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Journal of Conchology
V. 45, . 1
Published by the Conchological Society of Great Britain and Ireland, established 1874
ISSN 2755-3531
Revision of the genera of Scolodontidae, part 1:
disentangling Happia Bourguignat, 1890 from Austroselenites Kobelt, 1905,
Drepanostomella Bourguignat, 1890, Hirtudiscus Hylton Sco, 1973,
Luteostriatella gen. nov., and Systrophiella H.B. Baker, 1925
M T. R1 & A S.H. B2, 3, 4
1 Natural History Museum of Roerdam, Westzeedijk 345, NL-3015AA Roerdam, e Netherlands; hps://orcid.org/0000-
0001-7651- 6685
2 Natural History Museum, Invertebrate Division, London SW7 5BD London, U.K.
3 Royal Belgian Institute of Natural History, Vautierstraat 29, B-1000 Brussels, Belgium
4 Naturalis Biodiversity Center, P.O. Box 9517, NL-2300 Leiden, e Netherlands
Corresponding author: M.T. Roosen (marijn.roosen@gmail.com)
Abstract. Currently there are several problems with the genus-level taxa within the Scolodontidae. Multiple type-spe-
cies designations have been proposed for some genera, type species are poorly described, and in some cases no clear
decisions have been made in cases of homonymy or synonymy. is has resulted in wrongly identied species and
genera within this group, which, among other problems, hinders the discovery and description of new species as
well as the identication of known species. is paper is the rst in a series in which all scolodontid genera will be
redescribed based on type materials, starting with Happia Bourguignat, 1890 and its allies. Nomenclatural issues are
resolved where possible. One new genus and a new species are described: Luteostriatella gen. nov. and Austroselenites
pichinchense sp. nov. e following new combinations are made: Happia andia (Pilsbry, 1932) comb. nov., Systro-
phiella altivaga (Crawford, 1939) comb. nov., Systrophiella cayennensis (L. Pfeier, 1842) comb. nov., Systrophiella
pygmea (Spix in Spix & Wagner, 1827) comb. nov., Systrophiella snethlagei (F. Baker, 1913) comb. nov., Systrophiella
vitrina (J.A. Wagner in Spix & Wagner, 1827) comb. nov., and Luteostriatella variegata (F. Haas, 1949) comb. nov.
Key words. Neotropics, South America, Mollusca, Gastropoda, Eupulmonata, Scolodontina
ZooBank identier. urn:lsid:zoobank.org:pub:F7C305E8-616F-4F5C-9D70-24A4561335DB
DOI. hps://doi.org/10.61733/jconch/4511
I
Scolodontidae are a species-rich but understudied family
of land snails (Ramírez 1993). Recent genetic studies sug-
gest that the family forms its own suborder (Scolodontina)
within the Stylommatophora (Bouchet et al. 2017; Saadi
& Wade 2019). Scolodontids occur only in the Neotrop-
ical region and are mostly represented by small species
(Ramírez 1993; Barbosa 2014). Many of these small species
remain unknown to science, as shown by Wendebourg &
Hausdorf (2019) and Ramirez Perez & Hausdorf (2022),
who discovered many new species in the rainforests of the
Peruvian Amazon and Ecuadorian Chocó, but most of these
reported species could not be identied below the family
level. A similar high diversity of unknown Scolodontidae is
also recognised elsewhere in Ecuador (Roosen 2019).
Describing these taxa is currently hindered by two issues.
Most taxa are only known from their shells, which show few
characteristics for comparison, and the genera are poorly
dened and their limits oen seem confusing and overlap-
ping. A good example of this is the recently published case of
Hirtudiscus excisa (Reeve, 1854), which has been only gured
by Reeve (1854) and Breure et al. (2022) and was used to
dene Drepanostomella Bourguignat, 1890 by Baker (1925a),
Ramirez (1993), Hausdorf (2003), and Breure et al. (2022),
even though it is much smaller and has a dierent sculpture
than the type species of Drepanostomella (see Roosen 2023).
e most problematic genus is probably Happia Bour-
guignat, 1890, which is currently used for 42 species (Mol-
luscaBase 2023a) of morphologically distinct scolodontids.
17 A 2024
R & B: Revision of the genera of Scolodontidae, 1 92
Happia was introduced by Bourguignat (1890) as a replace-
ment name for Ammonoceras L. Pfeier, 1855 (non Lamarck,
1822). e type species of Ammonoceras is Helix ammon-
oeras Reeve, 1854, designated by absolute tautonymy in
accordance with International Code of Zoological Nomen-
clature Article 68.4 (ICZN 2023). However, Bourguignat
(1890) placed this species in Drepanostomella. Gude (1902)
either overlooked this error or followed Pfeier & Clessin
(1881) and designated Helix vitrina as the type species of
Happia, an interpretation later adopted by many authors
(e.g. Baker 1925a). To complicate maers, Baker (1925a)
used Helix ammonoceras along with Hirtudicus excisa to
establish the genus diagnosis of Drepanostomella (Roosen
2023) and Helix euspira Reeve, 1854, the type species of
Austroselenites Kobelt, 1905, is currently included in Hap-
pia (e.g. Simone 2006; Agudo-Pádron 2023; MolluscaBase
2023b). Austroselenites has been placed in the family Haplo-
trematidae H.B. Baker, 1925 since the publication of Baker
(1941).
ese unresolved issues within scolodontid taxonomy
hinder proper research of both known and unknown taxa.
is family needs an urgent revision, especially at the genus
level. Proper identication at the genus level is currently
nearly impossible, and placement of some taxa within the
Scolodontidae is doubtful at best. is is the rst paper of a
series we have in preparation that aims to properly diagnose
scolodontid genera based on their type species and other
relevant species. ese papers are organized in a manner
that facilitates disentangling the most prominent nomen-
clatural issues in this group and do not reect currently
accepted taxonomic order.
M M
To revise the known scolodontid genera, we undertook a
comprehensive study of all literature available to us con-
taining information on these genera, the type specimens of
the type species, and most species from Ecuador and Peru.
e laer were already located in museum collections and
imaged for unrelated projects and for that reason readily
available for the current study (Breure et al. 2022). Shell
characteristics useful for identication are highlighted and
a new species relevant for the diagnosis of the genera is pre-
sented in this paper.
Specimens were imaged by scanning electron microscopy
(SEM) and light microscopy under stereomicroscopes at
the institutes where the specimens are deposited at or at the
Royal Belgian Institute of Natural Sciences. Measurements
were taken to the nearest 0.1 mm during the imaging process
or with vernier callipers. e shell height (H) was measured
from the apex to the lower lip of the aperture, width (W) at
the widest section perpendicular to the coiling axis, height of
the aperture (HA) from the lowest point of the peristome to
the upper part of the whorl, and the umbilical width (UW)
at the widest section starting at the columella. All measure-
ments are in millimetres. Whorls were counted to the nearest
¼ whorl following Gienberger et al. (2004). Other abbre-
viations used adapted from original descriptions are largest
diameter (Diam. Maj.; D), smallest diameter (Diam. Min.;
D3), height (alt., long.), and width (lat.).
For many genera only the shells of the type species are
known. erefore, we only use conchological characteris-
tics of the type specimens to dene the genera. DNA and
anatomy should be studied, but this is beyond the scope of
this paper. We have analysed type specimens or clear pho-
tographs of type specimens for all species treated in this
paper. Taxa that we did not study should be treated as ten-
tative members of the genera in which they are currently
placed, with exception of Happia. e species of Happia not
examined in this study are tentatively moved to Systrophiella
unless indicated otherwise.
