Rediscovery of Obeliscus agassizi Pilsbry, 1906 (Gastropoda, Subulinidae, Obeliscinae), annotated checklist of species of Obeliscus Beck, 1837 and first description of the anatomy for the genus

Abstract

Obeliscus agassizi Pilsbry, 1906 was described from specimens collected in Brazil, by J. G. Anthony, during the Agassiz expedition, in 1865. The type locality of the species was given simply as Brazil, without further information. More than a century after its description, this species was rediscovered during an expedition to the Biological Reserve of Pedra Talhada, Alagoas/Pernambuco states, Brazil. In the present study we provide detailed description of the anatomy of the soft parts of O. agassizi specimens, collected during this expedition, besides more precise data concerning the species occurrence. This work presents the first anatomical description for the genus. An annotated checklist of Obeliscus Beck, 1837 species is also provided. The species of Obeliscus have been traditionally distinguished by shell characteristics, mainly the general shell shape, number of whorls proportionally to the shell size, shell ornamentation, the shape of the protoconch and aperture size and shape. The distinction between species is often subtle. Considering that shell traits may be not sufficient as single taxonomic characters, other evidence for species boundaries, especially anatomical characters, is needed. Presently, it is not possible to discuss the taxonomic meaning of the set of anatomical characters described for O. agassizi, at the intra-genus level, as there is no information on the anatomy of the soft parts of other species of Obeliscus. The comparative analysis of the genital system of O. agassizi, Neobeliscus calcareus (Born, 1780), Stenogyra terebraster (Lamarck, 1822) and Rectobelus birabeni (Hylton-Scott, 1946) pointed out anatomical characters with potential diagnostic significance for the genus Obeliscus. However, the establishment of a definitive differential diagnosis for this genus, based in both conchological and anatomical traits, depends on the analysis of a greater number of species of Obeliscus, as well as species of other genera of Obeliscinae Thiele, 1931.

Full text

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COMITÉ SCIENTIFIQUE / SCIENTIFIC BOARD :
James Carpenter (AMNH, New York, États-Unis)
Maria Marta Cigliano (Museo de La Plata, La Plata, Argentine)
Henrik Enghoff (NHMD, Copenhague, Danemark)
Rafael Marquez (CSIC, Madrid, Espagne)
Peter Ng (University of Singapore)
Norman I. Platnick (AMNH, New York, États-Unis)
Jean-Yves Rasplus (INRA, Montferrier-sur-Lez, France)
Jean-François Silvain (IRD, Gif-sur-Yvette, France)
Wanda M. Weiner (Polish Academy of Sciences, Cracovie, Pologne)
John Wenzel (The Ohio State University, Columbus, États-Unis)
COUVERTURE / COVER :
Obeliscus agassizi Pilsbry, 1906 living crawling specimen, shell length c. 6 mm.
