952 contains three species, which inhabit areas in
southern Brazil and northern Argentina (Simone 2018):
Pilsbrylia paradoxa Hylton Scott, 1952 (the type spe-
cies), P. hyltonae Fernandez & Rumi, 1980, and P. dalli
Simone, 2018 (Fig. 1). Members of this genus have high-
spired and narrow shells and the shell aperture bears a
number of teeth and lamellae. The shells are morpho-
logically similar to the members of family Cyclodontin-
idae (which was formerly part of Odontostomidae; see
Salvador et al. 2023 for the revised classification within
superfamily Orthalicoidea), in particular to members of
genera such as Clessinia Doering, 1875 and Cyclodontina
H. Beck, 1837. Thus, since its description, Pi/sbrylia has
been classified in that family (e.g., Hylton Scott 1952;
Breure 1974; Fernandez and Rumi 1980; Schileyko 1999;
Cuezzo et al. 2013; Simone 2018).

The molecular phylogenetic study of Breure and
Romero (2012) showed that Pilsbrylia did not belong
in Cyclodontinidae and those authors proposed the
genus was instead the sister taxon to all other Orthal-
icoidea. A more recent phylogenetic study (Salvador
et al. 2023), showed that Pilsbrylia did not belong in
Orthalicoidea at all, being instead related to the Clausi-
liidae, or door snails. That was a surprising result, con-
sidering that the door snails typically have sinistral
Shells (1.e., a shell that “coils” counter-clockwise or,
when seen with its aperture facing the observer and the
Spire top pointing upwards, a shell whose aperture is
on the left-hand side).

Furthermore, the two branches of Clausiliidae present
in the Americas are restricted to the Caribbean (subfamily
Neniinae) and northwest South America (subfamily Peru-
iniinae) (Uit de Weerd and Gittenberger 2013). Thus, the
genus Pilsbrylia is rather geographically removed from
the family’s range.

Copyright Sa/vador, R.B. & Breure, A.S.H. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits
unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

10

Figure 1. Shells of the three known species of Pilsbrylia in
apertural, lateral, and dorsal views. A. P. paradoxa, holotype
MLP-Ma 11337 (Museo de La Plata, Argentina); B. P. hyltonae,
lectotype MLP-Ma 3991-1; C. P. dalli, holotype MZSP 133161
(Museu de Zoologia da Universidade de Sao Paulo, Brazil).
Scale bar: 5 mm.

In the present study we include Pilsbrylia in a phylo-
genetic framework of the Clausiliidae to test if it really
belongs to this family and, if so, assess how it 1s related
to other South American door snails.

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Salvador, R.B. & Breure, A.S.H.: Reclassification of Pi/sbrylia in Clausiliidae

Materials and methods

DNA sequences of Pilsbrylia paradoxa used in previous
Orthalicoidea-focused studies (Breure and Romero 2012;
Salvador et al. 2023) were used for the present analysis.
They are available on GenBank under accession num-
bers JF514745 (28S) and JF514687 (H3) and stem from
a voucher specimen previously housed in the collection
of the Instituto Miguel Lillo, Tucuman, Argentina (regis-
tration number IFML-MOLL BD316, collected in north-
ern Argentina, Salta province, km 1650 of Salta-Jujuy
highway) but now housed in the Instituto de Biodiver-
sidad Neotropical (IBN) of the Universidad Nacional de
Tucuman, Argentina. No further specimen of Pi/sbrylia
spp. preserved in ethanol and suitable for DNA extraction
could be found in the present study.

The sequences of P. paradoxa were included in the
phylogenetic framework for the family Clausiliidae estab-
lished in the study of Uit de Weerd and Gittenberger (2013).
That study included a reasonable sample of South Ameri-
can (subfamily Neniinae) taxa across two distinct clades
(tribes), which enables us to test the phylogenetic position
of Pilsbrylia. Later molecular phylogenetic studies with
Clausiliidae focused on Eurasian taxa (e.g., Hausdorf and
Neiber 2022) and data from them was not included here.

A total of 67 species of Clausiliidae, belonging to all sub-
families and almost all tribes, was part of the analysis. Five
Species were selected as outgroup representing the families
Cerionidae, Chondrinidae, Enidae, Rhytididae, and Urocop-
tidae (data from Uit de Weerd 2008; Uit de Weerd and Git-
tenberger 2013; Saadi and Wade 2019). A complete list of
the species used 1n the analysis, with locality data and Gen-
Bank accession numbers is given in the Suppl. material 1.