Specimens from the following museum collections were
studied: Academy of Natural Sciences of Drexel University,
Philadelphia, Pennsylvania, USA (ANSP); Field Museum
of Natural History, Chicago, Illinois, USA (FMNH);
Museo de La Plata, La Plata, Argentina (MLP); Muséum
National d’Histoire Naturelle, Paris, France (MNHN), Nat-
ural History Museum, London, United Kingdom (NHM,
NHMUK); Ponticia Universidad Católica del Ecuador,
Quito, Ecuador (PUCE, QCAZI); Zoologische Staatssam-
mlung München, Munich, Germany (ZSM, SNSB). Other
institutes mentioned: Leibniz Institute for the Analysis of
Biodiversity Change, Hamburg, Germany (LIB); Royal
Belgian Institute of Natural Sciences, Brussels, Belgium
(RBINS).
S
Family Scolodontidae H.B. Baker, 1925
Scolodontidae Baker 1925b: 88.
In this paper we discuss six scolodontid genera with com-
plex nomenclatural histories. To avoid further confusion,
we list the most important conchological characteristics to
distinguish the genera in Table 1.
Genus Happia Bourguignat, 1890
Ammonoceras L. Pfeier 1855: 122; junior homonym of Am-
monoceras Lamarck, 1822.
R & B: Revision of the genera of Scolodontidae, 1 93
Table 1. Key dierences between the scolodontid genera discussed in this paper. e genera are ordered in accordance with their appear-
ance in the main text of the paper.
Genus
Parietal
incision
Maximum
diameter (mm) Sculpture
Happia Bourguignat, 1890 Yes 2.5–4.5 Spiral rows of papillae or striae
Drepanostomella Bourguignat, 1890 Yes 25–30 Absent or vestigial spiral sculpture
Hirtudiscus Hylton Sco, 1973 Yes 4–6 Protoconch with spiral lirae, teleoconch with periostracal hairs
and wavy axial ribs
Systrophiella F. Baker, 1925 No <18 Absent or vestigial spiral sculpture
Luteostriatella gen. nov. No <10 Absent, but periostracum with bands of colour
Austroselenites Kobelt, 1905 No 25–35 Fine axial ribbing
Happia Bourguignat 1890: 39. Type species: Helix ammonoc-
eras Reeve, 1854 (typication of the replaced name).
Diagnosis. Shell small, discoid to subdiscoid, transpar-
ent-whitish. First whorl without sculpture. Second, third,
and fourth whorls with weak growth striae and spiral rows
of papillae or microscopic spiral grooves. Umbilicus wide.
Aperture broadly lunulate. Peristome simple, with an acute
incision in parietal angle.
Species included in the genus. Happia ammonoceras (Reeve,
1854), Happia andia (Pilsbry, 1932) comb. nov.
Species tentatively included in the genus. ?Happia lyzarz-
aburui ( Jousseaume, 1887).
Geographic range. Colombia and Peru. Questionably Ec-
uador.
Comparisons. In size and general shape true Happia is most
similar to Hirtudiscus Hylton Sco, 1973 and Xenodiscula
Pilsbry, 1919. However, they do not have a punctate or stri-
ate microsculpture, which is typical of Happia. Moreover,
Xenodiscula does not have an acute incision in the parietal
angle. Drepanostomella Bourguignat, 1890 is also a group of
discoid scolodontids with an incision in the parietal angle
of the aperture, but this genus has no punctate or striate
microsculpture and the adult shells are much larger. For
more information on Xenodiscula, we refer to Pilsbry (1919)
and Roosen et al. (2023).
In this study, only species for which the spiral rows of
papillae (Fig. 1A–E) or microscopic spiral grooves are con-
rmed by detailed SEM photomicrographs are included
in Happia, as we consider this type of spirally oriented
microsculpture typical for the genus. Within the Scolodon-
tidae only Miradiscops H.B. Baker, 1925 and its subgenera
have similar pied or striate microsculpture, which was
especially well documented by Ravalo et al. (2023). How-
ever, Happia diers from Miradiscops by its sunken spire,
less convex whorls, and acute incision in parietal angle of
the peristome. e group of Miradiscops will be discussed in
a subsequent paper.
Remarks. e confusion about the identity of the type spe-
cies of Happia pre-dates the replacement of Ammonoceras
L. Pfeier, 1855 (non Lamarck, 1822) by Happia in Bour-
guignat (1890). No clear type designation was given in any
of Pfeier’s papers, but according to ICZN Article 68.4,
Helix ammonoceras Reeve, 1854 should be regarded as the
type species by absolute tautonymy. As several authors (e.g.
Baker 1925a; Breure et al. 2022) have acknowledged this
species as the type of Ammonoceras (and by extension Hap-
pia) and it is the oldest known type designation, it cannot
be replaced by a later designation, as stated in ICZN Article
70.2. As there is confusion around the identity of Happia
(see below), the decision to go back to the original type spe-
cies is needed to stabilise Happia.
So where did the original denition of Happia get lost?
As early as 1863, omas Bland considered Helix eu spira L.
Pfeifer, 1854the type species of Austroselenites Kobelt,
1905to be the type species of Ammonoceras (= Happia)
(Bland 1863). In a paper published aer Pfeier’s death
(Clessin & Pfeier 1881), Helix vitrina J.A. Wagner in Spix
& Wagner, 1827 is the proposed type species of Ammonoc-
eras. Aer this, Tryon (1885) further complicated maers
by regarding Helix ammoniformis d’Orbigny, 1835the
type species of Drepanostomella Bourguignat, 1890as
the type species of Ammonoceras. Bourgui gnat (1890)
later replaced Ammonoceras with Happia, but moved its
type species, Helix ammonoceras, to Scolodonta Doering,
1874, and Gude (1902) designated Helix vitrina J.A . Wag-
ner in Spix & Wagner, 1827 as the type species of Happia.
Baker (1925a) recognised the problems with Gude’s type
designation, but he followed it regardless and dened Dre-
panostomella using H. ammonoceras along with Helix excisa
Reeve, 1854, which is a species of Hirtudiscus Hylton Sco,
1973 according to Roosen (2023). Baker’s views were later
R & B: Revision of the genera of Scolodontidae, 1 94
adopted by Ramírez (1993). Because of all of the above,
several scolodontid genera became entangled with Hap-
pia, which resulted in all those species, including type spe-
cies of other genera, being lumped together in Happia.
Happia ammonoceras (Reeve, 1854)
Figure 1
Helix ammonoceras Reeve 1854: pl. 191 g. 1338; Pfeier
1855a: 54; Pfeier 1859: 104.
Figure 1. Happia ammonoceras (Reeve, 1854). A–E, NHMUK 20210342, syntype. B–E apical surface of same shell.
R & B: Revision of the genera of Scolodontidae, 1 95
Helix (Ammonoceras) ammonoceras PfeierPfeier 1855b:
122.
Streptaxis (Ammonoceras) ammonoceras (Pfeier)Pfeier &
Clessin 1881: 15.; Tryon 1885: 65, pl. 13, g. 52.
Scolodonta ammonoceras (Pfeier)Bourguignat 1890: 41.
Happia ammonoceras (Pfeier)Gude 1902: 235.
Drepanostomella ammonoceras (Pfeier)?Baker 1925: 23.;
Ramírez 1993: 21.
