![Compilation of shells representing most of the currently recognized genera of Ariantinae. 1. Arianta arbustorum (Linnaeus, 1758) B 24.1 mm [RMNH G2131] Austria, Steiermark, near Gstatterboden; E. Gittenberger leg., 10-IX-1964. 2. Ari-](https://www.researchgate.net/publication/334445391/figure/fig2/AS:780187486662658@1563022600717/Compilation-of-shells-representing-most-of-the-currently-recognized-genera-of-Ariantinae_Q320.jpg)
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Contributions to Zoology, 85 (1) 37-65 (2016)
Systematics of Ariantinae (Gastropoda, Pulmonata, Helicidae), a new approach to an old
problem
Dick S.J. Groenenberg1, 4, Peter Subai2, Edmund Gittenberger1, 3
1 Naturalis Biodiversity Center, P.O. Box 9505, Sylviusweg 70, 2333 BE Leiden, The Netherlands
2 Kronenberg 143, D-52074, Aachen, Germany
3 Institute of Biology, Leiden University, P.O. Box 9505, 2300 RA Leiden, The Netherlands
4 E-mail: Dick.Groenenberg@naturalis.nl
Key words: Ariantinae, classication, molecular phylogeny, morphology
Abstract
A new starting-point in Ariantinae systematics is presented by
combining data on traditional shell morphology and genital
anatomy, with phylogeny reconstructions based on DNA se-
quence data. For nearly all genera and subgenera one or more
shells are depicted and drawings of the proximal part of the
genital organs are shown to illustrate the morphological diver-
sication within the subfamily. For as much as our material al-
lowed it, partial sequences are presented for Histone H3 (H3),
Cytochrome c oxidase subunit I (COI) , Cytochrome B (CytB)
and 16S ribosomal RNA (16S). Some of the allegedly speciose
genera like Chilostoma and Campylaea (Zilch, 1960) do not
represent monophyletic groups of species, whereas most of the
remaining nominal taxa (e.g. Causa, Dinarica, Josephinella,
Faustina, Liburnica, Kosicia and Thiessea) warrant a separate
taxonomic status indeed. Sequence data from individual mark-
ers were informative at the species-level, but not for higher-
level phylogenetics. Insight in genus-level relationships was
obtained after concatenation of the individual datasets. The
Ariantinae are estimated to have or iginated during the late Cre-
taceous (Campanian), not later than ca. 80 million years ago.
The enigmatic and morphologically aberrant, monotypic genus
Cylindrus is shown as the sister-group of Arianta, a genus in-
cluding A. arbustorum, which is also unusual in shell-shape
and
habitat. Ariantopsis and Wladislawia are classied as sub-
genera of neither Campylaea nor Chilostoma, but Cattania.
Sabl jaria is considered a subgenus of Dinarica. The nominal
genus Superba is shown to be paraphyletic; additional data
should demonstrate whether Superba has to be synonymised
with Liburnica. The Ariantinae are here divided in 21 genera
(2
new) and 13 subgenera (3 new).
Contents
Introduction ..................................................................................... 37
Material and methods .................................................................... 39
Tax on sam pling ......................................................................... 39
Genital anatomy ....................................................................... 39
DNA extraction, PCR and sequencing ................................ 39
Phylogenetic analyses .............................................................. 41
Fossil occurrences and age of taxa ...................................... 41
Genetic distances ..................................................................... 43
Systematics ................................................................................ 43
Results ............................................................................................... 43
Discussion ........................................................................................ 44
Acknowledgements ........................................................................ 44
References ........................................................................................ 44
Appendix .......................................................................................... 49
Introduction
The classication of the Ariantinae Mörch, 1864 (Gas-
tropoda, Helicidae), a subfamily of terrestrial air-
breathing snails, with a primary radiation in southern
Europe, has been under debate for more than a century.
Apart from a few exceptions such as Cylindrus obtu-
sus, Helicigona lapicida and Isognomostoma isogno-
mostomos, most species within this subfamily are con-
chologically close to a basic bauplan (see Appendix),
with shells that are more or less depressed globular,
with an open umbilicus and no apertural teeth. All
species are characterized by a pair of accessory glands
in the genital system, inserting between the dart sac (=
bursa telae) and the bursa copulatrix. In the literature,
these glands are often referred to as mucous glands, a
term that should be preferentially used for glands in
the snail’s foot-sole, however. The accessory glands
can either be undivided or more or less completely
split and are always longer than the dart sac.
The conchological uniformity did not hamper the
description of new species, and higher taxa, what made
the delimitation of genera and subgenera increasingly
subjective. In the literature, more than once, taxon sta-
tus was changed from a generic to a subgeneric level,
or the other way round, without proper argumentation.
This led to the confusing situation of today, where
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Groenenberg et al. – Systematics of Ariantinae
there is neither consensus on the identication and
ranking of the taxa that should be accepted within the
Ariantinae, nor about their mutual phylogenetic rela-
tionships.
Thus, the main issue regarding the systematics of
this subfamily is not the delimitation of species, but
the distinction between genera and subgenera. This is
illustrated by the number of species-group taxa that
have been described after Mörch (1864) erected the
subfamily, and even more so by the number of genera
used by different authors to classify the same species.
Ariantopsis pelia, for example, has also been classied
in Arianta, Chilostoma, Faustina, and Helicigona. In
the taxonomic literature a variety of subdivisions of
the Ariantinae has been proposed (e.g. Sturany and
Wagner, 1914; Hesse, 1931; Knipper, 1939; Zilch, 1960;
Subai, 1984, 1996; Bank et al., 2001; Subai, 2002;
Sub
ai and Fehér, 2006; Schileyko, 2006, 2013), among
which the enumeration by Zilch (Table 1) has been
most frequently cited. The classication resulting from
this study will be compared in some detail with only
the latter. This article is an extended version of a pub-
lication by Groenenberg et al. (2012), which was pub-
lished only as a part of a doctoral thesis. In a recent
article Cadahia et al. (2013) published similar data on
the phylogeny of the Ariantinae, dealing with fewer
taxa, however, and without discussing the implications
for classication and nomenclature. Schileyko (2006,
2013) suggested classications of the Ariantinae on
the basis of morphological data. Initially (Schileyko,
2006) Marmorana Hartmann, 1844, with some gener-
ally accepted close relatives, and Theba Risso, 1826,
were considered to belong to the Ariantinae. Later on,
however (Schileyko, 2013), these genera were classi-
ed in other subfamilies, viz. Murellinae Hesse, 1918
and Thebinae Wenz, 1923. Interestingly, on the basis
of a preliminary DNA analysis, using COI sequences
in GenBank, Marmorana, Murella, and Tyrrheniberus
showed up as Ariantinae indeed, whereas Theba has to
be excluded as a genus of that subfamily. Thus, the sta-
tus of the so-called Murellinae has to be studied in
more detail.
Table 1. Selection of former classications of the Ariantinae by different authors.
Sturany and Wagner (1914) Zilch (1960) Bank et al. (2001) ‘Clecom’
Genus Subgenus Genus Subgenus Genus Subgenus
Campylaea Cattania Arianta Arianta
Campylaea Campylaea Ariantopsis Causa
Dinarica Campylaea Chilostoma Ariantopsis
Liburnica Delphinatia Campylaea
Cylindrus Dinarica Campylaeopsis
Helicigona Arianta Faustina (=Cattania) Cattania
Campylaeopsis Liburnica Chilostoma
Cingulifera Wladislawia Cingulifera
Drobacia Chilostoma Campylaeopsis Corneola
Helicigona Chilostoma Delphinatia
Thiessea Cingulifera Dinarica
Isognomostoma Drobacia Josephinella
Vidovicia Josephinella Kosicia
Kosicia Liburnica
Thiessea Thiessea
Cylindrus Wladislawia
Helicigona Cylindrus
Isognomostoma Drobacia
Vidovicia Faustina
Helicigona
Isognomostoma
Vidovicia
5 genera 10 subgenera 7 genera 14 subgenera 9 genera 14 subgenera
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Contributions to Zoology, 85 (1) – 2016
Material and methods
Taxon sampling
This study is based on 172 specimens (including 5 out-
group specimens) from 85 (sub)species of Ariantinae
from across Europe, representing about half of the
known species and all the currently accepted genera
(see Appendix). Specimens were collected in the
peri
od 1957-2012 and most material was xed and
conserved in 70% ethanol or isopropanol. Some (old)
specimens were stored in “spiritus” (methylated spir-
its), whereas the more recently collected specimens
were preserved in 97% ethanol.
Genital anatomy
For nearly all the (sub)genera the genital tract is illus-
trated (see Appendix). These gures are arranged ac-
cording to the type of accessory glands, i.e. undivided
versus one or both glands split. Only Josephinella
vikosensis, with undivided accessory glands, was il-
lustrated next to two congeneric species with split
glands. For all the (sub)genera our personal observa-
tion regarding the accessory glands is presented. We
refrained from an analysis of all the data that can be
found in the literature (often without information on
the actual number of individuals that was investigat-
ed). For the ease of comparison we only differentiate
between undivided vs. split accessory glands, i.e.
specimens in which only one of the glands was divid-
ed, as well as those with trifurcate glands, were con-
sidered split.
DNA extraction, PCR and sequencing
Total genomic DNA was extracted from small foot tis-
sue samples using a DNeasy Tissue Kit (Qiagen) fol-
lowing the manufacturer’s instructions. As a follow-up
of an earlier investigation (Gittenberger et al., 2004),
this study started with the amplifcation of COI, but
due to the poor quality of some of the DNA extracts,
mini-barcode primers (Hajibabaei et al., 2006; Meus-
nier et al., 2008) were occasionally used to amplify a
smaller fragment of COI (124 bp fragment excluding
primersites). These mini-barcode sequences grouped
with those of conspecics, or otherwise with conge-
ners, for which the 655 bp COI fragment was obtained.
Although most of the recognized (sub)genera formed
well supported clades (based on Bayesian phylogeny
inference), the relationships between the (sub)genera
were poorly supported. Therefore nuclear marker H3
and mitochondrial markers CytB and 16S were added.
PCR primers and references are given in Table 2.
PCRs were carried out in 25 µl volumes using 1.25
units of Taq DNA polymerase (Qiagen), 0.4 mM of
each primer and 0.2 mM dNTPs. For COI the nal
MgCl2 concentration occasionally had to be increased
to 2.5 mM (1x PCR buffer contains 1.5 mM; Qiagen).
For 16S, Q-solution (Qiagen; nal concentration 1 ×)
was added to most of the reactions. PCR thermopro-
le: inititial denaturation 3 min. @ 94˚C, followed by
Table 2. Primer information. * Amplicon length excluding primer sequence. ** Annealing temperature. *** Minibarcode Reverse primer
has been modied to be more specic for Ariantinae.
Primer name Sequence (5’ to 3’) Marker Length* Source AT**
H3-F ATGGCTCGTACCAAGCAGACVGC H3 328 Colgan, 2000 57
H3-R ATATCCTTRGGCATRATRGTGAC
L1490 GGTCAACAAATCATAAAGATATTGG COI 655 Folmer, 1994 45-50
H2198 TAAACTTCAGGGTGACCAAAAAATCA
MB-F TCCACTAATCACAARGATATTGGTAC COI mini-barcode 124 Meusnier, 2008 50
MB-R*** GAAAATTATKACAAARGCATGAGC
151-F TGTGGRGCNACYGTWATYACTAA CytB 361 Merritt, 1998 50
270-R AANAGGAARTAYCAYTCNGGYTG
Pal-F CGGCCGCCTGTTTATCAAAAACAT 16S 404-412 Palumbi, 1991 50
Pal-R GGAGCTCCGGTTTGAACTCAGATC
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Groenenberg et al. – Systematics of Ariantinae
Table 3. Information content per dataset. Inf. char. = The number of parsimony informative characters. Perc. inf. = The percentage of
informative characters, calculated as 100 × (Inform. char. / Total char.).