Data from three nuclear markers were used in the present
phylogenetic analysis, following Uit de Weerd and Gitten-
berger (2013): partial 28S rRNA gene (ca. 1700 bp), partial
H3 (histone 3) gene (ca. 270 bp), and partial H4 (histone 4)
gene (ca. 260 bp). Information on primers and PCR proto-
cols can be found in Uit de Weerd and Gittenberger (2013);
for Pilsbrylia, see Breure and Romero (2012).

The genetic sequences were aligned through the MUS-
CLE plugin (Edgar 2004) in Geneious Prime (v.2023.0.4,
Biomatters Ltd.), using default settings (1.e., optimised
for accuracy). The resulting alignments were visually
proofed for inconsistencies. The alignment of the 28S
marker was run through Gblocks (Talavera and Castresa-
na 2007), using the least restrictive settings, in order to
eliminate poorly-aligned or data-deficient positions that
could introduce noise into the analysis. The alignments
were then concatenated for a single phylogenetic analysis,
with each marker being treated as an individual partition.

A Bayesian inference phylogenetic analysis was per-
formed through MrBayes (v.3.2.7; Ronquist et al. 2012)
via the CIPRES Science Gateway (v.3.3; Miller et al.
2015). Two concurrent analyses, each with 4 Markov
chains of 80 million generations (the first 20% discarded
as “burn-in’), were run with the default priors, nst = 6,

Zoosyst. Evol. 100 (1) 2024, 9-14

rates = invgamma, temperature parameter = 0.1, sampling
every 1,000 generations. Substitution model parameters
were unlinked across the markers (28S, H3, and H4).
MCMC convergence was assessed using the standard de-
viation of split frequencies (<0.01) and the potential scale
reduction factor PSRF (~1.0), as well as by examining the
trace plots (Ronquist et al. 2009).

Results

The concatenated sequences of the three markers (af-
ter trimming the 28S marker using Gblocks) contained
2172 bp. The total-evidence tree resulting from the
Bayesian analysis contained 73 species (including

0.75

11 (0.68 <=!

Macedonica pangaionica
—S>
0.76

Pontophaedusa funiculum

1
0.54

11

Pilsbrylia paradoxa and the outgroup) and is shown here
in simplified format, with the branches of non-immediate
interest collapsed (Fig. 2). The full tree can be seen in the
Suppl. material 2.

All subfamilies of Clausiliidae are strongly support-
ed (posterior probability PP=1), except for Serrulininae,
which 1s paraphyletic (Fig. 2). As expected in a clade in-
flated by family-level names, many of the supposed tribes
were recovered as para- or polyphyletic (e.g., Cochlodini-
ni, Delimini; Suppl. material 2), as already noted by Uit
de Weerd and Gittenberger (2013). When also consider-
ing the existence of several monotypic tribes (e.g., Garn-
ieriini, Strumosini), the Clausiliidae would benefit from a
clean-up of names on the tribe level, as many could easily
be synonymized if supported by further research.

ef Baleini + Clausiliini

Oe <] Acrotomini + Strigileuxinini

| Filosini + Mentissoideini
0.68

Olympicola olympica Olympicolini

. — <A Boettgeriini

= Strumosa strumosa Strumosini

. — ames ie | Medorini

Montenegrina dennisi

0.96

AVNIMNSNV19

' Herilla bosniensis rex Alopiini

ALOPIINAE
Alopia bielzii

| Cochlodinini
) + Delimini

PHAEDUSINAE

SERRULININAE

ee | GARNIERIINAE

0.88

0.97
0.55

Temesa clausilioides
Cyclonenia gibber
Incaglaia adusta

Incania papillosa PERUINIINAE

Steeriana malleolata

Andinia taczanowskii

Cylindronenia cicatricosa

1 Peruinia peruana

0.99 1

Neniatlanta pauli

1 Bofilliella subarcuata

Zilchiella scala

Nenisca franzi
NENIINAE

Nenia tridens

| LAMINIFERINAE

Mastus cretensis

Rhytida stephenensis 0.02

Chondrina avenacea
Cerion striatellum

—_—_m_=__==mms A tocoptis menkeana

Figure 2. Bayesian inference tree of the Clausilioidea. The crown group 1s collapsed to facilitate visualization (see Suppl. material 2
for a full view). Posterior probabilities are shown on nodes. Scale bar is substitutions per site.