Misidentication:
Streptaxis (Ammonoceras) ammonoceras (Pfeier)Smith 1895:
315 (= unknown species).
Studied material. NHMUK 20210342 (syntypes, 3 shells,
dry), “New Grenada”.
Type locality. “Santa Anna, New Grenada”.
Measurements. “Diam. maj. 4½, min. 3¾, alt. vix 1½ mill.”
(Pfeier 1855a: 55).
Redescription. Shell medium-sized, discoid, translucent,
with a sunken spire. Embryonic whorl smooth. Aer one
whorl, eight slightly oblique spiral rows consisting of
densely spaced punctuations develop on last quarter of shell
between suture and periphery. Spiral rows visible on entire
shell and continue over periphery and onto base, only to dis-
appear on last quarter whorl in umbilicus. Flexuous growth
striae also present on entire shell but stronger towards aper-
ture. Aperture deected, rounded-trapezoidal in shape, with
a small incision at suture. Peristome simple. Umbilicus wide,
up to 32% of shell width.
Geographic range. Colombia.
Comparisons. Only “Ammonoceras” lyzarzaburui Jous-
seaume, 1887 is similar in shape and microsculpture. is
species diers from H. ammonoceras by its smaller size,
less distinct microsculpture that is only visible in high-res-
olution SEM images, and the absence of an incision in
the upper le corner of the aperture. In some respects
“Ammonoceras” lyzarzaburui is more similar to Miradiscops,
but it diers from this genus by its smaller microsculpture
and completely sunken spire. A new genus might have to be
established for this species, but this requires further study
of the type material and freshly collected shells from the
“Un poco del Chocó” nature reserve (Roosen 2019). For
dierences with H. andia (Pilsbry, 1932) comb. nov., see
that species.
Remarks. Happia sensu stricto is gured here for the rst
time since Reeve (1854). is is also the rst time that spi-
ral rows of papillae are documented for a scolodontid genus
other than Miradiscops. It is unclear whether the record of
this species by Baker (1925b) belongs to this taxon. If it
does, its periostracum has spiral rows of papillae.
Happia andia (Pilsbry, 1932) comb. nov.
Figure 2
Drepanostomella andia Pilsbry 1932: 400, g. 5–5bBaker
1963: 239; Richardson 1989: 118; Ramírez 1993: 21.
Studied material. ANSP 159924 (holotype, one shell, dry),
“Leymebamba, Department of Amazonas, Peru, at 7,000 ”.
Type locality. “Leymebamba, Department of Amazonas,
Peru, at 7,000 ” (Pilsbry 1932).
Redescription. Shell minute, transparent-whitish, with a
slightly sunken spire. Protoconch 1¼ whorl, without sculp-
ture. Teleoconch sculpture starts with a patch of 16 small,
densely crowded axial ribs and numerous indistinct spiral
grooves. Spiral grooves cover entire teleoconch, but axial
ribs become less crowded and distance between them more
variable. Aperture ovate-drop-shaped, with thin, simple
peristome. Umbilicus wide, up to 45% of shell width.
Dimensions. “Height 1 mm, Diam. 2.45 mm, 3¼ whorls”
(Pilsbry 1932).
Dierential diagnosis. It is most similar to Happia ammo-
no ceras (Reeve, 1854), which diers from H. andia by its
sculpture consisting of spiral rows of papillae and larger size.
Distribution. Peru: Amazonas department.
Remarks. Although the spiral sculpture and small size were
documented by Pilsbry (1932), at that time these charac-
ters were not recognised as a signicant in distinguishing H.
andia from Drepanostomella.
Genus Drepanostomella Bourguignat, 1890
Drepanostomella Bourguignat 1890: 42. Type species Helix
ammoniformis d’Orbigny, 1835 by monotypy.
Diagnosis. Shell large for family, discoid to subdiscoid, with
concave to slightly elevated spire. Younger whorls partially
cover older ones. Protoconch smooth; teleoconch with only
growth lines. Aperture ovate, not deected. Peristome sim-
ple, with an acute incision in parietal angle.
Species included in the genus. Drepanostomella ammoni-
formis (d’Orbigny, 1835), D. tucma Hylton Sco, 1948.
Species tentatively included in the genus. ?Drepanosto-
mella pinchoti Pilsbry, 1930.
Species not evaluated. Drepanostomella circumscripta Hyl-
ton Sco, 1948, D. stolli (E. von Martens, 1892), D. uru-
guayana Hylton Sco, 1979.
Geographic range. Panama, Brazil, Bolivia, Uruguay, and
Argentina.
Comparisons. Species belonging to Happia have oen erro-
neously be assigned to Drepanostomella, probably because
of the invalid subsequent type designation of Helix ammoni-
R & B: Revision of the genera of Scolodontidae, 1 96
formis d’Orbigny, 1835 as type species of Ammonoceras L.
Pfeier, 1854 (= Happia Bourguignat, 1890) by Tryon
(1885) (see the discussion of Happia). However, Drepanos-
tomella diers from Happia by its larger size and the absence
of punctate or striate microsculpture on the protoconch
and/or teleoconch.
As shown by Roosen (2023), a species of Hirtudis-
cus Hylton Sco, 1973 was used to provide a diagnosis of
Drepanostomella in several major papers on Scolodontidae
(e.g. Baker 1925a). Because of this, Hirtudiscus might be
confused with Drepanostomella. is genus diers from Hir-
tudiscus by its much larger size, smooth protoconch and sim-
ple periostracum (e.g. Hausdorf 2003; Hausdorf & Medina
2006; Roosen 2023).
Remarks. Recently Drepanostoma nautiliforme Porro, 1836,
was reported from Ecuador as Drepanostomella nautiliforme
in a paper on iridescence in terrestrial gastropods (González
& Teruel 2022). However, this is likely a misidentication,
as Drepanostoma Porro, 1836 is an unrelated European
genus. We noticed shells of Guestieria cf. powisiana (L.
Pfeier, 1848) misidentied as Drepanostomella nautiliforme
(Porro, 1836) on Femorale.com, the website of a Brazilian
shell vendor showcasing shells for sale (Femorale 2023).
Perhaps González & Teruel (2022) and the Femorale web-
site used the same erroneous source to identify Guestieria
Crosse, 1872 species as Drepanostoma nautiliforme. If so,
this source was not found by us. at said, it could also be a
simple error based on a misinterpretation of the similarity of
Drepanostoma to Drepanostomella, the tendency of authors
to describe the shape Guestieria species as “nautiliform” (e.g.
Jousseaume 1887) and the comparison of Guestieria to D.
nautiliforme in the original description of Guestieria (Crosse
1872). e genus Guestieria will be discussed in a later paper.
Drepanostomella ammoniformis (d’Orbigny, 1835)
Figure 3
Helix (Helicella) ammoniformis d’Orbigny 1835: 5.
Helix ammoniformisFérussac & Deshayes 1850: pl. 69B, g.
1; Gray 1854: 11.
Ammonoceras ammoniformisDoering 1875: 441; Pfeier
1876: 41; Pfeier & Clessin 1881: 15.
Figure 2. Happia andia (Pilsbry, 1932). A–C, ANSP 159924, holotype. B, C, apical surface of same shell.
R & B: Revision of the genera of Scolodontidae, 1 97
Hyalinia ammoniformisClessin 1888: 166.
Drepanostomella ammonitiformisBourguignat 1890: 43 (in-
correct subsequent spelling).
Happia (Drepanostomella) ammoniformisHaas 1949: 243.