Dataset Specimens Total char. Constant char. Inf. char. Perc. inf.
H3 161 328 266 46 14.0
COI 149 655 339 300 45.8
CB 91 361 128 222 61.5
16S 82 335 163 149 44.5
H3-COI-CB 89 1344 748 558 41.5
H3-COI-CB-16S 103 1679 912 711 42.3
Fig. 1. MrBayes phylogeny based on the ‘stringent’ H3-COI-CytB dataset. Branch values show posterior probabilities.
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Contributions to Zoology, 85 (1) – 2016
40 cycles of – denaturation 15 sec. @ 94˚C, annealing
30 sec. @ AT (Table 2), extension 40 sec. @ 72˚C –
and a nal extension of 5 min. @ 72˚C. PCR products
were cleaned with a Montage purication kit (Mil-
lipore) at Macrgoen Inc. Europe (Amsterdam), where
they were sequenced in both directions on an
ABI37730XL using the same primers as used for the
PCR. Forward and reverse sequences were assembled
with Sequencher 4.10.1 (Gene Codes Corporation) and
protein coding genes (H3, COI and CytB) were manu-
ally aligned in MacClade 4.08 (Maddison and Maddi-
son, 2005). The alignment for 16S was made with
MAFFT (Katoh and Standley, 2013) as implemented
in the software package Geneious Pro 7.0.6 using the
default settings with the G-INS-i algorithm. Non-
con
served blocks of sequence data were removed from
the alignment with Gblocks (Castresana, 2002) con-
ducted on the Gblocks Server (http://molevol.cmima.
csic.es/castresana/Gblocks_server.html), using only
the ‘more stringent selection’ option (which restricts
the introduction of contiguous nonconserved posi-
ti ons).
Phylogenetic analyses
For each dataset (each marker) a nucleotide substitu-
tion model was selected with MrModeltest 2.2 (Ny-
lander, 2004). For the mitochondrial datasets the mod-
el was GTR+I+G, for H3 it was HKY+I+G. Bayesian
analyses were done in MrBayes 3.2.1. (Ronquist and
Huelsenbeck, 2003) hosted on the CIPRES Science
Gateway (Miller, 2010). For each marker the analysis
consisted of two simultaneous, four chain, MCMC
runs (10 M generations). Trees were sampled every
1000 generations, the rst 2500 trees were discarded
as burnin (relburnin = yes, burninfrac = 0.25). Exami-
nation of the .p output les in Tracer v.1.5 (Rambaut
and Drummond, 2007) showed stationarity was
reached with proper effective sample sizes for all pa-
rameters (ESS > 200). Sumtrees (Sukumaran and
Holder, 2010) was used to calculate 25% majority rule
consensus trees. Subsequently the datasets for the indi-
vidual markers were combined into two concatenated
datasets (from hereon referred to as): the ‘stringent’
and ‘relaxed’ datasets. The stringent dataset (89 taxa)
consisted of only protein coding genes (i.e. H3, COI
and CytB) and had no missing data. The relaxed data-
set (103 taxa) consisted of all markers (H3, COI, CytB
and 16S); taxa for which only one marker was missing
were also included. A partitioned analysis was set up
in MrBayes (same version) for both datasets; for each
partition the GTR+I+G model was selected using the
above described procedure.
Fossil occurrences and age of taxa
In a recent check-list of fossil land snails of western
and central Europe, Nordsieck (2014) reviews fossil
taxa based on stratigraphic ranges. The oldest and only
indisputably identied Ariantinae fossil in that list is
of a Helicigona species from the late Burdigalian,
Early Miocene (17.5-16 MYA; references in Nordsieck,
2014). This fossil was used as a single calibration point
imposing a normal distribution prior (mean 16.75 MY,
stdev 0.375) allowing for soft minimum and maximum
age boundaries. Inital BEAST analyses were per-
formed with and without setting the monophyly of the
Ariantinae sensu auct. as a constraint. Species of the
genera Cepaea, Caracollina and Soosia were used as
outgroup taxa that are traditionally classied in closely
related taxa within the same superfamily Helicoidea.
If the monophyly of the Ariantinae sensu auct. was not
set as a constraint, many internodes appeared between
these outgroup taxa and the root of the tree. Re-rooted
with the outgroup taxa, the topology was virtually
identical to the ML (not shown) and MrBayes phylog-
enies. In our BEAST analyses, clades I, II and III (PP
≥ 0.86) were therefore used as a constraint. Three runs
consisting of 100 M generations were performed (for
both the stringent and relaxed datasets) using a relaxed
clock model (lognormal uncorrelated) and with the
Yule process (Yule, 1924; Gernhard, 2008) set as tree
prior (BEAUti; Drummond et al., 2012). After initial
inspection with Tracer v.1.5 (Rambaut and Drum-
mond, 2007), for each dataset the log and tree les
were combined with Logcombiner v.1.7.5 (Rambaut
and Drummond, 2007) disregarding 10 M generations
(10%) as burnin. ESS values were all above 200. Sub-
sequently TreeAnnotator v.1.7.5 (Rambaut and Drum-
mond, 2007) was used (burnin set to zero) to generate
the maximum clade credibility tree for both the com-
bined tree les. Given the use of a single calibration
point and the overall low posterior probabilities, the
obtained ages should be considered indicative only.
Condence intervals (node bars) were so large (espe-
cially for the deeper nodes) that they obscured the tree
and hence were omitted for clarity. Node ages were
rounded to the rst decimal to still visualize the differ-
ences in results between ‘stringent’ and ‘relaxed’ data-
set, not to imply accuracy.
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Groenenberg et al. – Systematics of Ariantinae
Fig. 2. BEAST phylogeny based on the ‘relaxed’ H3-COI-CytB-16S dataset. Branch values show posterior probabilities. Node values
indicate divergence estimates in MYA.
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Contributions to Zoology, 85 (1) – 2016
Genetic distances
Uncorrected pairwise (p) distances were calculated
with Paup 4.0b10 for Unix (Swofford, 2002) for each
protein coding gene. Sequence divergence percentages
were calculated as the uncorrected p-distance times
100.
Systematics
Genera and subgenera are taxonomic ranks. To make
the ranking less subjective, a cladistic approach is re-
quired. Throughout this study, clades that are sister-
groups of one another do not differ in their taxonomic
ranking. This leaves the option open to give the same
ranking also to clades that do not have a sister-group
relationship. As a consequence, a genus may have
more than two subgenera. Since there are many more
splitting points in evolutionary history than taxonomic
ranks, this cannot be avoided. Genera are always based
on monophyletic species groups. In some cases, at the
subgeneric level, paraphyletic taxa are accepted. Ge-
netic distances were not used as decisive in deciding
upon the status of genus versus subgenus.
When morphologically cryptic taxa are unequivo-
cally brought to light by the molecular analyses, these
taxa are not neglected but formally characterized
and
named, as advocated by Cook et al. (2010),
Gittenber ger and Gittenberger (2011) and Jörger and
Schrödl (2013). Abbreviations: PP = posterior proba-
bility, MYA = million years ago.
Results
In total 483 sequences were obtained: 161, 149, 91 and
82 for the markers H3, COI, CytB and 16S, respec-
tively. A summary of the character statistics (as calcu-
lated with Paup 4.0b10 for Unix; Swofford, 2002) for
each dataset is given in Table 3. Table 4 (Appendix)
gives a summary of uncorrected p-distances.
Both the phylogeny reconstructions for the individ-
ual markers (Figs S1-S4; online supplementary infor-
mation), as well as those for the concatenated datasets
(Figs 1, 2, S5, S6) distinguish most of the (sub)genera
that were traditionally characterized by subtle concho-
logical differences and geographic origin (e.g. Arian-
ta, Cattania, Corneola, Chilostoma, Cingulifera, Di-
narica, Faustina, Helicigona, Josephinella, Kosicia
and Liburnica). Although sister-group relationships
between some taxa were shown explicitly (e.g. Jose-
phinella - Thiessea, Ariantopsis - Wladislawia), deeper
nodes were hardly supported, particularly in the phy-
logenies based on the individual markers (Figs S1-S4).
The H3 dataset differs most from the other datasets by
its relatively low percentage of parsimony informative
characters (Table 3). The phylogeny based on this
marker is not discriminative below the genus level, but
can be useful for the assignment of species (or subgen-
era) to genera (e.g. Ariantopsis pelia, Campylaeopsis
moellendorffii, Superba spec., Wladislawia sztolcma-
ni). The Histone gene cluster is multicopy (slight inter-
copy variation might exist and H3 pseudogenes have
been reported; Rooney et al., 2002) and has been used
in higher-level phylogenetics (Armbruster et al., 2005;
Colgan et al., 2007 and references therein). For some
species of mostly Chilostoma, double peaks were ob-
served at a few positions within the H3 sequence. This
genus which is shown as monophyletic in nearly all
phylogeny reconstructions (Figs 1, 2, S2, S3, S5 and
S6), turns out paraphyletic in the phylogeny for H3
(Fig. S1). It might be argued that the more extensive
sampling of Chilostoma (Cingulifera) increased the
chance of observing this apparent intercopy variation,
but it was not observed in other genera for which mul-
tiple species were sequenced (e.g. Arianta, Cattania,
Josephinella and Liburnica) either. In agreement with
Colgan et al. (2000) we therefore conclude that inter-
copy variation in H3 will not signicantly interfere
with the phylogeny reconstructions.
Initially no amplicons were obtained with the COI
mini-barcode primers of Meusnier et al. (2008); to get
these working for Ariantinae, the reverse primer was
modied (Table 2). Hajibabaei et al. (2006) showed (in
silico) that COI mini-barcodes (109 bp; compared to
the full length barcode of 654 bp) are 3% less effective
in the correct identication of closely related species
and pointed out that mini-barcodes might be less use-
ful for the classication of specimens in larger species
assemblages. Based on those taxa for which both a
complete and a mini-barcode sequence were obtained
(C. (Cattania) faueri, Corneola desmoulinsii, Heli-
cigona lapicida andorrica and Vidovicia caerulans),
we conclude that the mini-barcodes (despite their short
length) are placed correctly in the COI phylogeny (Fig.
S2).
The phylogeny reconstructions for the concatenated
datasets (Figs 1, 2, S5-S6) show a basal split within the
Ariantinae, differentiating the ancestor of the genera
Causa, Isognomostoma and Helicigona, referred to
here as group A, from that of all other genera (except
Campylaea and Corneola), collectively referred to as
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Groenenberg et al. – Systematics of Ariantinae
group B. It remains unclear whether Campylaea and
Corneola belong to A (Fig. 2) or B (Figs 1, S5), or
should be considered a group on their own (Fig. S6).
All of the obtained sequences were deposited in
GenBank; a complete list of taxa, sampling informa-
tion and accession numbers is given in Table S7.