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12

Considering that our ingroup was identical to that of
Uit de Weerd and Gittenberger (2013) — barred the inclu-
sion of Pilsbrylia paradoxa — our tree is largely similar to
the one presented by those authors. However, a major dif-
ference can be observed: while Uit de Weerd and Gitten-
berger (2013) had the Neniinae as the sister group to all
other clausiliids (though unsupported), our tree places the
Laminiferinae in that position (Fig. 2). The Laminiferinae
are sister to the remaining clausiliids (PP=0.97) and, in-
side the latter clade, the Neniinae are sister to all others,
which are joined in an unsupported polytomy (PP=0.54).
This polytomy includes three strongly supported branch-
es (PP=1: Peruiniinae, Garnieriinae, and a clade formed
by Phaedusinae and a paraphyletic Serrulininae) and an
unsupported branch (PP=0.75) containing the Alopiinae
and the Clausiliinae (each PP=1) as sister clades (Fig. 2).

Our taxon of interest, Pilsbrylia paradoxa, 1s included
in the Peruiniinae, in a derived position (Fig. 2). It was
recovered as the sister taxon to the Peruvian /ncaglaia
adusta (O. Boettger, 1880), although with low support
(PP=0.88). Both species are the sister clade to Incania
papillosa Neubert & Nordsieck, 2005 (PP=1), also from
Peru. The relationships within Peruiniinae recovered by
our analysis are largely the same as those in Uit de Weerd
and Gittenberger (2013), with a single exception: those
authors recovered Zilchiella scala Neubert & Nordsieck,
2005 and Peruinia peruana (Troschel, 1847) as sister taxa
(although unsupported), while in the present tree they
form a trichotomy with the remaining Peruintinae (Fig. 2).

Discussion

The results of the present phylogenetic analysis allow the con-
clusion that Pilsbrylia in fact belongs in the family Clausili-
idae, more specifically to the South American subfamily Pe-
ruiniinae, as expected from a biogeographical perspective.
Pilsbrylia, however, has a much different shell morphol-
ogy from other members of the Peruiniinae. First of all, the
Shell is dextral. Dextral shells are rare in Clausiliidae, but
are more frequent in subfamily A/opiinae (Nordsieck 2007;
Pall-Gergely et al. 2019). Within the genera A/opia Adams
& Adams, 1855 and Albinaria Vest, 1867, eniantomorph
taxon pairs are known, and dextral lineages have evolved
from sinistral ancestors multiple times independent-
ly (Fehér et al. 2013; Kornilios et al. 2015). While those
shells are usually near mirror images of their congeners or
conspecifics, Pilsbrylia, on the other hand, presents a shell
morphology that is more similar to Cyclodontinidae than to
most Clausiliidae, notably by not having a complete peri-
stome (Fig. 1), which justified its previous classification.
Hylton Scott (1952) described the radular and genital
anatomy of P. paradoxa, the only species in the genus for
which anatomical data is available. Notably, that author
did not mention a clausilium, which would be a reasonably
obvious structure if present (unless it was present as a re-
duced structure). The single columellar lamella of Pi/sbry-
lia spp. only extends about half whorl inside the shell along
the columella (Hylton Scott 1952; Simone 2018), which

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Salvador, R.B. & Breure, A.S.H.: Reclassification of Pi/sbrylia in Clausillidae

could indicate a reduced or absent clausilium. Clausiliidae
have two columellar lamellae that stretch throughout the
entire body whorl (Nordsieck 2007, 2015).

Species of the sister genus to Pi/sbrylia in the pres-
ent phylogeny, /ncaglaia Pilsbry, 1949 (Fig. 2), have a
typical clausiliid shell morphology. However, members
of the next closest genus, /ncania Polinski, 1922, can
exhibit a simplified peristome, reduced lamellae and
reduced clausilial apparatus (Neubert and Nordsieck
2005). Notably, some fossil species of European Ryllia
Munier-Chalmas, 1883 and Rillyarex Nordsieck, 1985
have reduced or absent lamellae and a peristome with
similar configuration to Pilsbrylia spp. (Nordsieck 2015).