Drepanostomella ammoniformisRamirez 1993: 19; Roosen
2023: 108, g. 5
Studied material. NHMUK 1854.12.4.105 (syntypes, 5
shells, dry), Yungas Province, Bolivia; MNHN-IM-2000-
25753 (syntype, 1 shell, dry), Yungas Province, Bolivia.
Type locality. “provincia Yungasensi (republica Boliviana)”.
Measurements. “Long. 6 millim.; latit. 18 millim., amplit.
27 millim.” (d’Orbigny 1835).
Redescription. Shell discoid, rather large for the family (up
to 27 mm wide), with a sunken spire. Protoconch–teleo-
conch transition unclear. Microsculpture of minute growth
striae on teleoconch. Some specimens with vestigial spiral
sculpture on earlier whorls. Aperture ovate; peristome thin,
with an incision in upper le corner of aperture. Umbilicus
wide, up to 37% of shell width.
Geographic range. Bolivia and Argentina.
Comparisons. is species is most similar to Drepanos-
tomella tucma Hylton Sco, 1948, which is distinguished
from D. ammoniformis by its smaller size. Drepanostomella
tucma Hylton Sco, 1948 was recently revised by Cuezzo &
Miranda (2009).
Figure 3. Drepanostomella ammoniformis (d’Orbigny, 1835). A–C, NHMUK 1854.12.4.105, syntype. B, C, apical surface of same shell.
R & B: Revision of the genera of Scolodontidae, 1 98
Genus Hirtudiscus Hylton Sco, 1973
Hirtudiscus Hylton Sco 1973: 128. Type species Hirtudiscus
hirtus Hylton Sco, 1973 by original designation.
Redescription. Shell small, up to 6 mm wide, subdiscoid
to discoid, with a at or only slightly raised spire. Protoconch
oen with more or less distinct, scalloped spirals; teleoconch
sculpture of thin, wavy axial ribs. Periostracum brownish, with
thin, oen spirally arranged periostracal hairs. Aperture drop-
shaped to trapezoidal, with an acute incision in parietal angle.
Species included in the genus. Hirtudiscus antioquiensis
Hausdorf & Medina, 2006, H. boyacensis Hausdorf, 2003, H.
comatus Hausdorf, 2003, H. curei Hausdorf, 2003, H. excisa
(Reeve, 1854), H. hirtus Hylton Sco, 1973, H. triserialis
Hausdorf & Medina, 2006.
Geographic range. Colombia, Ecuador, and Peru.
Comparisons. Hirtudiscus can be distinguished from all
other Scolodontidae by its small shell size, waved spirals on
the protoconch and complex periostracal structures.
Remarks. Until recently Hirtudiscus excisa (Reeve, 1854)
was regarded as a typical species of Drepanostomella. is
was resolved by Roosen (2023), by redescribing the species
while comparing it to Hirtudiscus hirtus Hylton Sco, 1973
and Drepanostomella ammoniformis (d’Orbigny, 1835). For
the dierences between Hirtudiscus and Drepanostomella,
see the discussion of Drepanostomella.
e record from Peru is based on an unnamed species
reported by Wendebourg & Hausdorf (2019) from the
Panguana reserve in the Peruvian Amazon. Even though
the species is likely undescribed, this record is worth men-
tioning, as it presents a signicant range extension for the
genus.
Hirtudiscus hirtus Hylton Sco, 1973
Figure 4
Hirtudiscus hirtus Hylton Sco 1973: 129, g. 2Hausdorf
2003: 179, g. 4A, 5A, B, 6; Hausdorf & Medina 2006: 211;
Roosen 2023: g. 4.
Studied material. MLP 3965, holotype, “Monte Redondo,
Colombia oriental, entre Bogotá y Villavicencio, 3800 m
alt.” (images only).
Type locality. “Monte Redondo, Colombia oriental, entre
Bogotá y Villavicencio, 3800 m alt.” (Hylton Sco 1973;
Hausdorf 2003).
Measurements. “D: 3.9; D3: 3.1; H: 1.8 mm” (Hausdorf
2003).
Redescription. Shell small, nearly discoid, with a at spire.
Protoconch of 1¾ whorls, separated from teleoconch by
a distinct scar. Spiral striae distinct on protoconch. Teleo-
conch with dense growth striae and, on damaged base,
spirally arranged periostracal hairs. Body whorl rounded,
Figure 4. Hirtudiscus hirtus Hylton Sco, 1973. A, B, MLP 3965, holotype. B, SEM images of same shell, not at the same scale as A.
R & B: Revision of the genera of Scolodontidae, 1 99
aperture broadly lunulate. Peristome sharp, neither ex-
panded nor thickened, with an incision in parietal angle of
aperture near suture. Umbilicus wide, occupying about 42%
of shell width.
Geographic range. Colombia.
Comparisons. Hirtudiscus hirtus diers from other species
by its at spire, comparatively smaller axial ribbing and
comparatively shorter periostracal hairs. Hirtudiscus triseri-
alis Hausdorf & Medina, 2006 is somewhat similar in shape,
but H. hirtus has more and shorter periostracal hairs (H.
triserialis only has three spirally rows of periostracal hairs)
and has a smaller adult size. For an in-depth description and
comparison of all Hirtudiscus species, see Hausdorf (2003),
Hausdorf & Medina (2006), and Roosen (2023).
Remarks. Periostracal hairs seem to worn of on the type,
but they were present when Hylton Sco (1973) examined
the specimen and are visible on the original gures.
Genus Systrophiella H.B. Baker, 1925
Systrophiella H.B. Baker 1925a: 15, 31. Type species: Scolo-
donta (Systrophiella) eudiscus H.B. Baker, 1925 by original
designation.
Redescription. Shells small to medium-sized, subdiscoid to
discoid, with a at or only slightly elevated spire and thin,
yellow-brown to brown periostracum. Protoconch with-
out sculpture; teleoconch with some vestigial spiral sculp-
ture most oen present on the teleoconch. Whorls slowly
increase in diameter. e umbilicus wide to very wide, aper-
ture subcircular, peristome simple. e last section of the
body whorl can be slightly deected.
Species included in the genus. Systrophiella alicea (Guppy,
1871), S. altivaga (Crawford, 1939) comb. nov., S. cayennen-
sis (L. Pfeier, 1842) comb. nov., S. eudiscus (H.B. Baker,
1925), S. lobaterita (H.B. Baker, 1925) S. pygmea (Spix in
Spix & Wagner, 1827) comb. nov., S. snethlagei (F. Baker,
1913) comb. nov., S. starkei (H.B. Baker, 1925), S. viridis
(H.B. Baker, 1925), S. vitrina (J.A. Wagner in Spix & Wag-
ner, 1827) comb. nov., S. weyrauchi (F. Haas, 1951).
Geographic range. Venezuela, French Guinea, Brazil, and
Peru.
Comparisons. Conchologically Systrophiella is most similar
to Scolodonta, but it diers from this genus by its larger size
and presence of vestigial spiral sculpture on the teleoconch.
For more information on Scolodonta, see Hausdorf (2006).
Remarks. Based on current information, this genus cannot
be separated from the Helix vitrina group (= the dominant
denition of Happia since the incorrect subsequent desig-
nation of H. vitrina as type species by Gude (1902)). No
properly identied specimen of H. vitrina was available to us
for dissection and comparison to the anatomy of S. eudiscus
as reported by Baker (1925a).