Discussion
Left aside some exceptions, Ariantinae shells are rather
monomorphic (Fig. 3, Appendix). Because of the limit-
ed number of conchological characters, many authors
studied the genital tract for morphological characters
that could discriminate species and especially higher
taxa. However, the genital morphology within this sub-
family is also very homogeneous, what is uncommon
among pulmonates. The form of the accessory glands,
which are either undivided or more or less completely
split (Fig. 4, Appendix), has been used by some authors
as a (partial) basis for the systematics of the Ariantinae
(Sturany and Wagner, 1914; Schileyko, 2006), although
according to other authors both types of accessory
glands can occur within the same genus, or even species
(Hesse, 1931; Knipper, 1939; Schileyko, 2013). A clas-
sication of the species of Ariantinae in only two gen-
era, as for example Helicigona and Campylaea (sensu
Sturany and Wagner, 1914; Table 1), or Chilostoma and
Campylaea (sensu Zilch, 1960; Table 1) is an oversim-
plication according to all modern authors, but what
classication should be accepted alter natively remains a
matter of dispute. Recently it has been suggested that
the structure of the penial papilla might be a useful
character to clarify the phylogenetic relationships be-
tween the (sub)genera within the subfamily (Schileyko,
2013), but that view still has to be conrmed.
Obviously, given the actual situation, a new ap-
proach
is necessary, as was realized most recently by
Groenenberg et al. (2012) and Cadahia et al. (2013),
who tried to escape from the confusion by the use of
molecular phylogenetics. Despite its shortcomings in
the completeness of the molecular data, this article ex-
pands the reliability of the molecular phylogeny recon-
structions, enabling a still better founded discussion
regarding the subdivisions of the Ariantinae.
Our analyses do not support an evolutionary signi-
cance of the transformation series based by Schileyko
(2013) on the structure of the penial papilla in several
Ariantinae genera. Dinarica and Cattania are not
closely related to Helicigona, for example, so that the
depicted morphocline Cattania - Helicigona - Dinarica
(Schileyko, 2013) cannot be interpreted in an evolu-
tionary context.
Aiming at a general classication of the Ariantinae,
the shape of the accessory glands is equally uninform-
ative. The transition from undivided to split gland(s),
or the other way round, must have occurred more than
once.
When the phylogeny reconstructions obtained with
this study are compared to generic classications
based on conchology and geography, nearly all the
named (sub)genera are recovered as distinct clades. A
few additional (sub)generic groups were discovered
and described, viz. Campylaea (Oricampylaea),
Chilostoma (Achatica), Cattania (Cattaniella), Pseu-
dotrizona, Kollarix (Table 5, Appendix). The phyloge-
netic relationships above the genus-level, as indicated
by the lower posterior probabilities, remain less cer-
tain in many cases. For a limited number of genera,
sister-group relationships were disclosed, i.e. Arianta-
Cylindrus, Causa-Isognomostoma, Josephinella-
Thiessea
and Kosicia-Faustina (PP ≥ 0.95; Figs 1, 2,
S5, S6). In particular the close relationship between
Cylindrus and Arianta is intriguing. Clearly both gen-
era are part of a lineage that was less restricted in the
development of conchological novelties. The classi-
cation of Cylindrus as a member of the Ariantinae is
now conrmed genetically.
We agree with Cadahia et al. (2013) that indications
of evolutionary age are uncertain, to say the least. The
fossil record is very incomplete indeed, and a molecu-
lar clock model is also not easily applicable. The un-
at
tractive alternative would have been to omit such
speculations altogether.
Acknowledgements
We thank all the colleagues who put specimens at our disposal
(Table S7). We are indebted to H.A. Thomassen and R. Glas
(Leiden) for some of the labwork carried out in the early stages
of this proje ct. Finally we like to thank Dr. Neuber t (Dr. E. Neu-
bert (Bern), Dr. A.A. Schileyko (Moscow) and two anonymous
reviewers for their remarks and constructive comments on an
earlier version of this manuscript.
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Online Supplementary Information
S1. MrBayes phylogeny based on the H3 dataset
S2. MrBayes phylogeny based on the COI dataset
S3. MrBayes phylogeny based on the CytB dataset
S4. MrBayes phylogeny based on the 16S dataset
S5. BEAST phylogeny for the stringent H3-COI-CytB dataset
S6. MrBayes phylogeny based on the relaxed H3-COI-CytB-16S dataset
S7. Taxa and sampling information
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Contributions to Zoology, 85 (1) – 2016
Appendix
Systematics
Most nominal genus-group taxa, viz. Cattania, Cor-
neola, Delphinatia, Dinarica, Faustina, Josephinel-
la, Kosicia, Liburnica and Thiessea, are shown as dis-
tinct clades in our molecular analyses. These taxa are
closely related to neither Campylaea nor Chilostoma
and should no longer be regarded as subgenera of one
of these genera. We consider the Ariantinae a subfam-
ily with 21 genera (Table 5), ve of which are subdi-
vided into two (Campylaea, Dinarica, Liburnica),
three (Chilostoma) or four (Cattania) subgenera. To
some extent, the genetic distances may be used as a
measure for the amount of (dis)similarity between
taxa. For the status of taxa in the taxonomic hierarchy
we use cladistic arguments, i.e. using identical ranking
for clades that are sister-groups. Based on the phylo-
geny reconstructions we recognize the following gen-
era and subgenera, listed in alphabetical order.
Subgenus nov. Achatica (monotypic), genus
Chilostoma
Type species: Helix achates Rossmässler, 1835
Abbreviations. PS = P. Subai; RMNH = Naturalis Bio-
diversity Center, Leiden; SMF = Forschungsinstitut
Senckenberg, Frankfurt am Main
Diagnosis. The diagnosis of this monotypic subge-
nus is by denition identical with that of its type spe-
cies, i.e. Chilostoma (Achatica) achates (Rossmässler,
1835). Achatica subgen. nov. is also differentiated by a
unique combination of nucleotides in the 655 bp COI
sequence obtained with general barcoding primers
(Folmer, 1994) at the following relative positions: 79
C, 88 C, 181 C, 197 A, 211 C, 272 G, 319 C, 352 A, 538
G, 595 C.
Description. Shell strongly depressed, nearly dis-
coid, rather dark, yellowish brown, with a brown spiral
band; umbilicus wide (Kerney and Cameron, 1979: 201,
pl. 21 g. 2; Boschi, 2011: 570-571; Welter-Schultes,
2012: 580). See “Chilostoma (Chilostoma) achates” in
Schileyko (2006: g. 2264B, C; 2013: 143, g. 13) for
details regarding the genital morphology. The acces-
sory glands are undivided.
Molecular data. Two individuals have been used
for the molecular analyses, viz. (a) a specimen col-
lected in the northern limestone Alps (Berchtesgaden,
Bayern, Germany), and (b) a specimen from the south-
ern limestone Alps (Greifenburg, Kärnten, Austria).
The sequence divergences between these specimens,
based on all four markers, is less than 0.2% (Table 4).
When compared to sequence divergences of 3.3-6.5%
between subspecies of Chilostoma (Cingulifera), there
is at least no genetic support for a classication of
these two populations of C. (Chilostoma) achates as
different subspecies (as suggested by Falkner, 1998).
Age. The unresolved sister-group relationships be-
tween the subgenera of Chilostoma (se e Chilostoma
[Chilostoma]) do not allow for an unequivocal esti-
mate for the emergence of Chilostoma (Achatica). If it
dates back to the most basal node within the genus
(Fig. 2), it is estimated at ca. 38.5 MYA (Fig. 2). When
C. (Achatica) and C. (Cingulifera) are sister-groups
(Fig. S5) the most recent common ancestor is estimated
at ca. 24.7 MYA (Fig. S5).
Distribution. Austria, E Switzerland, S Germany
(Bayern, Berchtesgadener Alps), N Italy.
Remarks. It is surprising that only a single, poly-
typic species is classied in Achatica, because Chilos-
toma (C.) adelozona (Strobel, 1857) and Chilostoma
(C.) zonatum (Studer, 1820) have brown shells that
look similar to C. (A.) achates at rst sight. See
Chilostoma (Chilostoma).
Derivatio nominis. The name Achatica is supposed
to recall the name of the type species.
Genus Arianta Turton, 1831
Type species: Helix arbustorum Linné, 1758
Molecular data. Four Arianta species could be stud-
ied, viz. A. aethyops (Bielz, 1851), A. arbustorum s. lat.
(with ve subspecies, two of which are considered
separate species by some authors [Welter-Schultes,
2012]), A. chamaeleon (Pfeiffer, 1868), and A. schmidtii
(Rossmässler, 1836). The monophyly of this broadly
accepted genus is supported in all molecular phyloge-
ny reconstructions (PP = 1.0; Figs 1, 2, S1-S6). The
position of A. arbustorum stenzii (see Gittenberger et
al. 2004) or A. arbustorum stenzii-arbustorum in phy-
logeny reconstructions presented in this study does not
give any support for the introduction of Altarianta
Schileyko, 2013, as a subgenus of Arianta. This is in
accordance with the fact that A. a. stenzii and A. a.
arbustorum, hybridize where they are in contact. Ari-
anta chamaeleon, which is shown as the sister-group
of the other Arianta species, is less closely related. The
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Groenenberg et al. – Systematics of Ariantinae
Fig 3. Compilation of shells representing
most of the currently recognized genera of
Ariantinae. 1. Arianta arbustorum (Lin-
naeus, 1758) B 24.1 mm [RMNH G2131]
Austria, Steiermark, near Gstatterboden;
E. Gittenberger leg., 10-IX-1964. 2. Ari-
anta chamaeleon wiedermayeri (Kobelt,
1903) B 18.3 mm [RMNH G2608] Aus-
tria, East Tirol, S of Kartitsch; E. Gitten-
berger leg., VIII-1974. 3. Cylindrus ob-
tusus (Draparnaud, 1805) B 13.7 mm
[RMNH 73877] Austria, Oberösterreich,
Bledigupf; W.H. Neuteboom leg., 22-VII-
1966. 4. Isognomostoma isognomostomos
(Schröter, 1784) B 9.7 mm [RMNH 74226]
Austria, Kärnten, Plöckenpass; W.H. Neu-
teboom leg., 14-IX-1952. 5. Causa holos-
ericea (Studer, 1820) B 10.5 mm [RMNH
74311] Austria, Salzburg, Amerthal; W.H.
Neuteboom leg., 16-VII-1968. 6. Chilosto-
ma (Achatica) achates (Rossmässler,
1835) B 21.9 mm [RMNH G2410] Austria,
Steiermark, E of Brandtriedl; A. and E.
Gittenberger leg., 19-V-1972. 7. Chilo-
stoma (Chilostoma) zonatum rhaeticum
(Strobel, 1857) B 25.2 [RMNH G54412]
Switzerland, Graubünden, E of Martins-
bruck; E. Gittenberger leg., IX-1963. 8.
Chilostoma (Chilostoma) tigrinum (De
Cristofori and Jan, 1832) B 24.7 mm
[RMNH 73434] Italy, Como, Pasturo;
W.H. Neuteboom leg., 3-VIII-1954. 9.