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159ZOOSYSTEMA • 2020 • 42 (12) © Publications scientifi ques du Muséum national d’Histoire naturelle, Paris. www.zoosystema.com
urn:lsid:zoobank.org:pub:BDBBCF4F-D739-422C-AF7F-17166F5966C8
D’ávila S., Simone L. R. L., Oliveira L. F. Cappa De, Charles L. & Maestrati P. 2020. — Rediscovery of Obeliscus agassizi
Pilsbry, 1906 (Gastropoda, Subulinidae, Obeliscinae), annotated checklist of species of Obeliscus Beck, 1837 and fi rst
description of the anatomy for the genus. Zoosystema 42 (12): 159-172. https://doi.org/10.5252/zoosystema20v42a12.
http://zoosystema.com/42/12
ABSTRACT
Obeliscus agassizi Pilsbry, 1906 was described from specimens collected in Brazil, by J.G. Anthony,
during the Agassiz expedition, in 1865.  e type locality of the species was given simply as Brazil,
without further information. More than a century after its description, this species was rediscovered
during an expedition to the Biological Reserve of Pedra Talhada, Alagoas/Pernambuco states, Brazil. In
the present study we provide detailed description of the anatomy of the soft parts of O.agassizi speci-
mens, collected during this expedition, besides more precise data concerning the species occurrence.
 is work presents the  rst anatomical description for the genus. An annotated checklist of Obeliscus
Beck, 1837 species is also provided.  e species of Obeliscus have been traditionally distinguished by
shell characteristics, mainly the general shell shape, number of whorls proportionally to the shell size,
shell ornamentation, the shape of the protoconch and aperture size and shape.  e distinction between
Sthefane D’ÁVILA
Museu de Malacologia Prof. Maury Pinto de Oliveira, Instituto de Ciências Biológicas,
Universidade Federal de Juiz de Fora, Rua José Lourenço Kelmer, s/n - Martelos,
Juiz de Fora, 36036-330 Minas Gerais (Brazil)
*corresponding author: sthefane.davila@ufjf.edu.br, sthefanedavila@hotmail.com
Luiz Ricardo L. SIMONE
Museu de Zoologia da Universidade de São Paulo,
Avenida Nazaré 481, 04218-970 São Paulo (Brazil)
lrsimone@usp.br; lrlsimone@gmail.com
Luiz Fernando Cappa de OLIVEIRA
Núcleo de Espectroscopia e Estrutura Molecular, Instituto de Ciências Exatas,
Universidade Federal de Juiz de Fora, Rua José Lourenço Kelmer, S/n - Martelos,
Juiz de Fora, 36036-330 Minas Gerais (Brazil)
luiz.oliveira@ufjf.edu.br
Laurent CHARLES
Muséum d’Histoire naturelle, 5 place Bardineaux, F-33000 Bordeaux (France)
l.charles@mairie-bordeaux.fr
Philippe MAESTRATI
Muséum national d'Histoire naturelle de Paris
Direction générale déléguée aux Collections, pôle Expédition
case postale 51, 57 rue Cuvier, F-75231 Paris cedex 05 (France)
philippe.maestrati@mnhn.fr
Submitted on 30 November 2018 | Accepted on 11 June 2019 | Published on 21 April 2020
Rediscovery of Obeliscus agassizi Pilsbry, 1906 (Gastropoda,
Subulinidae, Obeliscinae), annotated checklist of species
of
Obeliscus Beck, 1837 and fi rst description of the anatomy
for the genus