Baker (1925a) regarded the presence of vestigial spiral
sculpture on the teleoconch as the distinguishing charac-
ter in separating shells of the H. vitrina group from Systro-
phiella. However, we found some vestigial spiral sculpture
on the teleoconch of the syntypes of Systrophiella eudiscus,
H. vitrina, and H. nana, so this character cannot be used to
separate these taxa at the generic level. erefore, we ten-
tatively include the H. vitrina group in Systrophiella, while
acknowledging they might be dierent. Pending further
study, all orphaned species previously included in Happia
based on their similarities with Systrophiella vitrina comb.
nov. are moved to Systrophiella, except for Prohappia besckei
(Dunker, 1847), which likely should be maintained in a sep-
arate genus (Roosen & Breure in prep.).
Baker (1925a) noted that Helix thomasi L. Pfeier, 1855
from Marmato (Colombia) and some allied species are
similar to Systrophiella, but they dier in their minute spiral
sculpture. We have seen images of the syntypes of H. thomasi
and tentatively agree with Baker that these shells require fur-
ther study. e syntypes of this species will be discussed in a
later paper, along with some striate shells currently included
in Miradiscops and poorly known scolodontids described by
Preston (1914).
Systrophiella eudiscus (H.B. Baker, 1925)
Figure 5
Scolodonta (Systrophiella) eudiscus Baker 1925a: 31 pl. 8 g. 43;
Baker 1926: 5.
Systrophiella eudiscusTillier 1980: 68, 194, 286, 292, gs
399–403.
Systrophia eudiscaRichardson 1989: 128.
Systrophia (Systrophiella) eudiscusRamírez: 68, g. 14B.
Studied material. ANSP 140959, holotype, near La Fría,
Venezuela.
Type locality. “near La Fría”, Venezuela.
Measurements. “alt. 5.0, Shell maj. diam. 16.3, min. diam.
15.3, Aperture alt. 4.4, diam. 4.4 mm, Whorls 6¼” (Baker
1925a).
Redescription. Shell large for genus, discoid, semitranslu-
cent, greenish brown. Protoconch of circa two whorls, with
only rather indistinct growth wrinkles. Teleoconch whorls
evenly rounded, slightly aened above; suture shallow but
distinct. Sculpture on teleoconch consists of rather promi-
nent growth wrinkles. Aperture broadly lunulate, with thin,
simple peristome. Umbilicus wide, up to 47% of shell width.
R & B: Revision of the genera of Scolodontidae, 1 100
Geographic range. Venezuela.
Comparisons. is species diers from others by its wide
um bilicus and low spire. Systrophiella weyrauchi comb. nov.
from the Peruvian Andes is most similar but has a higher
spire and thicker, dark-brown periostracum.
Remarks. e anatomy of this species was described by
Baker (1925a).
Systrophiella vitrina (J.A. Wagner in Spix & Wagner,
1827) comb. nov.
Figure 6
Solarium imperforatum Spix in Spix & Wagner 1827: 25, pl. 17
g. 6; Gray 1847: 170.
Helix vitrina Wagner in Spix & Wagner 1827: 25, pl. 17, g. 6
Pfeier 1848: 109.
Helix (Helicella) imperforata (Spix)Beck 1837 in 1837–1838:
7.
Helix (Patula) vitrina WagnerAlbers 1850: 65.
Ammonoceras vitrina (Wagner)Pfeier & Clessin 1881: 15.
Streptaxis (Ammonoceras) vitrina (Wagner)Tryon 1885: 64.
Happia vitrina (Wagner)Bourguignat 1890: 40; Gude 1902:
234; iele 1927: 318; Baker 1928: 125; Haas 1953: 205;
Ramírez 1993: 35; Salgado & Coehlo 2003: 169; Agu-
do-Padrón 2008: 165; Viana & Santos 2008: 104; Santos
et al. 2010: 514, 536; Nunes & Santos 2012: 82; Oliveira
2015: 40, 43, g. A2; Birckolz et al. 2016: 150; Salvador
2019: 93; Salvador 2020: 64, gs 7–9.
Streptaxis (Happia) tumescens Suter 1900: 330, pl. 3 g. 4–4b.
Scolodonta (Happia) vitrina (Wagner)Kobelt 1905 in 1905–
1906: 49, pl. 48 gs 13, 14.
Happia (Happia) vitrina (Wagner)Schileyko 2000: 756–757,
g. 986A.
Misidentications:
Helix vitrina WagnerHidalgo 1870: 36 (= “Helix” circum plexa
Deshayes, 1839 in Férussac & Deshayes, 1820–1851?).
Helix vitrina WagnerHidalgo 1893: 84 (= “Helix” circum -
plexa Deshayes, 1839 in Férussac & Deshayes, 1820–1851?).
Happia vitrina (Wagner)Simone 2006: 228, g. 871 (=
Figure 5. Systrophiella eudiscus (H.B. Baker, 1925), ANSP 140959, paratype.
Figure 6. Systrophiella vitrina (J.A. Wagner in Spix & Wagner, 1827), ZSM 20020666, holotype.
R & B: Revision of the genera of Scolodontidae, 1 101
“Helix” circumplexa Deshayes, 1839 in Férussac & Deshayes,
1820–1851?); Barbosa 2014: 58-65, g. 5–7 (= Scolodonti-
dae sp.?); Breure & Araujo 2017: 110, g. 40D–F (=“Helix”
circumplexa Deshayes, 1839 in Férussac & Deshayes, 1820–
1851?); Salvador et al. 2018: 116–119, g. 11A–C (= ?);
Silva et al. 2019: 179, g. 2L–N (= Scolodontidae sp.); Ran-
gel et al. 2021: 4–6, g 7e (= Scolodontidae sp.).
Streptaxis tumescensSimone 2006: 194, g. 722 (= Streptaxi-
dae sp.?).
Studied material. ZSM 20020666, holotype, “Brasilien”,
leg. Spix 1827.
Type locality. “Habitat cum precedente [Provinciis aus-
tralioribus Brasiliae]”.
Measurements. “Longitudo 2½ lin.; lat. 5 lin.” (Spix & Wag-
ner 1827).
Redescription. Shell medium-sized for genus, whitish-
transparent, with some remnants of a brownish periostra-
cum and a slightly raised spire. Suture rather deep, pro-
toconch–teleoconch transition indistinct. Sculpture of
indistinct, slightly exuous growth lines. Vestigial spiral
sculpture absent. Aperture rather large, subcircular, peri-
stome too damaged for description. Umbilicus of medium
width, up to 30% of shell width.
Geographic range. Brazil.
Comparisons. It is similar to S. pygmea (Spix in Spix & Wag-
ner, 1827), but that species has a much smaller size at the
same whorl count and a less circular aperture.
Aside from S. pygmea, S. vitrina has been confused with
several poorly known and possibly undescribed species of
Scolodontidae from Brazil. For instance, Simone (2006),
Barbosa (2014), Silva et al. (2019), and Rangel et al. (2021)
gured several species with a aer spire and broadly lunu-
late aperture under the name “Happia vitrina”. ese species
can be separated from each other by the shape of the aper-
ture, colour, and dierences in size. Moreover, Salvador et
al. (2018) gured a small species unknown to us under the
name “Happia vitrina”; it has a narrow umbilicus, at to con-
cave spire, and a last whorl that partially covers the preced-
ing whorls.