Chilostoma (Cingulifera) cingulatum cin-
gulatum (Studer, 1820) B 20.5 mm
[RMNH H1938] Switzerland, Tessin,
Mel ide along Lago di Lugano; J.T. Henrard
leg., 28.VIII.1938. 10. Chilostoma (Cingu-
lifera) cingulatum gobanzi (Frauenfeld,
1867) B 22.8 mm [RMNH 73408] Italy,
Brescia, Val Toscolano; W.H. Neuteboom
leg., 05-VIII-1954. 11. Delphinatia fon-
tenillii alpina (Michaud, 1831) B 19.6 mm
[RMNH G3646] France, Isère, SSE of Laurent-du-Pont; E. Gittenberger leg., 12-IX-1975. 12. Faustina faustina (Rossmässler, 1835) B
19.6 mm [RMNH 53576] Hungary, Bükk, Szalajkavölgy; Agócsy leg., 21-V-1921. 13. Campylaea (Campylaea) planospira planospira
(Lamarck, 1822) B 25.9 mm [RMNH 73625] Italy, Torino, Santvaris di Montebruno; W.H. Neuteboom leg., 8-VII-1977. 14. Campylaea
(Oricampylaea) illyrica (Stabile, 1864) B 25.4 mm [RMNH 11124] Italy, Friuli, SSE of Tarvisio; E. Gittenberger leg., VI-1992. 15. Kosi-
cia ambrosi (Strobel, 1852) B 12.5 mm [RMNH 24940] Italy, Vicenza, Valstagna; W.H. Neuteboom leg., 22-VII-1968. 16. Kosicia inter-
media (Pfeiffer, 1828) B 15.3 mm [RMNH 54440] Austria, Kärnten, Deutschpeter; E. Gittenberger leg., IX-1964. 17. Kosicia ziegleri
(Rossmässler, 1836) B 17.7 mm [RMNH G2350] Slovenia, Kamniške Alps, Igla Studenec; A. and E. Gittenberger leg., 26-VIII-1971.
conchologically unexpected sister-group relationship
(PP = 1.0; Figs 1-2, S1, S3-S6) between Arianta and
Cylindrus (Fig. 3.3) was shown by Groenenberg et al.
(2012) and has recently been conrmed by Cadahia et
al. (2013).
Age. The most recent common ancestor for Arianta
is estimated at ca. 23.1-22.4 MYA (Figs. S5, 2).
Distribution. Arianta arbustorum (Fig. 3.1) has the
largest distribution range of all the species within the
subfamily Ariantinae. It occurs in north and central
Europe, from Iceland, Norway, Sweden, N.-Ireland,
Great Britain, and central France eastwards to the Bal-
tic countries, Poland, Ukraine and Romania (Carpathi-
ans). The southern border ranges from the N.-Italian
Alps through Slovenia, Croatia and Serbia into Bul-
garia (up to Stara Planina); except for some localities
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51
Contributions to Zoology, 85 (1) – 2016
18. Campylaeopsis moellendorffii (Ko -
belt, 1871) B 18.4 mm [RMNH AJW898]
Bosnia-Herzegovina, Vrelo Bosne, near
Ilidza; A.J. de Winter leg., 17-IX-1980. 19.
Kollarix kollari (Pfeiffer, 1856) B 24.3
mm [RMNH YU.429] Serbia, along Ovcar
Banja; W.J.M. Maassen leg., V-1984. 20.
Liburnica (Liburnica) setosa setosa (Fé-
russac, 1832) B 23.6 mm [RMNH 94272]
Croatia, Lovrec, S of Imotski; W.J.M.
Maassen leg., IV-1989. 21. Dinarica (Sab-
ljaria) stenomphala (Menke, 1830) B 30.7
mm [col. PS.21228] Croatia, Velebit Mts,
near Krasno Polje; P. Subai leg., 29-VII-
2002. 22. Dinarica (Dinarica) pouzolzii
(Deshayes, 1830) B 40.1 mm [RMNH
53508 / 413] Croatia, Dalmatia, E of Biok-
ovo Mts; J.J. ter Pelkwijk leg., 12-VIII-
1939. 23. Liburnica (Superba) skipetarica
skipetarica (Subai, 1995) B 19.6 mm [col.
PS.20215] Albania, Periferi Berat, Tom-
morit; P. Subai leg., VIII-2004. 24. Cor-
neola desmoulinsii (Farines, 1834) B 17.2
mm [RMNH 93925] Andorra, Canillo,
northern wall; W.J.M. Maassen leg., VII-
1990. 25. Helicigona lapicida lapicida
(Linnaeus, 1758) B 17.6 mm [SMF3254
26/1] Germany, Hessen, Schlüchtern; M.
Pfenninger leg. 26. Drobacia banatica
(Rossmässler, 1838) B 29.5 mm [RMNH
54500 / 485] Romania, Siebenbürgen; H.
de Wever leg. 27. Cattania (Cattania) tri-
zona (Rossmässler, 1834) B 24.4 mm
[RMNH 99615] Romania, Banat Mts, Mt
Domogled; Kroupa leg., 21-VI-1985. 28.
Cattania (Cattania) subaii (Fauer, 1991) B
22.9 mm [RMNH GU.9921 / EK5558]
Greece, Makedonia, W of Kozani; E. Git-
tenberger and D. Uit de Weerd leg., 23-V-
1999. 29. Vidovicia caerulans (Pfeiffer,
1828) B 15.4 mm [RMNH 93836] Croatia,
Velebit near Starigrad; W.J.M. Maassen
leg., IX-1982. 30. Cattania (Ariantopsis) pelia (Hesse, 1912) B 17.9 mm [col. PS.23572] Bulgaria, Vitosha, Bistrisko branishte; I. Dedoy
leg., 8-VII-2004. 31. Cattania (Wladislawia) polinskii (Wagner, 1928) B 16.4 mm [RMNH G3749] Bulgaria, Pirin Mts, Mt Vihren; A.
Riedel leg., 24-VI-1977. 32. Cattania (Wladislawia) sztolcmani (Wagner, 1928) B 10.6 mm [RMNH G3749] Bulgaria, Pirin Mts, Mt
Vihren; A. Riedel leg., 24-VI-1977. 33. Josephinella vikosensis (Subai, 1990) B 18.8 mm [R MNH EG.9703 / DK8112] Greece, Ipiros, Vikos
valley; E. Gittenberger leg., 23-VII-1997. 34. Josephinella hemonica (Thiesse, 1884) B 19.1 mm Greece, Makedhonia, SE of Grevena; E.
Gittenberger leg., 18-VII-1986. 35. Thiessea sphaeriostoma (Bourguignat, 1857) B 21.0 mm [RMNH 75078] Greece, Sterea Ellas, SE of
Mariolates; E. Gittenberger and D. Uit de Weerd leg., 19-V-2000.
in the Spanish Pyrenees south of the watershed (A. ar-
bustorum xatarti Farines, 1834) it does not occur in
the Iberian peninsula.
Remarks. Four or ve Arianta species have been
described, some of which are polytypic. Campylaea
apfelbecki Sturany, 1901, which was considered a sub-
species of A. chamaeleon by Knipper (1939), could not
be investigated; it might be either a fth Arianta spe-
cies or belong to Cattania (Cattaniella). The excep-
tionally widespread Arianta arbustorum arbustorum
is aberrant also in terms of shell morphology and in its
ecological requirements, occurring independently of
limestone from the lowland to high in the mountains.
The other Arianta species are restricted to (high) al-
pine habitats. While nearly all Ariantinae have a de-
pressed shell and an open umbilicus, A. a. arbustorum
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Groenenberg et al. – Systematics of Ariantinae
has a globular shell with a closed umbilicus (Gitten-
berger et al., 2004). Other Arianta species, such as A.
chamaeleon (Fig. 3.2) and A. schmidtii retained the
plesiomorphic, depressed shell phenotype. Some sub-
species of A. arbustorum that are characterized by de-
pressed shells (Gittenberger et al., 2004; Haase and
Misof, 2009) might have evolved that character state
by reversal. See also Arianta in Schileyko (2013) for
details regarding the genital morphology. The acces-
sory glands are undivided (Fig. 4.1). Specimens of A.
arbustorum stenzii from several localities should be
dissected to investigate the status of Altarianta Schi-
leyko, 2013, in more detail.
Subgenus Ariantopsis Wagner, 1928 (monotypic), ge-
nus Cattania
Type species: Helicigona (Arianta) pelia Hesse, 1912
Molecular data. Both MrBayes and BEAST phyloge-
ny reconstructions for the relaxed dataset highly sup-
port a sister-group relation (PP = 1.0) between Arian-
topsis and Wladislawia (Figs 2, S6), which are here
considered subgenera of Cattania Brusina, 1904.
Age. The most recent common ancestor of Cattania
(Ariantopsis) and C. (Wladislawia) is estimated at ca.
7.3 MYA (Fig. 2).
Distribution. Ariantopsis is endemic to SW and W
Bulgaria. The eastern boundary of its distribution is
situated near Plovdiv, the northern boundary is near
Lakatnik in the Iskar-valley, and its western distribu-
tion is formed by Mt Vitosha and the Rila Mts.
Remarks. The taxonomic position of Cattania
(Ariantopsis) pelia (Fig. 3.30) has long been uncertain.
Conchologically it somewhat resembles Arianta aethy-
ops. It has been assigned to various genera, viz. Arianta
by Kroupa (1994) and Dedov (1998), Helicigona by
Hesse (1912), Chilostoma by Bank et al. (20 01) a n d
Faustina by Damjanov and Likharev (1975). See also
Campylaea (Ariantopsis) and Ariantopsis in Schileyko
(2006, 2013) for details regarding the shell and genital
morphology. The accessory glands can be undivided,
but are mostly split up to half of their length (Fig. 4.29).
Subgenus Campylaea Beck, 1837 (monotypic?), genus
Campylaea
Type species: Campylaea planospira Lamarck, 1822
Molecular data. Campylaea (C.) planospira (Fig.
3.13)
is the sister-group of a clade with three species,
referred to below as Campylaea (Oricampylaea) (PP
> 0.92; Figs S1, S2). Together, the subgenera Campy-
laea and Oricampylaea, form a monophyletic group
(PP = 1.0; Figs 2, S1, S2, S4, S6), viz. the genus Campy-
laea. The genetic distances between C. (Campylaea)
and C. (Oricampylaea) are comparatively large (COI
sequence divergence up to 22.1%; Table 4). It is unclear
to which genus Campylaea is most closely related. Ex-
cept for the phylogeny reconstruction according to
COI, which suggest a sister-group relationship between
Campylaea and Kollarix (PP = 0.86; Fig. S2), none
of
the other datasets provides information regarding
possible sister-group relationships of Campylaea. In
the phylogenies based on the concatenated datasets
Campylaea branches off early in either group A (Figs
1, S6) or group B (Fig. 2).
Age. The most recent common ancestor of Campy-
laea is estimated at ca. 34.0 MYA (Fig. 2).
Distribution. Campylaea (C.) planospira is repre-
sented in S Austria, N Balkans, mainland Italy, and the
island of Sicily.
Remarks. See also Campylaea (Campylaea) and
Chilostoma (Campylaea) in Schileyko (2006, 2013)
for details regarding the shell and genital morphology.
The accessory glands can be undivided (Fig. 4.14), but
Sturany and Wagner (1914) and Knipper (1939) showed
split accessory glands in C. (Campylaea) planospira.
Penial papilla small, conical, with ne transverse ridges
and an obtuse apex with a short, transverse, slit-like
pore. Secondary ureter entirely open. For the moment
being, only a single, polytypic species is accepted in
Campylaea s. str. Some of the so-called subspecies
could be considered separate species, however.
Genus Campylaeopsis Sturany and Wagner, 1914
(monotypic)
Type species: Helicigona moellendorffii Kob elt, 18 71.
Molecular data. Only an H3 sequence was obtained for
this taxon. In the respective phylogeny Campylaeopsis
is placed in a clade with Delphinatia, Drobacia,
Pseudotrizona, and Vidovicia (PP = 0.8; Fig. S1).
Campylaeopsis moellendorffii shares a substantial
part of its distribution area with Pseudotrizona in-
flata.
Age. Not enough sequence information was obtained
to include Campylaeopsis in the time calibrated analy-
ses (Figs. 2, S5).
Distribution. The mountains of Bosnia-Herzegowi-
na and Montenegro.
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Contributions to Zoology, 85 (1) – 2016
Remarks. Campylaeopsis moellendorffii (Fig. 3.18)
has a characteristic shell with regularly arranged,
widely spaced hairs. It has been assigned to Helicigo-
na by Knipper (1939) and to Chilostoma by Bank
(2 0 01).