160 ZOOSYSTEMA • 2020 • 42 (12)
D’ávila S. et al.
species is often subtle. Considering that shell traits may be not su cient as single taxonomic characters,
other evidence for species boundaries, especially anatomical characters, is needed. Presently, it is not pos-
sible to discuss the taxonomic meaning of the set of anatomical characters described for O.agassizi, at the
intra-genus level, as there is no information on the anatomy of the soft parts of other species of Obeliscus.
 e comparative analysis of the genital system of O.agassizi, Neobeliscus calcareus (Born, 1780), Stenogyra
terebraster (Lamarck, 1822)
and Rectobelus birabeni (Hylton-Scott, 1946)
pointed out anatomical characters
with potential diagnostic signi cance for the genus Obeliscus. However, the establishment of a de nitive dif-
ferential diagnosis for this genus, based in both conchological and anatomical traits, depends on the analysis
of a greater number of species of Obeliscus, as well as species of other genera of Obeliscinae  iele, 1931.
RÉSUMÉ
Redécouverte d’ Obeliscus agassizi Pilsbry, 1906 (Gastropoda, Subulinidae, Obeliscinae), liste commentée des
espèces d’ Obeliscus et première description de l’anatomie du genre.
Obeliscus agassizi Pilsbry, 1906 a été décrit à partir de spécimens recueillis au Brésil par J.G. Anthony, pen-
dant l’expédition Agassiz en 1865. La localité type de cette espèce a tout simplement été donnée comme
«Brésil », sans autre information. Plus d’un siècle après sa description, cette espèce a été redécouverte
au cours d’une mission à la Réserve biologique de Pedra Talhada, Alagoas/Pernambuco, au Brésil. Nous
donnons ici une description détaillée de l’anatomie de O.agassizi, ainsi que des données plus précises sur
la distribution de l’espèce. Nous donnons également la première description de l’anatomie du genre Obe-
liscus Beck, 1837. Une liste commentée des espèces du genre est également donnée. Les espèces du genre
Obeliscus ont toujours été distinguées d’après les caractères conchyliologiques, comme la forme générale, le
nombre de tours proportionnellement à la taille, l’ornementation, la forme et la taille de la protoconque et
de l’ouverture. La distinction entre les espèces est souvent subtile. Considérant que les caractères conchy-
liologiques sont insu sants comme seuls caractères taxonomiques, d’autres sources d’information sur les
limites entre espèces, en particulier les caractères anatomiques, deviennent nécessaires. Actuellement, il
n’est pas possible d’argumenter sur la signi cation taxonomique des caractères anatomiques décrits pour
O. agassizi, au niveau intragénérique, puisqu’il n’y a pas d’informations sur l’anatomie des parties molles
chez d’autres espèces d’Obeliscus. L’analyse comparative du système génital d’Obeliscus agassizi, Neobeliscus
calcareus (Born, 1780), Stenogyra terebraster (Lamarck, 1822)
et Rectobelus birabeni (Hylton-Scott, 1946)
font ressortir des caractères anatomiques pouvant peut-être servir à la diagnose d’Obeliscus. Cependant, la
dé nition d’une diagnose di érentielle pour le genre nécessitera l’étude d’un plus grand nombre d’espèces
d’Obeliscus, et de genres de la sous-famille Obeliscinae  iele, 1931.
INTRODUCTION
Subulinidae
Fischer & Crosse, 1877
is a near pan-tropical family
of land snails, including nine subfamilies, 81 genera and about
820 species (Pilsbry 1906; Schileyko 1999).  e systematics of
this family has been entirely based on shell characteristics and
the attempts to add new data about this family have been very
scarce throughout the last century. Hence, Subulinidae  gures
as a neglected group of land snails, contrasting with its diversity.
Species delimitation based only on shell characters has been
proved to be inadequate for several families of pulmonate
snails, e.g. Lymnaeidae Ra nesque, 1815
(Pfenninger etal.
2006), Bradybaenidae Pilsbry, 1934 (Chiba& Davison 2008),
Helicidae
Ra nesque 1815
(Elejalde etal. 2008), Hygromiidae
Tryon, 1866
(Dépraz etal. 2009), Camaenidae
Pilsbry, 1895
(Köhler& Burghardt 2015), Enidae
Pilsbry, 1895
(Modica
etal. 2016), and Pyramidulidae Kennard & Woodward, 1914
(Razkin etal. 2017); since it leads to some taxonomic issues
as the di cult detection of cryptic species or, on the other
hand, the overestimation of species diversity.  is observation
pinpoints the need of revisiting the anatomy of soft parts,
which has been often associated to molecular data, for a more
precise delimitation of the species.
Considering the lack of other criteria of discrimination
besides the shell (available data on the internal anatomy of
Subulinidae concern 30 genera and no more than 32 species
[Schileyko 1999; Medeiros etal. 2013]), and taking into ac-
count that no full systematic revision of the family has been
provided during more than a century, the description of the
anatomy of the soft parts as well as the characterization of
life history traits of subulinid species is a necessary challenge.
Without these two sources of information, studies on ecology,
evolution, phylogeny and conservation of subulinid species
will remain in a state of impediment.
Obeliscinae  iele, 1931 comprises 12 American, one
Asian and three Paci c genera (Schileyko 1999). Among the
12 American genera,  ve were recorded in Brazil, including
at least 13 endemic species (Pilsbry 1906; Simone 2006).
Only Neobeliscus calcareus (Born, 1780) (Pilsbry 1899, 1906),
Stenogyra terebraster (Lamarck, 1822) (Schileyko 1999) and
Rectobelus birabeni (Hylton-Scott, 1946) were characterized
from an anatomical perspective. So far, all other species in-
cluded in this subfamily are known only from their shells.
More than a century ago, Pilsbry (1906) stated that besides
the information on the anatomy of N.calcareus given by
him, nothing was known of the soft parts of the other genera.
MOTS CLÉS
Morphologie,
étude anatomique,
mollusque terrestre,
subulinides,
Achatinoidea.
KEY WORDS
Morphology,
anatomical study,
land snail,
subulinids,
Achatinoidea.