“Helix” circumplexa Deshayes, 1839 is a species from the
Atlantic rainforest of Brazil, which diers from S. vitrina by its
low spire that is partially covered by the last whorl (Férussac
& Deshayes 1820–1851, 1 (30): 19, pl. 84 gs 5, 6). ere-
fore, it is not a synonym of S. vitrina, as suggested by sev-
eral authors, including Gude (1902) and Ramírez (1993).
e specimens listed as H. vitrina by Hidalgo (1870, 1893),
Simone (2006), and Breure & Araujo (2017) might be “H.”
circumplexa, but we have not had the opportunity to com-
pare these shells with the type material. e re-identication
in this paper of the specimens described by Hidalgo (1870,
1893) was based on the gure by Breure & Araujo (2017),
who imaged the specimen used in Hidalgo’s publication.
Another species oen listed as a synonym of S. vitrina,
“Helix valvaeformis ‘Nyst’”, seems to be an unpublished
manuscript name rst used by Pfeier (1847 in 1847–1848:
110) in the synonymy of Helix vitrina. Here we regard it as
a nomen nudum.
Remarks. Beck (1837 in 1837–1838) used Solarium imper-
foratum Spix, 1827 as the valid name for this species. Beck
met all the requirements of rst reviser according to ICZN
Article 24.2. Cowie et al. (2004) also considered Spix’s
names to take precedence over Wagner’s. However, Spix’s
Solarium imperforatum has not been used as valid since Gray
(1847), so replacing Wagner’s Helix vitrina with Spix’s name
would threaten the stability of the taxon in our opinion.
As the conditions of ICZN Article 23.9.1. are not met, we
cannot make the decision to maintain the name ourselves
but must refer the case to the International Commission on
Zoological Nomenclature (Roosen & Breure submied).
Meanwhile, we tentatively use Helix vitrina following pre-
vailing usage in recent papers.
Gray (1847) designated Solarium imperforatum Spix, 1827
as type species for Solarium “Spix”. However, Spix never
made this name available, but seems to have misapplied
Solarium Lamarck, 1799 (= Architectonica Röding, 1798). As
the genus itself is not valid, neither is the type designation.
Systrophiella pygmea (Spix in Spix & Wagner, 1827)
comb. nov.
Figure 7
Solarium pygmeum Spix in Spix & Wagner 1827: 25, pl. 17 g. 7.
Helix nana Wagner in Spix & Wagner 1827: 25, pl. 17 g. 7 (non
Pennant, 1777).
Happia nanaBarbosa 2014: 66–73, gs 8–10; Oliveira 2015:
40, 44, g. A3.
Studied material. ZSM 20020657, syntypes (two shells,
dry), “Brasilien”.
Type locality. “Habitat cum precedentibus [Provinciis aus-
tralioribus Brasiliae]”.
Measurements. “Longitudo 1½ lin.; lat. 3½ lin.” (Spix &
Wagner 1827).
Redescription. e type material is likely juvenile, so not all
diagnostic characteristics could be studied. Shell small for
genus, subdiscoid, with a slightly raised spire, semitranspar-
ent, yellowish brown. Distinction between protoconch and
teleoconch not clear. First whorls smooth, without visible
sculpture. Later whorls with indistinct growth lines; base
R & B: Revision of the genera of Scolodontidae, 1 102
of last whorl with indistinct vestigial spiral sculpture. Aper-
ture subcircular, with thin, simple peristome. Umbilicus of
medium width, up to 20% of shell width.
Geographic range. Brazil.
Comparisons. Systrophiella vitrina is similar, but it is larger
at the same number of whorls and has a more elevated spire
and a nearly circular aperture.
Remarks. is species has most oen been regarded as a
synonym of S. vitrina, but re-examination indicates that it
is likely a valid species, so some nomenclatural issues need
to be addressed. Both Helix nana J.A. Wagner, 1827 and
Solarium pygmeum Spix, 1827 were made available by Spix
& Wagner (1827), and as there is no ocial rst revisor
Helix nana was only treated as valid in unpublished theses
(Barbosa 2014; Oliveira 2015)that decision is made here.
As Cowie et al. (2004) explained at length, Spix’s names
should get priority over Wagner’s names in most cases.
Moreover, Wagner’s Helix nana is a junior primary homo-
nym of Helix nana Pennant, 1777 (= Planorbarius corneus
(Linnaeus, 1758)). erefore, we follow Cowie et al. (2004)
and ICZN Article 57.2 and select Solarium pygmeum Spix in
Wagner & Spix, 1827 as the valid name for this species.
e specimen from Ilha Grande (Rio de Janeiro State,
Brazil) gured by Barbosa (2014) diers only slightly from
the type material.
Genus Luteostriatella gen. nov.
ZooBank identier. urn:lsid:zoobank.org:act:3971CFD2-
C22A-46CC-9FA6-B828A5AAD314
Type species. Austroselenites variegatus F. Haas, 1949.
Description. Shell strongly depressed, conical-globular,
medium-sized for family. Periostracum with thin, yellow-
ish-white and reddish-brown axial streaks, three per 1 mm.
Sculpture of weak growth lines. Aperture subcircular, with
thin, simple peristome. Umbilicus of moderate width, c.
30% of shell width.
Species included in the genus. Luteostriatella variegata (F.
Haas, 1949) comb. nov.
Geographic range. Peru.
Derivation of the name. e name of the genus is com-
posed of the Latin word luteo and striata, which means yel-
low-striped. Its gender is feminine.
Comparisons. is species has been questionably included
in Austroselenites, but species of Austroselenites can be sep-
arated from Luteostriatella gen. nov. by their stronger axial
sculpture, larger size, lower spire, and uniformly coloured
periostracum. Ramírez (1993) placed the species in Systro-
phiella H.B. Baker, 1925, which diers from Luteostriatella
gen. nov. by its vestigial spiral sculpture, uniformly coloured
periostracum and lower spire.
Figure 7. Systrophiella pygmea (Spix in Spix & Wagner, 1827). A–C, ZSM 20020657, syntype. B, apical surface of same shell. C, basal
surface of same shell, in umbilicus (not to scale).
R & B: Revision of the genera of Scolodontidae, 1 103
Remarks. Over 70 years aer the original description of
L. variegatus, an additional specimen was photographed in
the Peruvian Amazon. As it is unlikely more material will
become available soon, so we are describing the genus here
based on the available information. We are placing it in the
Scolodontidae based on its conchological similarities to Sys-
trophiella, but we acknowledge that studies of its DNA and
anatomy may reveal that it belongs to the Streptaxidae.
Luteostriatella variegata (F. Haas, 1949) comb. nov.
Figure 8
Austroselenites (subgenus?) variegatus Haas 1949: 247, g. 59.
Systrophia variegataRichardson 1989: 140.
Systrophia (Systrophiella) variegataRamirez 1993: 72.
Studied material. FMNH 30037, holotype (dry), Peru,
Loreto department, near Rio Ucayali.
Type locality. Peru: “Cerro Azul, on Rio Ucayali, Depart-
ment of Loreto”.
Measurements. “Diameter 10.3 mm, height 5.5 mm, diam-
eter of umbilicus 3.9 mm” (Haas 1949).
Redescription. Shell strongly depressed, conical-globular,
medium-sized for family; suture shallow. Periostracum with
thin, yellowish-white and reddish-brown axial streaks, three
per 1 mm. Spire whorls slightly damaged, without sculp-
ture. Teleoconch sculpture with minute growth lines. Aper-
ture subcircular, with thin, simple peristome. Umbilicus of
medium width, up to 31% of shell width.
Geographic range. Peru: Loreto Department: near Rio Ucay-
ali and Maynas Province (iNaturalist observation 15694700,
hps://www.inaturalist.org/observations/15694700).