See also Helicigona (Campylaeopsis) and Chilos-
toma (Campylaeopsis) in Schileyko (2006, 2013) for
details regarding the shell and genital morphology.
The accessory glands are undivided (Fig. 4.10).
Subgenus Cattania Brusina, 1904, genus Cattania
Type species: Helix trizona Rossmässler, 1835
Molecular data. Five of the ca. 9 Cattania (C.) species
are included in this study. The genus Cattania is shown
to be monophyletic (PP = 1.0; all pylogenies except
S1). It includes the subgenera: Cattania (Ariantopsis),
C. (Cattania), Cattania (Cattaniella) and C. (Wladis-
lawia). Cattania is the sister-group of the clade Jose-
phinella-Thiessea (PP ≥ 0.8; Figs 1, 2, S1-S3 and S6).
Cattania (Cattania) constitutes a monophyletic group
within Cattania (PP = 1.0; Fig. S2) and is in this study
represented by C. (C.) faueri, C. (C.) kattingeri, C.
(C.) pseudocingulata, C. (C.) subaii and the type spe-
cies C. (C.) trizona. It is the sister-group of the clade
C. (Ariantopsis)-C. (Wladislawia) (PP ≥ 0.95; Figs
S2, S6). The COI sequence divergences between C.
(Cattania) and C. (Ariantopsis), and between the
former and C. (Wladislawia) are 13.4% and 14.7%,
re
spectively. The data of Cadhaia et al. 2013 show that
C. (Cattania) haberhaueri belongs to this subgenus as
well. Future research will have to make clear whether
C. balcanica and C. rumelica should also be classied
here. All phylogeny reconstructions indicate that the
species referred to as Cattania inflata (Kobelt, 1876)
by Subai (1995) represents a separate lineage (Figs 1,
2, S1-S6) that is clearly distinct from Cattania. We
consider this lineage a separate genus, referred to be-
low as Pseudotrizona gen. nov.
Age. The most recent common ancestor of Cattania
is estimated at ca. 27.6-26.9 MYA (Figs 2, S5).
Distribution. Central Balkans, SW Romania, E and
S Serbia, SW Bulgaria and N Greece (Thraki).
Remarks. See Campylaea (Cattania) and Chilos-
toma (Cattania) in Schileyko (2006, 2013) for details
Tab le 4. Uncorrected p-distances (in percentages) for a selection of taxa.
From To H3 COI CB
25 Campylaea (Campylaea) planospira 26 Campylaea (Oricampylaea) illyrica 2,4 22,1 -
25 Campylaea (Campylaea) planospira 30 Campylaea (Oricampylaea) lefeburiana 2,4 20,5 -
41 Cattania (Cattania) subaii 34 Cattania (Ariantopsis) pelia 0 13,4 18
41 Cattania (Cattania) subaii 46 Cattania (Wladislawia) sztolcmani 0,3 14 ,7 -
41 Cattania (Cattania) subaii 45 Cattania (Cattaniella) thateensis 0,3 15,4 -
54 Causa holosericea 125 Isognomostoma isognomostomos 1,2 20,2 23
63 Chilostoma (Chilostoma) zonatum adelozona 83 Chilostoma (Cingulifera) cingulatum preslii 0 14,5 18 ,6
63 Chilostoma (Chilostoma) zonatum adelozona 57 Chilostoma (Achatica) achates 0 14,7 18,6
83 Chilostoma (Cingulifera) cingulatum preslii 57 Chilostoma (Achatica) achates 0 14,8 18
57 Chilostoma (Achatica) achates 58 Chilostoma (Achatica) achates 0 0,2 0,3
72 Chilostoma (Cingulifera) cingulatum peregrini 74 Chilostoma (Cingulifera) cingulatum preslii 0,3 0,6 0,6
72 Chilostoma (Cingulifera) cingulatum peregrini 86 Chilostoma (Cingulifera) cingulatum preslii 0,3 0 0
90 Corneola squamatinum 88 Corneola desmoulinsii 0,6 16,9 -
94 Cylindrus obtusus 18 Arianta chamaeleon 0,9 18,8 24,9
94 Cylindrus obtusus 9 Arianta arbustorum arbustorum 0,9 20 23,8
100 Dinarica (Dinarica) pouzolzii 102 Dinarica (Dinarica) serbica 0,9 11 13,6
100 Dinarica (Dinarica) pouzolzii 103 Dinarica (Sabljaria) stenomphala 1,8 16 24,7
110 Faustina faustina orba 111 Faustina kiralikoeica 0 17,4 -
107 Faustina faustina associata 108 Faustina faustina faustina 0 10,7 13
113 Helicigona lapicida andorrica 115 Helicigona lapicida andorrica 0 0,3 0,3
119 Helicigona lapicida lapicida 117 Helicigona lapicida lapicida 0,6 1,8 0,6
113 Helicigona lapicida andorrica 117 Helicigona lapicida lapicida 0,6 11,8 14,1
154 Liburnica (Liburnica) setosa 149 Liburnica (Liburnica) albanograeca 0 3,4 -
150 Liburnica (Liburnica) dunjana 153 Liburnica (Liburnica) setigera setigera 0 6,3 -
158 Liburnica (Superba) skipetarica 156 Liburnica (Superba) kulmankana 0 2,1 -
153 Liburnica (Liburnica) setigera setigera 157 Liburnica (Superba) skipetarica 0 6,9 -
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Groenenberg et al. – Systematics of Ariantinae
regarding the shell and genital morphology. In C.
(Cattania) the accessory glands are usually split (Fig.
4.27); occasionaly one of the glands is undivided.
Subgenus nov. Cattaniella, genus Cattania
Type species: Helix cingulata olympica Roth, 1855
Diagnosis. The two Cattaniella species share a unique
combination of nucleotides in the 655 bp COI sequence
obtained with general barcoding primers (Folmer,
1994) at the following relative positions: 87 A, 95 T, 235
A, 331 G, 365 G, 406 T, 499 G, 542 A, 543 G, 583 A.
Description. Shell depressed globular to low coni-
cal, nearly discoid, whitish, corneous or brown, with
one to three brown spiral bands. The accessory glands
in C. (Cattaniella) thateensis are split (Subai, 2012).
Molecular data. Cattania (Cattaniella) is repre-
sented by C. (Cattaniella) olympica and C. (Catta-
niella) thateensis (Subai, 2012). It is the sister-group of
the combined three other subgenera (Figs 2, S2, S3,
S6) of Cattania. Clearly C. (Cattaniella) olympica
should no longer be considered a subspecies of C.
(Cattania) trizona (see Knipper, 1939). Likewise, C.
(Cattaniella) thateensis cannot be classied in Wladis-
lawia (see Subai, 2012). Future research will have to
show whether Campylaea apfelbecki Sturany, 1901
and Campylaea zebiana Sturany, 1907 belong to Cat-
tania (Cattaniella) as well (Subai, 2012).
Distribution. Higher montane areas of E Albania
(Thäte mountains); Olympos and Ossa mountain areas
of Thessaly, Greece.
Remarks. For the moment being, only the two spe-
cies that could be investigated for this study are classi-
ed in Cattaniella.
Derivatio nominis. Cattaniella refers to Cattania.
Genus Causa Schileyko, 1971 (monotypic)
Type species: Glischrus (Helix) holosericea St ude r,
1820
Molecular data. The sister-group relationship, as well
as a substantial genetic distance between Causa and
Isognomostoma are established (PP = 1.0; Figs 1, 2,
S2-S6; Table 4). Only the H3 data failed to show a di-
rect sister-group relation, but still placed both genera
in the same clade (PP = 0.42, Fig. S1).
Age. The most recent common ancestor of Causa
and Isognomostoma is estimated at ca. 33.0-30.1 MYA
(Figs 2, S5).
Distribution. Alps, Sudetes and W Carpathians
(Tatra Mts), isolated in S Germany (Franconian Jura).
Remarks. Conchologically, Causa holosericea (Fig.
3.5) and Isognomostoma isognomostomos (Fig. 3.4)
are both aberrant among the Ariantinae by the dentate
aperture. These species were considered congeneric
until Schileyko (1971), primarily based on differences
in genital anatomy, introduced Causa as a new genus.
See also Causa in Schileyko (2006, 2013) for details
regarding the shell and genital morphology. The ac-
cessory glands in both Causa and Isognomostoma are
undivided (Fig. 4.7-4.8).
Subgenus Chilostoma Fitzinger, 1833, genus Chilos-
toma
Type species: Glischrus (Helix) foetens Studer, 182 0
Molecular data. Four or ve species can be classied
in Chilostoma s. str., three of which are included in
this study. Within the genus Chilostoma (PP = 1.0;
Figs 1, 2, S2, S3, S5, S6), three well supported clades
can be discerned: I) Chilostoma (Chilostoma), II)
Chilostoma (Cingulifera) Held, 1838 and III) Chilos-
toma (Achatica) subgen. nov. The COI sequence diver-
gences between each of the three subgenera are about
15%. Chilostoma is the sister-group of all other Arian-
tinae taxa in group B (PP ≥ 0.6, Figs 1, 2, S6). Only
Figs S3 and S5 specically indicate Corneola as its
sister-group (PP = 0.94). The latter relationship is not
observed if both of the studied Corneola species are
included (Figs 2, S1, S2, S6). The phylogenetic rela-
tionships between the subgenera of Chilostoma are not
resolved. Figs 2, S4 and S6 support (PP ≥ 0.95) a sister-
group relationship between C. (Chilostoma) and C.
(Cingulifera), whereas Figs 1, S3 and S5 indicate
Chilostoma (Achatica) as the sister-group of C. (Cin-
gulifera) (PP ≥ 0.95).
Age. The most recent common ancestor of the ge-
nus Chilostoma is estimated at ca. 38.5-31.3 MYA,
whereas that of C. (Chilostoma) is estimated at ca.
20.3-18.4 MYA (Figs 3 and S5).
Distribution. The Alps (SE France, S Switzerland,
N Italy).
Remarks. Unexpectedly, from a conchological per-
spective, C. (Chilostoma) zonatum (Fig. 3.7) turns out
to be more closely related to C. (Chilostoma) frigidum
and C. (Chilostoma) tigrinum (Fig. 3.8), than to C.
(Achatica) achates (Fig. 3.6), which shares the chest-
nut brown colour of the shell. That colour might be the
plesiomorphic character state in Chilostoma. Nowa-
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days, Helix foetens is either synonymised with C.
(Chilostoma) zonatum, as by Turner et al. (1998) or it
is considered a subspecies of that species (Bank et al.,
2001). In the past many subgenera have been assigned
to Chilostoma (Zilch, 1960; Bank et al., 2001). It is
unclear which, if any, character states of the genital
tract are diagnostic for the subgenera of Chilostoma.
The accessory glands are undivided (Fig. 4.3, 4.4).
Subgenus Cingulifera Held, 1838 (monotypic), genus
Chilostoma
Type species: Glischrus (Helix) cingulata Stude r,
1820. The type species can be subdivided into several
subspecies, ve of which are included in this study
Molecular data. See Chilostoma (Chilostoma).
Age. The most recent common ancestor of C. (Cin-
gulifera) is estimated at ca. 7.9-7.0 MYA (Figs 2, S5).
The split between C. (Cingulifera) and either C.
(Chilostoma) or C. (Achatica) subgen. nov. (see
Chilostoma) is estimated at ca. 29 and 24.7 MYA,
respectively.