Thus, the epithet ‘paradoxa’ given to the first species
described in this genus reaffirms itself as a very appropriate
name: genetically, it is a Clausiliidae, but it does not pos-
sess the two typical traits of the family, 1.e., a sinistral shell
and a clausilial apparatus (or might present it in a reduced
state). Nevertheless, the genetic similarity of Pilsbrylia
with clausiliids is observed across all markers analysed
and thus, a fortuitous similarity of a single marker biasing
the phylogenetic analysis is unlikely. In view of the present
evidence, we conclude that Pilsbrylia is a member of Peru-
iniinae defined by its dextral shell and hypothesize that the
simplification of the dextral shell (i.e., aperture and lamel-
lae) is related to the reduction or loss of the clausilium.

The Clausilioidea are considered to have arisen in Eu-
rope, as both the extinct Filholiidae and Palaeostoidae,
as well as one of the first branches of Clausiliidae (..e.,
Laminiferinae; Fig. 2) are from that continent (Nordsieck
2007, 2015; Uit de Weerd and Gittenberger 2013). The
Palaeostoidae date back to the Late Cretaceous of western
Europe and so does Clausiliidae thanks to the extinct tribe
Rillyini Nordsieck, 1985 (Nordsieck 2015).

Still, Clausiliidae has a clear Laurasian component,
with Neniinae in the Caribbean and Peruiniinae in South
America (Fig. 2) and an origin in the latter cannot be en-
tirely excluded (Uit de Weerd and Gittenberger 2013). It
has since become known that the presence of the family
Clausiliidae (and purportedly of subfamily Peruiniinae) in
South America is equally old. Namely, there is a fragmen-
tary fossil from the Late Cretaceous of Uruguay classified
as Clausiliidae indet. (Salvador et al. 2018; Cabrera et al.
2020). The next oldest fossil is Temesa? magalhaesi (Trin-
dade, 1953) from the Early Eocene of Rio de Janeiro, Bra-
zil (Salvador and Simone 2013; Salvador et al. 2018). The
latter species was provisionally assigned to a recent genus,
but it likely belongs to a different (and still undescribed)
genus, potentially close to the basal node of Peruiniinae.

Conclusion

Considering the present findings, we propose a revised
classification of the genus Pi/sbrylia Hylton Scott, 1952,
placing it inside subfamily Peruiniinae of the Clausili-
dae. Thus, this extends the known range of this subfamily
in South America to the south (Argentina and Uruguay)
and to the east (Brazil, Minas Gerais state).

Zoosyst. Evol. 100 (1) 2024, 9-14

Acknowledgements

We are very grateful to Diego E. Gutierrez Gregoric
(MLP), Eugenia S. Orofio (IFML), Fernanda S. Silva
(MZSP), M. Gabriela Cuezzo (Universidad Nacional de
Tucuman) for the photos of the specimens of Pilsbrylia,
including the types; and to the reviewers and editor Frank
Kohler for their comments. Our study used genetic se-
quences from previous papers that are freely available in
GenBank (Suppl. material 1) and thus, we are very grate-
ful to the authors of those works.

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Supplementary material |

Table listing all species used in the present
analysis, including information on their
locality of origin and GenBank accession
numbers

Authors: Rodrigo B. Salvador, Abraham S. H. Breure
Data type: xlsx

Salvador, R.B. & Breure, A.S.H.: Reclassification of Pi/sbrylia in Clausillidae

Supplementary material 2

Bayesian inference tree of the Clausilioidea
showing the complete set of terminal taxa

Authors: Rodrigo B. Salvador, Abraham S. H. Breure

Data type: png

Explanation note: Posterior probabilities are shown on
nodes. Scale bar is substitutions per site.

Copyright notice: This dataset is made available under

Copyright notice: This dataset is made available under

the Open Database License (http://opendatacommons.
org/licenses/odbl/1.0/). The Open Database License
(ODbL) is a license agreement intended to allow us-
ers to freely share, modify, and use this Dataset while
maintaining this same freedom for others, provided

the Open Database License (http://opendatacom-
mons.org/licenses/odbl/1.0/). The Open Database
License (ODbL) is a license agreement intended to
allow users to freely share, modify, and use this Data-
set while maintaining this same freedom for others,
provided that the original source and author(s) are

that the original source and author(s) are credited.
Link: https://do1.org/10.3897/zse.100.110105.suppl1

zse.pensoft.net

credited.
Link: https://doi.org/10.3897/zse.100.110105.suppl2