Comparisons. ere are no similar species currently known.
Remarks. e placement of L. variegata in Scolodontidae
is tentative and should be corroborated by anatomical and
genetic research.
Genus Austroselenites Kobelt, 1905
Scolodonta (Austroselenites) Kobelt 1905 in 1905–1906: 49,
70. Type species: Helix euspira Reeve, 1854 by subsequent
designation (Baker 1925a: 15).
Redescription. Shell medium-sized to large, discoid to
subdiscoid, broadly umbilicate. Spire strongly depressed to
slightly elevated. Protoconch smooth, no clear distinction
between protoconch and teleoconch. Teleoconch with nar-
row, dense axial ribs, which become weaker on later whorls.
Periostracum yellow-brown. Whorls usually convex or at-
tened above. Aperture broadly lunulate to rounded-trape-
zoidal, with simple peristome. Living animal yellowish, with
red antennae (Nicole Büner pers. comm. 2020; Fig. 9).
Species included in the genus. Austroselenites euspira
(Reeve, 1854), A. ora (L. Pfeier, 1850), A. moyobambensis
(J. Moricand, 1858), A. pichinchense sp. nov.
Species tentatively included in the genus. ?Austroselenites
cyclina (Cousin, 1887)
Geographic range. Venezuela, Brazil, Peru, and Ecuador.
Comparisons. Species included in Austroselenites were of-
ten assigned to Happia (e.g. Breure et al. 2022). However,
Happia contains discoid species that, to our knowledge,
only reach 4.5 mm wide when adult. Moreover, all Happia
Figure 8. Luteostriatella variegata (Haas, 1949). A, FMNH 30037, holotype. B, living specimen (iNaturalist observation 15694700).
R & B: Revision of the genera of Scolodontidae, 1 104
species have spiral rows of microscopic papillae, which do
not occur in Austroselenites. Austroselenites are most similar
to Systrophiella, which diers by its smaller, semitransparent
shells lacking strong axial sculpture.
Austroselenites ora was included in Mesomphix Ran-
esque, 1819 in MolluscaBase (2023c). However, Mesomphix
is known from North America, and all species have smaller
shells with a narrow to nearly closed umbilicus. In addition,
Mesomphix is not known to have the yellowish body, as in
Austroselenites. e source which persuaded MolluscaBase
editors to move the species to Mesomphix is unknown.
Remarks. Austroselenites Kobelt, 1905 was described for
A. euspira (Reeve, 1854) and A. moyobambensis, but Baker
(1941) and Schileyko (2000) included only A. euspira in
this genus. However, based on re-examination of several
shells of A. euspira, A. ora, A. pichinchense sp. nov., and A.
moyobambensis, they are all similar in microsculpture on the
early whorls, shape of the aperture, and colour of the perio-
stracum. e only shell characteristic hinting towards sepa-
ration of A. euspira and the other taxa at the generic level is
the slightly smaller size of A. euspira.
Its yellow mantle, along with a few shell characteristics,
suggests true Austroselenites belongs to the Scolodontidae
rather than Haplotrematidae. is agrees with the inclu-
sion of its species in Happia sensu lato by most authors. e
species studied by Baker (1941) seem to dier from Aus-
troselenites sensu stricto and were correctly referred to Zophos
Gude, 1911 by him. In our opinion, Baker (1941) provided
sucient evidence to place Zophos in the Haplotrematidae.
erefore, we follow Schileyko (2000) in treating Zophos as
a genus in the Haplotrematidae. As Austroselenitinae Baker,
1941 was largely based on the examination of Zophos spe-
cies (Baker 1941), we refrain from using it here pending
further study.
To beer illustrate the microsculpture of this genus, we
include SEM photomicrographs of an unidentiable juve-
nile Austroselenites sp. from Cotopaxi province, Ecuador
(Fig. 10; QCAZ-I-274879).
Figure 9. Austroselenites sp., living specimen, on road around Las Tolas (Gualea), Pichincha Province, Ecuador, at 1700 m (Nicole Bü -
ner pers. comm. 2020).
Figure 10. Austroselenites sp., QCAZ-I-274879.
R & B: Revision of the genera of Scolodontidae, 1 105
Austroselenites euspira (Reeve, 1854)
Figure 11
Helix euspira Reeve 1854: pl. 185, g. 1277Pfeier 1855a:
54; Brown 1866: 15.
Ammonoceras euspira (Pfeier, 1855)Albers & Martens
1860: 72.
Hyalina (Ammonoceras) euspira PfeierMartens 1873: 167.
Macrocyclis euspira (Pfeier)Binney 1876: 247.
Happia euspira (Pfeier)Gude 1902: 233; Simone 2006:
226, g. 863.
Scolodonta (Austroselenites) euspira (Pfeier)Kobelt 1905 in
1905–1906: 70, pl. 52 gs 1–3.
Haplotrema (Austroselenites) euspira (Pfeier)Baker 1941:
423.
Haplotrema euspira (Pfeier)Richardson 1988: 22.
Austroselenites euspira (Pfeier)Bouchet et al. 2017: 44.
Studied material. NHMUK 20230164, syntypes (3 shells,
dry), “Brazil”.
Type locality. “ Br az i l”.
Measurements. “Diam. maj. 26, min. 22½, alt. 9 mill.”
(Pfeier 1855a).
Redescription. Shell small for genus (but large for family),
subdiscoid, with a distinct suture and a yellow-brown perio-
stracum. Protoconch–teleoconch transition unclear. Sculp-
ture of rather strong, slightly exuous growth wrinkles, that
weaken on base of shell. Aperture broadly lunulate, with
Figure 11. Austroselenites euspira (Reeve, 1854). A–C, NHMUK 20230164, syntype. B, C, apical surface of same shell.
R & B: Revision of the genera of Scolodontidae, 1 106
thin, simple peristome. Umbilicus wide, up to 26% of shell
width.
Geographic range. Venezuela (Martens 1873) and Brazil.
Comparisons. Austroselenites euspira (Reeve, 1854) diers
from all other Austroselenites species by its smaller size. e
axial ribbing of the teleoconch is stronger in Andean taxa,
like A. ora and A. pichinchense nov. sp.
Remarks. We have been unable to check the Venezuelan shells
in the ZMB to verify the record of Martens (1873). Examina-
tion of the type material in the NHM may help researchers
rediscover this species, so the systematic position of Austrose-
lenites and the distribution of A. euspira can be claried.
Austroselenites pichinchense sp. nov.
Figure 12
ZooBank identier. urn:lsid:zoobank.org:act:09ED417F-
772A-4952-9617-EF8677FC8B2A
Type locality. Ecuador: Pichincha Province, between Cala-
calí and Nanegalito at 2100 m.
Type material. Holotype: dry shell, QCAZ-I 274877; from
the type locality.
Paratype: 1 dry shell, QCAZ-I 274878; Ecuador: Pichin-
cha Province: La Favorita, “under log”.
Measurements.
H (mm) W (mm) HA (mm) WA (mm) Whorls
Holotype 14.6 25.4 12.2 11.9 4¼
Paratype 17.8 30.6 14.7 14.4 4½
Description. Shell large (Table 2), depressed, with a slightly
elevated spire; whorls convex; periostracum yellow. Nucleus
small, partially covered by succeeding whorl. Protoconch
smooth, with transition between protoconch and teleo-
conch indistinct. Teleoconch sculpture of slender, densely
spaced, slightly exuous growth lines, which are more dis-
tinct on apical surface and fade away in umbilicus. Umbi-
licus of medium width, up to 20% of shell width. Aperture
subcircular, with peristome simple.