Distribution. NE Italy, SW Austria, SE Switzer-
land, locally in the French Alps, Central Italy and S
Germany (partly introduced). Chilostoma (Chilosto-
ma) and C. (Cingulifera) have a parapatric distribu-
tion. Generally the former subgenus is distributed in
the western Alps, whereas the latter one has its main
range in the eastern Alps. Additionally, our prelimi-
nary results indicate a strong separation between the
Chilostoma species east versus west of the Camonica
valley (Valcamonica, Italy).
Remarks. Chilostoma (Cingulifera) is a generally
accepted subgenus of Chilostoma (Zilch, 1960; Bank
et al., 2001). Taxonomically it was supposed to encom-
pass only a single species, i.e. Chilostoma (Cinguli-
fera) cingulatum (Studer, 1820) (Fig. 3.9, 3.10) with a
large number of alleged subspecies (Pfeiffer, 1951),
some of which are here classied differently, however,
viz. Chilostoma (C.). frigidum and Chilostoma (C.)
tigrinum (De Cristofori and Jan, 1832; Fig. 3.8).
Chilostoma (Cingulifera) cingulatum peregrini
Falkner, 1998 was introduced as a replacement name
for Chilostoma (Cingulifera) cingulatum cingulina
(Strobel, 1844), not Helix cingulina Deshayes, 1839 (in
Férussac and Deshayes). Contrary to the prevailing
view, Falkner suggested that the northern alpine popu-
lations of Chilostoma (Cingulifera) cingulatum might
belong to two instead of only a single subspecies, viz.
Chilostoma (Cingulifera) c. peregrini from near Inns-
bruck (Austria) and Chilostoma (Cingulifera) c. pres-
lii from near Berchtesgaden (Falkner, 1998; Kierdorf-
Traut, 2012). COI and CytB sequences for specimens
from both northern alpine localities (Table S7, 72-73
versus 83-86) are virtually identical and differ about
0.5 % (Table 4) from sequences of southern alpine, un-
disputed Chilostoma (Cingulifera) c. preslii (78-82,
Table S7). These data indicate that Chilostoma (Cin-
gulifera) c. peregrini is a junior synonym of Chilosto-
ma (Cingulifera ) c. preslii, which has a disjunct range,
occurring in both the northern and the southern lime-
stone Alps. See Helicigona (Cingulifera) and Chilos-
toma (Cingulifera) in Schileyko (2006, 2013) for de-
tails regarding the shell and genital morphology.
Chilostoma (Cingulifera) has undivided accessory
glands (Fig. 4.4).
Genus Corneola Held, 1838
Type species: Helix cornea Draparnaud, 1801
Molecular data. Corneola squamatinum (Rossmässler,
1835) and C. desmoulinsii (Farines, 1834) together are
monophyletic (PP = 0.86 and 0.77; Figs 2, S6). The
COI sequence divergence between these species is
16.9% (Table 4). The phylogeny reconstructions are
indistinct regarding the position of this genus. In the
concatenated analyses it is shown between Campylaea
and Chilostoma (Fig 1), as the sister-group of either of
these (Figs S6 and S5) or as the sister-group of Causa,
Isognomostoma and Helicigona (Fig. 2). Corneola is
here regarded as a genus.
Age. The most recent common ancestor of Corneola
is estimated at ca. 52.5 MYA (Fig. 2).
Distribution. Corneola acrotricha (Fischer, 1877)
and C. desmoulinsii are mainly found in the Pyrenees.
Corneola squamatinum extends also further into
southern and central France, along the Atlantic coast
up to Brittany, whereas C. crombezi (Bourguignat,
1880) inhabits the Alpes-Maritimes (Falkner et al.,
20 02).
Remarks. In the most recent literature (Bank et al.,
20 01; Falkner et al., 2002) Corneola is regarded as a
subgenus of Chilostoma, with four species. Two of
these, viz. Corneola desmoulinsii (Fig. 1.24) and C.
squamatinum, are included in this study. See Corneo-
la in Schileyko (2013) for details regarding the shell
and genital morphology. The accessory glands are un-
divided (Fig. 4.5)
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Groenenberg et al. – Systematics of Ariantinae
Genus Cylindrus Fitzinger, 1833 (monotypic)
Type species: Pupa obtusa Draparnaud, 1805
Molecular data. The phylogeny reconstructions for
the combined datasets show strong support for a sister-
group relation between Cylindrus and Arianta (PP =
1.0, Figs 1, 2, S5, S6). Only the phylogeny based on
COI fails to indicate that these taxa are sister-groups,
thus sharing a unique common ancestor. The sequence
divergence between Cylindrus and Arianta is ca. 19%
for COI and up to 25% for CytB (table 4).
Age. The common ancestor of Arianta and Cylin-
drus is estimated to have diverged at ca. 47.6-46.4
MYA (Figs 2, S5).
Distribution. Endemic to the Austrian Alps (be-
tween 1600 and 2500 m), known from Oberösterreich,
Niederösterreich, Salzburg, Steiermark and Kärnten.
Remarks. Among the (sub)genera of Ariantinae that
can be distinguished by shell-morphology, Cylindrus
is the most distinctive because the shell is cylindrical
and much higher than broad (Fig. 3.3). Its sister-group,
the genus Arianta, is characterized by much larger
shells that vary in shape between attened and globu-
lar. This close relationship, which is surprising in view
of the morphological data, was reported by Groenen-
berg et al. (2012) and later on conrmed by Cadahia et
al. (2013). Despite the long geological history of Cylin-
drus that is indicated by the molecular data and is also
suggested by its aberrant shell morphology, no clear
fossil representatives of this genus, or forms that are
transitional in shell-shape, are known from before the
Würm (Zilch, 1960; Frank, 2006). See Cylindruini in
Schileyko (2006, 2013) for details regarding the struc-
ture of the genital tract. Cylindrus has undivided ac-
cessory glands (Fig. 4.2).
Genus Delphinatia Hesse, 1931
Type species: Helix alpina Michaud, 1831
Molecular data. Delphinatia fontenillii alpina
(Michaud, 1831), and D. glacialis (Férussac, 1832) to-
gether are monophyletic (PP ≥ 0.93, Figs 2, S1, S2, S6)
and form a clade with Drobacia and Vidovicia in the
phylogeny reconstructions for the combined datasets
(0.5 ≥ PP ≥ 0.86, Figs 1, 2, S5, S6). However, in the
trees based on individual markers, this clade is only
observed with H3 (PP = 0.8, Fig. S1). There is no con-
sensus regarding the sister-group relations of these
three taxa. Only of D. f. alpina sufcient sequence
data were obtained to include it in the phylogeny re-
constructions of the stringent dataset (Figs 1, S5).
Age. The most recent common ancestor of the
com
bined group Delphinatia-Drobacia-Vidovicia is
estimated at ca. 59.1-53.3 MYA (Figs 2, S5); that of
Delphinatia is estimated at ca. 17.5 MYA (Fig 2).
Distribution. French Alps (departments of Hautes-
Alpes, Haute-Savoie, Isère and Savoie) to the adjacent
Italian Alps (Alpi Cozie and Graie) (Gavetti et al.,
20 08).
Remarks. Delphinatia is considered a subgenus of
Chilostoma by Bank et al. (2001), but has been classi-
ed as a subgenus of Campylaea as well (Zilch, 1960).
Only two species are generally recognized in Delphi-
natia, viz. D. fontenillii (Michaud, 1829), and D. gla-
cialis, which are both included in this study. Falkner et
al. (2002) distinguished D. f. fontenillii and D. f. alpi-
na (Fig. 3.11) next to the monotypic D. glacialis. See
Campylaea (Delphinatia) and Chilostoma (Delphina-
tia) in Schileyko (2006, 2013) for details regarding the
shell and genital morphology. The accessory glands
(Fig. 4.12) are undivided, or one of them is split for up
to 25-50% of its length.
Subgenus Dinarica Kobelt, 1902, genus Dinarica
Type species: Helix pouzolzii Deshayes, 1830
Molecular data. Two subgenera of Dinarica can be
recognized, viz. Dinarica (Dinarica) and D. (Sablja-
ria). In this study, the former taxon is represented by
D. (Dinarica) pouzolzii (Fig. 3.22) and D. (D.) serbica
Kobelt, 1872. Dinarica (Dinarica), as well as the ge-
nus itself, are shown to be monophyletic (PP = 1, Figs
1, 2, S5, S6). The COI sequence divergence between D.
(Dinarica) serbica and D. (Dinarica) pouzolzii is
about 10% (Table 4).
The phylogenies based on the concatenated data-
sets slightly differ regarding to the position of Dina-
rica. Figures 2, S5 and S6 suggest a sister-group rela-
tionship between Dinaricia and Liburnica (0.44 ≤ PP
≤ 0.78). In these gures Dinarica - Liburnica has a
sister-group relation with the clade Kollarix - Pseudo-
trizona - Cattania - Thiessea - Josephinella. Basi-
cally Fig 1 shows the same topology, but here Libur-
nica is the sister-group of the latter genera including
Dinarica.
Age. The most recent common ancestor of Dinarica
is estimated at ca. 37.1-36.2 MYA (Figs 2, S5).
Distribution. Along the NE coast of the Adriatic
sea, in SE Croatia, Bosnia-Herzegowina, S Servia,
Montenegro, Kosovo, Albania, the western border of
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Contributions to Zoology, 85 (1) – 2016
Macedonia and NW Greece. Dinarnica (D.) pouzolzii
and D. (D.) serbica have partially overlapping ranges.
Where the former dominates the coastal region of
Montenegro, the latter occurs more inland.
Remarks. See Campylaea (Dinarica) and Chilos-
toma (Dinarica) in Schileyko (2006, 2013) for details
regarding the shell and genital morphology. Dinarica
(Dinarica) has split accessory glands (Fig. 4.23).
Genus Drobacia Brusina, 1904
Type species: Helix banatica Rossmässler, 1838
Molecular data. Both Drobacia species, viz. D. ba-
natica (Fig. 3.26) and D. cf maeotica Wenz, 1926 (in
Krejci and Wenz, 1926), are included in this study. The
taxon is shown to be monophyletic (PP = 1.0 Figs 1, 2,
S5, S6). The position of Drobacia within the subfami-
ly Ariantinae is still unclear. The phylogeny recon-
structions indicate that Drobacia forms a clade with
Delphinatia and Vidovicia (se e Delphinatia). Only the
phylogeny for COI supports a sister-group relationship
with Liburnica (PP = 0.86; Fig. S2).
Age. For an age estimation of the most recent com-
mon ancestor of Drobacia, Delphinatia and Vidovicia,
see Delphinatia. Drobacia banatica and D. cf maeot-
ica are estimated to have diverged ca. 8.7-7.8 MYA
(Figs 2, S5).
Distribution. W and SW Romania and locally in E
Hungary. In the Pleistocene Drobacia reached as far
as the Harz Mts in Thüringen, Germany (Jaeckel,
1962).
Remarks. See Helicigona (Drobacia) and Droba-
cia in Schileyko (2006, 2013) for details regarding the
shell and genital morphology. Drobacia has undivided
accessory glands (Fig. 4.9).
Genus Faustina Kobelt, 1904
Type species: Helix faustina Rossmässler, 1835
Molecular data. Faustina is shown monophyletic in
all our phylogeny reconstructions (PP ≥ 0.99), but
CytB and 16S sequences were only obtained for sub-
species of F. faustina. Consequently the monophyly of
the genus could only be assessed with the data for H3
and COI. Sequence divergences within Faustina are
generally large; between F. faustina orba (von Kima-
kowicz, 1890) and F. kiralikoeica (von Kimakowicz,
1890) the sequence divergence for COI is 17.4%. Even
between the alleged subspecies F. f. faustina (Ross-
mässler, 1835) and F. f. associata (Rossmässler, 1835)
divergences reach up to 10.7% (Table 4). A sister-group
relationship between Faustina and Kosicia is shown
with the phylogeny reconstructions for the concatenated
datasets (PP ≥ 0.95; Figs 1, 2, S5, S6). The phylogeny
for H3 indicates Faustina as the sister-group of all
other Ariantinae, but this is not supported by any of
the other phylogeny reconstructions.