Derivation of the name. e new species is named aer the
province of Pichincha, where the specimens were found.
Geographic range. Ecuador: Pichincha province, known
form the type locality and La Favorita.
Comparisons. e new species is most similar to large
individuals of Austroselenites ora and ?A. cyclina from
which it diers by its higher spire, more circular aperture,
smaller umbilicus, and less excavated suture. Other species
lack periostracum, are much smaller at the same number of
whorls, and/or have a more elongate aperture.
Remarks. Austroselenites ora might occur in somewhat the
same area as A. pichinchense sp. nov.
A living specimen photographed near Las Tolas (Gualea)
in Pichincha Province, Ecuador, likely belongs to A. pi chin-
chense, but its identity cannot be conrmed from the photo-
graphs (Fig. 9).
D
Pfeier, Reeve, Spix, or Wagner?
One issue that needs to be addressed is the authorship of
some of the type species and other taxa in this paper. Sev-
eral taxa were previously aributed to Pfeier based on what
seems to be the intended order of publication (e.g. Roosen
2023), for instance Austroselenites ora (Reeve, 1854), Hap-
pia ammonoceras (Reeve, 1854), and Hirtudiscus excisa
(Reeve, 1854). During our study we concluded that Reeve
(1854) was published before Pfeier (1855), even though
Figure 12. Austroselenites pichinchense sp. nov., QCAZ-I 274878, holotype.
R & B: Revision of the genera of Scolodontidae, 1 107
Reeve (1854) clearly cited Pfeier (1855) as the original
source. As all taxa by Pfeier (1855) are also treated by
Reeve (1854), Reeve is the rst person to describe these
species and, thus, has priority over Pfeier.
Determining correct authorship and applying the prin-
ciple of priority to the names published by Spix & Wagner
(1827) is more dicult. First, Spix and Wagner’s paper is
oen aributed to only Wagner, although from the title and
further analysis it is clear Spix made substantial contribu-
tions. erefore, the paper should be cited as Spix & Wagner
(1827), as done by Salgado & Coelho (2003) and suggested
to us by Francisco Welter-Schultes (pers. comm. 2023). We
can assume that the names on the plates of Spix & Wagner
(1827) were contributed by Spix, as Wagner nished the
paper aer the plates were already made (Cowie et al. 2004).
We agree with Cowie et al. (2004) that new names and spell-
ings introduced in the main text of Spix & Wagner (1827)
can be aributed to Wagner. All names are made available
in the same paper, so the rst revisor decides which name
should be treated as valid (ICZN Article 24.2.1; Francisco
Welter-Schultes pers. comm. 2023).
Decisions regarding Happia
Some of our decisions regarding Happia Bourguignat, 1890
need further explanation. Most authors consider Helix vitrina
Wagner, 1827 as the type species (e.g. Ramírez 1993; Salvador
et al. 2018). Although, if we do not accept Pfeier’s (1855b)
type designation by absolute tautonomy for Ammonoceras
(= Happia), the subsequent designation of Helix euspira
Reeve, 1854 as type species for Ammonoceras by Bland
(1863) would have priority over later designations. is
would make Austroselenites Kobelt, 1905 a junior objective
synonym of Happia, leaving Helix ammonoceras without a
valid genus and still causing Helix vitrina to be moved to Sys-
trophiella. If Helix ammonoceras is retained as type species
for Ammonoceras Pfeier, 1855, and by extension Happia,
at least Austroselenites and Helix ammonoceras remain stable.
at said, until now, many more species have been
included in Happia. Discussing all these species is beyond
the scope of our study, but they do need to be placed in a
genus. As the dominant denition for Happia was the cur-
rent denition of Systrophiella in this paper, we propose that
all “orphaned” Happia species are placed in Systrophiella
sensu lato pending further study.
Systrophiella vitrina (Wagner in Spix & Wagner, 1827)
Strict adherence to the ICZN would mean that Helix vitrina
Wagner, 1827 should be replaced with Solarium imperfora-
tum Spix, 1827, because Beck (1837 in 1837–1838: 7) used
the laer name as valid and his paper meets all requirements
to be First Revisor. However, as shown in the systematic
section, this name was not used since Gray (1847). All sub-
sequent authors regarded Helix vitrina as the valid name,
but not enough papers have been published to trigger an
automatic reversal of precedence. As we believe using Solar-
ium imperforatum would cause unnecessary confusion and
threaten the stability of this already oen confused taxon,
we will petition the ICZN to retain the name.
Austroselenites and the Austroselenitinae
Another decision with high impact made in this paper is
the tentative move of Austroselenites sensu stricto to the
Scolodontidae. However, its conchological similarity to Sys-
trophiella and yellow body colour support this view. Even
the variation within the genus, which seems largely limited
to microsculpture, aperture shape, and spire height, ts
within our current expectations of the Scolodontidae.
at said, it is unlikely that the entire Austroseleniti-
nae of Baker (1941) should be a subfamily of the Scolo-
dontidae. Zophos, a genus oen regarded as a subgenus of
Austroselenites and the genus on which Austroselenitinae
was based, has a grey body and its anatomy, as described
by Baker (1941), does seem to support placement in the
Haplotrematidae. Because of all this, we propose to regard
Zophos as a separate haplotrematid genus and not to use the
Austroselenitinae until the DNA and anatomy of the type
species of Austroselenites are known.
General discussion and conclusion
In this paper we redescribe several, oen poorly known,
genera of Scolodontidae and their type species, and where
necessary also redescribe or describe additional species to
further stabilise the genera. However, this is only one small
step. e anatomy of the type species, except for Systrophiella
eudiscus (Baker, 1925), is still unknown, and their DNA has
not been sequenced yet. Although many authors have tried
to nd relationships by shell and radular morphology (e.g.
Gude 1902; Ramírez 1993; iele 1927; Tillier 1980), we
refrain from doing so, as another potentially wrong interpre-
tation of Scolodontidae is not needed. Besides, such a study
is also beyond the scope of this paper.
We sincerely hope that our redescriptions provide a
baseline for further research and are useful for malacologists
working on this taxonomically dicult group.
A
First, we thank Francisco Welter-Schultes for advice on how
to handle the nomenclatural issues concerning Systrophiella
R & B: Revision of the genera of Scolodontidae, 1 108
vitrina, Jonathan Able (NHM) for checking the grammar of
an earlier version of the manuscript and providing all images
of type specimens deposited in the NHMUK, and Enrico
Schwabe (SNSB) for loaning us the syntypes of Helix vitrina
and H. pygmea. We are also grateful to Yves Baree (RBINS)
for photographing the type material of Systrophiella vitrina
and S. pygmea and Laetitia Despontin (RBINS) for SEM
imaging the syntype of S. pygmea. Diego Eduardo Gutiér-
rez (MLP) made the photographs of the type specimen of
Hirtudiscus hirtus and provided SEM images of the same
shell originally made by Bernhard Hausdorf (LIB). Last, we
would like to thank Paul Callomon (ANSP) for the detailed
images of the type specimen of Happia andia and Systro-
phiella eudiscus, Fernanda Salazar (PUCE) for allowing us to
study part of the QCAZI collection, Lee Doolan for allow-
ing us to use his photograph of living Luteostriatella varie-
gata, and the anonymous reviewers for greatly improving
our manuscript.
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