Age. The most recent common ancestor of the in-
vestigated Faustina specimens is estimated at ca. 13.4 -
11.3 MYA. The split between Faustina and Kosicia is
estimated at 56-51.7 MYA (Figs 2, S5).
Distribution. The Carpathian Mts, E Czech, Slova-
kia, S Poland, W Ukraine and Romania; also in NE
Hu ng a r y. Faustina faustina (Fig. 3.12) has the widest
distribution, F. rossmaessleri (Pfeifer, 1848) and F.
cingulella (Rossmässler, 1837) are mainly found in
Slovakia, F. barcensis (von Kimakowicz, 1890) and F.
kiralikoeica are found in Romania.
Remarks. There are at least 5 Faustina species, 3 of
which are included in this study, viz. the nominate sub-
species of F. faustina, two additional subspecies [F.
faustina associata and F. faustina orba], and F. kira-
likoeica. See Campylaea (Faustina) and Faustina in
Schileyko (2006, 2013) for details regarding the shell
and genital morphology. In Faustina both types of ac-
cessory glands occur. Faustina cingulella and F. ross-
maessleri have undivided glands, whereas they are
split up to half their length in F. faustina (Fig. 4.19), F.
barcensis and F. kiralikoeica.
Genus Helicigona Férussac, 1821
Type species: Helix lapicida Linnaeus, 1758
Molecular data. In the past this generic name has been
used for many taxa of the Ariantinae (Hesse, 1931;
Knipper, 1939; Zilch, 1960; Subai, 1984). None of our
phylogeny reconstructions support these views. The
phylogeny reconstructions of the concatenated data-
sets indicate that Causa and Isognomostoma together,
are most likely the sister-group of Helicigona (group
A; Figs 1, 2, S5, S6). The monophyly of the two alleged
subspecies of H. lapicida is beyond dispute (PP = 1.0;
Figs 1, 2, S1-S6). COI and CytB sequence divergences
within each subspecies are less than 2.5% (n = 4), but
between both subspecies they reach up to 12% and
14%, respectively.
Age. The only fossil that can be indisputably as-
signed to any of the currently recognized Ariantinae is
a representative of Helicigona (see Nordsieck, 2014
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Groenenberg et al. – Systematics of Ariantinae
Fig. 4. Genital anatomy for most of the currently recognized genera of Ariantinae.The simplied diagram of the genital morphology
was reproduced and adapted from Koene and Schulenburg (2005; Creative Commons Attribution License 2.0).
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59
Contributions to Zoology, 85 (1) – 2016
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Groenenberg et al. – Systematics of Ariantinae
and references therein) from the Late Burdigalian
(~17.5-16.0 MYA). This date was the only calibration
point used in our BEAST analyses. The split between
Helicigona and the lineage Causa-Isognomostoma is
estimated at ca. 62.6-61.2 MYA (Figs 2, S5).
Distribution. The nominate subspecies is widely
distributed in W and N Europe, from S Scandinavia
and central England to the south up to S France, to the
east up to Czech and W Poland. Helicigona lapicida
andorrica (Bourguignat, 1876) is restricted to the east-
ern Pyrenees.
Remarks. Helicigona is considered a monotypic ge-
nus with only two clearly differentiated subspecies,
viz. Helicigona l. lapicida (Fig. 3.25) and H. l. andor-
rica, which are both included in this study. See Heli-
cigona (Helicigona) and Chilostoma (Helicigona) in
Schileyko (2006, 2013) for details regarding the shell
and genital morphology. The accessory glands are un-
divided (Fig. 4.6).
Genus Isognomostoma Fitzinger, 1833 (monotypic)
Type species: Helix personata Lamarck, 1792 [= Isog-
nomostoma isognomostomos (Schröter, 1784)]
Molecular data. Isognomostoma isognomostomos
(Fig. 3.4) and Causa holosericea (Fig. 3.5) have long
been regarded as congeneric. All phylogeny recon-
structions, except the one based on H3 (Fig. S4), ex-
plicitly show Causa and Isognomostoma together as a
monophyletic group (PP ≥ 0.99; Figs 1, 2, S2-S6). See
also the paragraph on Causa.
Distribution. Mountains of central Europe, S of the
line Eifel, Sauerland and the Harz Mts. From E France
eastwards in Zwitserland, Austria, N Italy, Slovenia,
Croatia, Czech, Slovenia, S Poland (Carpathians), NE
Hungary and Rumania.
Remarks. See Isognomostoma in Schileyko (2006,
2013) for details regarding the shell and genital mor-
phology. The accessory glands are undivided (Fig.
4.8).
Genus Josephinella Haas, 1936
Type species: Helix hemonica Thiesse, 1884 (Fig. 3.34)
Molecular data. Based on 11 included species (2 un-
described; Table S7), Josephinella is considered a
monophyletic group (PP = 1.0; Figs 1, 2, S2-S6). The
phylogeny reconstructions for the combined datasets
show Thiessea as the sister-group of Josephinella (PP
= 1.0; Figs 1, 2, S5, S6). Josephinella reischuetzi
(Su
bai, 1990) and J. vikosensis (Subai, 1990) together,
which were once classied in Superba by Subai and
Fehér (2006) are shown to belong to Josephinella (PP
≥ 0.87; Figs S1, S2).
Age. The most recent common ancestor of Josephi-
nella (based on four taxa) is estimated at ca. 22-21.6
MYA (Figs S5, 2).
Distribution. Southern half of Albania, the SW bor-
der area of Macedonia (FYROM), the Ionian islands,
mainland Greece and the Peloponnese.
Remarks. With at least 18 named species, and more
than 10 still to be described (Subai, in prep.), Josephi-
nella is the most speciose genus of the Ariantinae. For
this study 11 species were included.
See Helicigona (Josephinella) and Chilostoma (Jo-
sephinella) in Schileyko (2006, 2013) for details re-
garding the shell and genital morphology. The acces-
sory glands are generally split from halfway up to 2/3
of their length (Fig. 4.22); specimens with one or both
glands undivided (J. vikosensis and J. reischuetzi; Fig.
4.26 and Fig. 4.18) or trifurcate accessory glands are
rare (n=60: 4 undivided, 56 split, of which 3 specimens
had only one divided gland).
Genus nov. Kollarix
Type species: Helix kollari Pfeiffer, 1856 (monotypic)
Diagnosis. The diagnosis of this monotypic subgenus
is by denition identical with that of its type species,
i.e. Kollarix kollari (Pfeiffer, 1856). Kollarix gen. nov.
is also differentiated by a unique combination of nu-
cleotides in the 655 bp COI sequence obtained with
general barcoding primers (Folmer, 1994) at the fol-
lowing relative positions: 16 C, 67 A, 68 A, 84 G, 94 T,
357 G, 475 A, 493 G, 556 G, 625 C.
Description. Shell strongly depressed, nearly dis-
coid, with 4¾-5¼ whorls; umbilicus wide, measuring
1/5-1/6 of the total shell width; corneous brown, with a
brown spiral band in a whitish zone (see Welter-Schul-
tes, 2012: 595, Helicigona kollari); surface nely gran-
ulated, with growth lines and hairs. Height 8.5-12.0
mm; width 18.5-27.0 mm. The accessory glands are
always undivided (Fig. 4.15). Stimulator broad, at-
tened, lling the genital atrium and reaching far into
the vagina; an extension ends at the insertion of the
penis. Penial papilla small, conical, with ne trans-
verse ridges and an obtuse apex with a short, trans-
verse, slit-like pore. Secondary ureter closed for 0.5-
1.0 mm and open for the remaining 2-3 cm.
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Contributions to Zoology, 85 (1) – 2016
Molecular data. Genet i c a l l y, Kollarix kollari can-
not be assigned to Liburnica (Subai, 2002), nor any
other of the described genera. Kollarix is shown to be
a separate lineage within the Ariantinae, which is
more closely related to Pseudotrizona gen. nov., than
to Liburnica Kobelt, 1904 (Figs 1, 2, S5, S6).
Age. The most recent common ancestor of Kollarix
and the clade (Pseudotrizona - Cattania - Josephinella
- Thiessea) is estimated at ca. 55.9-55.8 MYA (Figs 2,
S5).
Distribution. Endemic to Serbia, S of the Donau up
to Aleksinac, between Šabac and Bor districts.
Remarks. Helix kollari Pfeiffer, 1856 (Fig. 3.19) has
been classied in Campylaea (by Tomić, 1959), in
Helicigona (by Knipper, 1939; Maassen, 1985) and
most recently in Liburnica (by Subai, 2002; Bank,
2012). In his treatise on Liburnica, Subai (2002) hy-
pothesized that Kollarix kollari might be the oldest
representative of Liburnica. Since Kollarix kollari
cannot be assigned to any of the described genera, nei-
ther genetically nor morphologically, it is here given
generic status. The name Kollarix has been used by
Groenenberg et al. (2012) and Schileyko (2013: 146),
but in both cases with the explicit note that this was not
for purposes of zoological nomenclature.
Derivatio nominis. Kollarix refers to kollari.
Genus Kosicia Brusina, 1904
Type species: Helix intermedia Pfeiffer, 1828
Molecular data. Kosicia is usually regarded as a sub-
genus of Chilostoma (Zilch, 1960; Bank et al., 2001),
but should be given generic status based on our phy-
logeny reconstructions. Its three species, viz. Kosicia
ambrosi (Strobel, 1852) (Fig. 3.15), K. intermedia (Fig.
3.16) and K. ziegleri (Rossmässler, 1836) (Fig. 3.17)
form a monophyletic group (PP ≥ 0.94; Figs 1, 2, S1-
S6). Kosicia ambrosi, which is much smaller than the
other two species, is the sister-group of K. intermedia
and K. ziegleri together (Figs 1, 2, S2, S3, S5, S6). The
phylogeny reconstructions for the concatenated data-
sets show Faustina as the sister-group of Kosicia (PP
= 1.0; Figs 1, 2, S5 and S6).
Age. The most recent common ancestor of Kosicia
is estimated at ca. 30.1-28.0 MYA (Figs 2, S5); that of
K. intermedia and K. ziegleri is estimated at ca. 7
MYA (Figs 2, S5).
Distribution. Kosicia intermedia is most widely
distributed; it occurs in NE Italy, S Austria (Kärnten),
NE Italy, Slovenia and NW Croatia. Kosicia ambrosi
has the smallest range; it is endemic to E Trentino and
the Prealps of Veneto (Italy). Kosicia ziegleri occurs in
S Kärnten (Austria) and in the border area between
Italy and Slovenia.
Remarks. All three known Kosicia species were
in
cluded in this study. See Helicigona (Kosicia) and
Kosicia in Schileyko (2006, 2013) for details regarding
the shell and genital morphology. The accessory
glands are always undivided (Fig. 4.13).
Subgenus Liburnica Kobelt, 1904, genus Liburnica
Type species: Helix setosa Férussac, 1832 (Fig. 3.20)
Molecular data. Liburnica has been regarded a subge-
nus of Campylaea (Zilch, 1960) and Chilostoma by
Bank et al. (2001), but none of our phylogeny recon-
structions indicate a close relationship between any of
these taxa. Liburnica respresents a distinct, monophy-
letic lineage (PP = 1.0; Figs 1, 2 and S1-S6), which in-
cludes Superba (Subai and Fehér, 2006) according to
H3 and COI sequences (Figs S1, S2; see Superba). The
position of Liburnica is only partly resolved; our data
hint at a sister-group relation with Dinarica (0.44 ≤ PP
≤ 0.78; Figs 2, S5, S6; see Dinarica). Only the phylo-
geny based on COI explicitly supports another sister-
group relation; see Drobacia. Six species of Liburnica
(Liburnica) could be investigated. The subgenus is
shown as a monophyletic group in the phylogeny based
on COI (PP = 0.75; Fig. S2). COI sequence divergences
within L. (Liburnica) range from 6.3% to 3.4%. Be-
tween L. (Liburnica) and L. (Superba) the COI se-
quence divergence is ≤ 6.9% (Table 4).
Age. The most recent common ancestor of Libur-
nica and Dinarica is estimated at ca. 53.2-51.4 MYA;
that of Liburnica (Liburnica) is estimated at ca. 5.9-
3.1 MYA (Figs 2, S5).
Distribution. Mts. along the NE coast of the Adri-
atic Sea in Croatia, Bosnia-Herzegowina, Montenegro,
Kosovo, Albania, W Macedonia, southwards to Epirus
in NW Greece.
Remarks. With over 15 described species, Liburnica
is among the most speciose genera of Ariantinae. Con-
chologically Liburnica is quite var iable (Fig. 3.20, 3.23).
The 6 species used in this study (10 including Super-
ba), suggest that these forms radiated rapidly (< ~ 6
MYA;
see Age). See Subai (2002) and Schileyko
(2013) for details regarding the shell and genital mor-
phology. The upper 1/3 to 2/3 of the accessory glands
in L. (Liburnica) are generally split (Fig. 4.20); occa-
sionally one (Fig. 4.17) or both glands are undivided.
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62
Groenenberg et al. – Systematics of Ariantinae
Subgenus nov. Oricampylaea, genus Campylaea
Type species: Faustina (Campylaea) illyrica Stabile,
186 4
Diagnosis. Oricampylaea subgen. nov. is differentiat-
ed by a unique combination of nucleotides in the 655
bp COI sequence obtained with general barcoding
primers (Folmer, 1994) at the following relative posi-
tions: 88 T, 187 T, 220 T, 301 T, 385 A, 409 C, 556 T,
575 T, 578 C, 650 C.
Molecular data. This clade (PP = 1.0; Fig. S2) con-
sists of at least the species Campylaea (Oricampy-
laea) illyrica and C. (Oricampylaea) lefeburiana (Fé-
russac, 1821). After our H3 sequence, Helicigona (Ari-
anta) ljubetenensis Wagner, 1914 (in Sturany and
Wagner, 1914), which was regarded as a subspecies of
Cattania (C.) trizona by Knipper (1939) and Bank
(2012), has to be added as a third species.
Age. Based on the intraspecic divergence in C.
(Oricampylaea) illyrica, the most recent common an-
cestor of C. (Oricampylaea) is estimated at, at least,
ca. 19-17.9 MYA (Figs 2, S5).
Distribution. Southern Germany (introduced),
southern Austria, from Slovenia southwards to W and
N Croatia. Campylaea (Oricampylaea) illyrica also
occurs along the SW Hungarian border, in N Serbia
and in SW Romania. Campylaea (Oricampylaea)
lju
betenensis is restricted to the Šar Mts (between Ko-
sovo and NW Macedonia).
Remarks. The phylogeny reconstructions based on
H3 and COI show a clade within Campylaea that sepa-
rates C. (Oricampylaea) illyrica and C. (Oricampy-
laea) lefeburiana from C. (Campylaea) planospira.
Here we denoted this group Oricampylaea subgen.
nov., because this clade persists even in case the un-
timely inclusion of C. (Oricampylaea) ljubetenensis
would turn out to be incorrect. Observing C. (Ori-
campylaea) ljubetenensis in a clade (data for H3 on ly)
with C. (Oricampylaea) illyrica is surprising both
morphologically as well as geographically; in shell
shape C. (Oricampylaea) ljubetenensis resembles C.
(Cattania) trizona more than C. (Oricampylaea) il-
lyrica, whereas it occurs ca. 250 km south of the distri-
bution area of the latter species. Future research has to
show if the provisional assignment of C. ljubetenensis
to C. (Oricampylaea) will uphold and whet her Campy-
laea hirta ( Me n ke, 183 0), C. m acrostoma (Rossmässler,
1836), C. schlaerotricha (Bourguignat, 1870), and C.
sadleriana (Rossmässler, 1838) should be assigned to
this new subgenus as well. The accessory glands for C.
(Oricampylaea) lefeburiana and C. (Oricampylaea)
ljubetenensis (Fig. 4.25) are split, whereas those for C.
(Oricampylaea) illyrica are undivided (Knipper,
1939). The name Ljubotenia has been used for C. (Ori-
campylaea) ljubetenensis by Groenenberg et al. (2012)
and Schileyko (2013: 146), but in both cases with the
explicit note that this was not for purposes of zoologi-
cal nomenclature.
Derivatio nominis. The epithet Oricampylaea is used
for a group of oriental Campylaea species, which can-
not yet be diagnosed with morphological characters.
Genus nov. Pseudotrizona
Type species: Helix inflata Kobelt, 1876 (monotypic)
Diagnosis. The diagnosis of this monotypic subgenus
is by denition identical with that of its type species,
i.e. Pseudotrizona inflata (Kobelt, 1876). Shell light
corneous with three brown spiral bands and a narrow
umbilicus. Pseudotrizona gen. nov. is also differenti-
ated by a unique combination of nucleotides in the 655
bp COI sequence obtained with general barcoding
primers (Folmer, 1994) at the following relative posi-
tions: 22 A, 181 A, 265 G, 271 G, 304 A, 325 A, 413 C,
481 G, 616 A, 649 C.
Description. Shell depressed conical, whitish to
light corneous, with three brown spiral bands (Welter-
Schultes, 2012: 594, Helicigona inflata). Surface with
growthlines only. With 4¾-5½ whorls; umbilicus nar-
row, measuring c. 1/10 of the total shell width. Height
10.5-18.0 mm; width 20.3-31.5 mm.
The accessory glands (Fig. 4.28) may be split for 1/3
to 1/2 of their length, but occasionally specimens with
both an undivided and a split glandula occur as well.
Stimulator more or less rounded triangular, promi-
nently protruding obliquely in the central part of the
genital atrium. Penial papilla slender conical, some-
times narrowed in the middle, with ne transverse
ridges. Secondary ureter closed for 0.1-0.15 mm and
open for the remaining 3.5-4.0 cm.
Molecular data. In the phylogeny reconstructions
based on the concatenated datasets this species is al-
ways the sister-group of the clade Cattania-Josephi-
nella-Thiessea (PP = 1.0; Figs 1, 2, S5, S6). None of
the phylogenies show a species group exclusively con-
sisting of Pseudotrizona and Cattania, thus Pseudotri-
zona inflata is not a species of Cattania.
Age. The lineage that gave rise to Pseudotrizona is
estimated to have diverged from the common ancestor
of Cattania-Josephinella-Thiessea at ca. 53.4-48.8
MYA (Figs 2, S5).
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63
Contributions to Zoology, 85 (1) – 2016
Distribution. N Albania, Montenegro, Kosovo, SW
Serbia.
Remarks. Pseudotrizona inflata (Kobelt, 1876) has
long been considered a subspecies of Cattania trizona,
which was classied in Campylaea by Sturany and
Wagner (1914), and in Helicigona by Knipper (1939)
and Subai (1995).
Derivatio nominis. The epithet Pseudotrizona re-
fers to the former incorrect classication of the type
species as a subspecies of Cattania (C.) trizona.
Subgenus Sabljaria Brusina, 1904 (monotypic), genus
Dinarica
Type species: Helix stenomphala Menke, 1830
Molecular data. All phylogenies based on the con-
catenated datasets depict Dinarica (Sabljaria) and
D. (Dinarica) as a monophyletic group, see Dinarica
(genus-level sister-group relations are also discussed).
The subgenera D. (Sabljaria) and D. (Dinarica) are
genetically clearly different. The COI and CytB
sequence divergences between D. (Sabljaria) and
D. (Dinarica) are 16.3% and 23.8%, respectively
(Table 4).
Age. See D. (Dinarica) for the estimated age of the
genus.
Distribution. Endemic to the Velebit Mts along the
coast of Croatia.
Remarks. Dinarica (Sabljaria) differs from D. (Di-
narica) both conchologically (Fig. 3.21, 3.22) and in
genital anatomy (Fig. 4.23, 4.24). These subgenera are
allopatrically distributed. See Chilostoma (Sabljaria)
in Schileyko (2013) for details regarding the shell and
genital morphology. Dinarica (Sabljaria) has split ac-
cessory glands (Fig. 4.24).
Subgenus Superba Subai and Fehér, 2006, genus Li-
burnica
Type species: Helicigona skipetaricus [sic] Subai,
1995
Molecular data. No CytB or 16S sequences were ob-
tained for L. (Superba) and H3 does not discriminate
between the alleged subgenera of Liburnica (Fig. S1).
Therefore the taxonomic status of L. (Superba) could
only be assessed with COI. The phylogeny based on
that marker shows L. (Liburnica) as a monophyletic
group (n = 4) and L. (Superba) as paraphyletic (n = 4).
Which of these subgenera is monophyletic depends on
the selected outgroup. A phylogeny in which both are
monophyletic, was not obtained. Partly based on these
results, Subai (2012) synonymized Superba with Li-
burnica. The COI sequence divergences within L. (Su-
perba) are less than 2.1% (Table 4).
Age. Due to missing data, L. (Superba) was not in-
cluded in the BEAST analyses. Given the limited
amount of sequence divergence within Liburnica (and
the larger intraspecic divergence in L. (Liburnica);
Table 4), we expect L. (Superba) not to be older than
L. (Liburnica); see L. (Liburnica).
Distribution. Albania, Tomor and Kulmakës Mts.
Remarks. Liburnica (Superba) contains three spe-
cies, viz. L . (S.) skipetarica (Subai, 1995) (Fig. 3.23),
L. (S.) grisea (Subai and Fehér, 2006) and L. (S.) kul-
mankana (Subai and Fehér, 2006), which are all in-
cluded in this study (for remarks on J. reischuetzi and
J. vikosensis; see sub Josephinella). See Subai and
Fehér (2006) for details regarding the shell and genital
morphology. The accessory glands are generally split
(Fig. 4.21), one gland undivided is also observed (Fig.
4.17 ).
Genus Thiessea Kobelt, 1904
Type species: Helix cyclolabris Deshayes, 1839 (in
Férussac and Deshayes, 1819-1851)
Molecular data. Thiessea is generally considered a
subgenus of Chilostoma (Zilch, 1960; Bank et al.,
2001). This view cannot be accepted, since both taxa
are not shown to be closely related in any of our phy-
logeny reconstructions. The data obtained for Thiessea
are limited; for three out of the four included species,
only H3 sequences were obtained. The H3 phylogeny
indicates the four Thiessea species as a monophyletic
group (PP = 0.49; Fig. S1). A sister-group relation is
shown between Thiessea and Josephinella (PP = 1.0;
Figs 1, 2, S1, S5, S6).
Age. The most recent common ancestor of Thiessea
and Josephinella is estimated at ca. 39-36.6 MYA
(Figs 2, S5).
Distribution. Mainland SE Greece, NE Pelopon-
nese, Aegean Islands and SW Turkey.
Remarks. With at least 16 species (of which only 4
included in this study), Thiessea