161
Rediscovery of Obeliscus agassizi
ZOOSYSTEMA • 2020 • 42 (12)
 e more recent revision of the family Subulinidae was
performed by Schileyko (1999), who has provided new in-
formation on the anatomy of the soft parts for several genera.
Schileyko (1999) illustrated the anatomy of the genital system
of S.terebraster (new data) and of N.calcareus after Pilsbry
(1906). However, the author did not describe the anatomy
of the other genera, including Obeliscus Beck, 1837, the type
genus. Schileyko (1999) argued that the characters previ-
ously used to de ne Obeliscinae are not su cient, proposing
the maintenance of this subfamily until information on the
anatomy of the soft parts of their representatives is available.
 e genus Obeliscus Beck, 1837 includes about
16 spe-
cies, distributed throughout Tropical South America and
Greater Antilles. So far, none of these species was anatomi-
cally characterized.  e diagnosis of the genus reviewed by
Pilsbry (1906) includes only shell characteristics and some
fragmentary information on the radula of O.obeliscus, taken
from the remains washed from a dry shell. After the work of
Pilsbry (1906), other additional  ve species of Obeliscus were
described based on shells only (Vanatta 1918; Pilsbry 1944;
Simone& Salvador 2016;  ach 2017).
Obeliscus agassizi Pilsbry, 1906 was described from speci-
mens collected in Brazil by J.G. Anthony, during the Agassiz
expedition.  e type locality of the species was given simply
as Brazil, without further information. More than a century
after its description, this species was rediscovered by Maestrati
etal. (2015) during an expedition to the Biological Reserve of
Pedra Talhada, Alagoas/Pernambuco states, Brazil. In the pre-
sent study we provided detailed description of the anatomy of
O.agassizi, based on specimens collected during that expedition.
An annotated checklist of Obeliscus species is also provided.
MATERIALS AND METHODS
 e material consists of four specimens of Obeliscus agassizi,
preserved in ethanol 70% (MZSP 133507; MZSP 133635),
and deposited in the malacological collection of the Museu
de Zoologia da Universidade de São Paulo (MZSP). All speci-
mens were collected in the Pedra Talhada Biological Reserve,
located in the municipalities ofQuebranguloin the state
ofAlagoasand ofLagoa do Ouroin the state ofPernambuco.
Pedra Talhada is a federally administered biological reserve in
North East Brazil, which contains a representative sample of
the remaining Atlantic Rainforest ecosystem in the Serras of
Guaribas, Pedra Talhada and Serra do Cavaleiro.
 e snails were dissected immerse in ethanol and drawn un-
der an Olympus stereo microscope, model SZX7As, equipped
with camera lucida. Details of the shell were acquired through
tabletop scanning electron microscope EDS. Shell specimens
were analyzed without any preparation.
Identi cation was conducted based on the species origi-
nal description provided by Pilsbry (1906) as well as on the
revision of the family Subulinidae provided by Schileyko
(1999), images of the shells of type specimens available in
the online collection database of the ANSP, and the catalogue
by Simone (2006).
 e terminology employed to designate the structures of
the reproductive system is mostly in accordance to recent pa-
pers (Gómez 2001; Bojat etal. 2001; Koene& Schulenburg
2005; Beese etal. 2006; Garefalaki etal. 2010; Simone 2011;
Schmera etal. 2016).
To provide an annotated checklist of the species of the genus
Obeliscus an intensive research was performed using both the
literature and collections databases of the Academy of Natu-
ral Sciences of Philadelphia, Bailey-Matthews National Shell
Museum, California Academy of Science, Carnegie Museum
of Natural History, Field Museum of Natural History, Florida
Museum of Natural History, Illinois Natural History Survey,
Löbbecke Museum Dusseldorf, Museu Nacional do Rio de
Janeiro, Muséum national de Histoire naturelle, Museum of
Comparative Zoology, Naturalis Center of Biodiversity, Ohio
State Museum of Biological Diversity, Royal Belgian Institute
of Natural Sciences, South African National Biodiversity
Institute, United States National Museum – Smithsonian
Institution, University of Michigan Museum of Zoology
and Zoologische Staatssammlung München – Mollusca, and
MZSP.  e data were obtained from the Institutions websites
or from data portals as Global Biodiversity Information Fa-
cility, Invertebase Data Portal, Sys Tax – Zoological Collec-
tions and SpeciesLink (Pimenta 2017; Grant& Jones 2017;
Academy of Natural Sciences 2018; Morris 2018; Orrell&
Hollowell 2018; Paulay& Brown 2018; Royal Belgian In-
stitute of Natural Sciences 2017; Systax 2018).  e search in
these online databases allowed us to update the geographical
distribution of some species. All museum depositits numbers
were provided in the checklist, including information on type
specimens. Museum deposit numbers were also referenced
in the topic “other localities” when occurrence data were
obtained from the databases consulted online. We also pro-
vided the website links for the species original descriptions,
illustrations, museum deposits and bibliographic references
cited in the checklist.
All the species are listed according to their current taxo-
nomic status.  e citations of the specimens from the collec-
tions databases are in accordance with the format proposed
by Chester et al. (2019). Some information were sometimes
not available in the databases, such as the number or the kind
of available specimens (dry shell or ethanol preserved shell
and soft parts).
ABBREVIATIONS
In  gures
ac albumen chamber;
ag albumen gland;
al allospermoduct;
at autospermoduct;
au auricle;
bc bursa copulatrix;
cm columellar muscle;
cv pulmonary vein;
da anterior lobe of digestive gland;
dg posterior lobe of digestive gland;
di diaphragm;
es esophagus;
ey eye;

162 ZOOSYSTEMA • 2020 • 42 (12)
D’ávila S. et al.
ft foot;
fo free oviduct;
fp female pore;
gg genital gland;
gm genital muscle;
go gonad;
hd hermaphrodite duct;
he head;
ki kidney;
mb mantle border;
om ommatophore;
pm penial muscle;
pn pneumostome;
pt prostate;
pu pulmonary (pallial) cavity;
rt rectum;
so spermoviduct;
sr seminal receptacle;
st stomach;
ur ureter;
vd vas deferens;
ve ventricle.
Institutions
ANSP
Academy of Natural Sciences of Philadelphia, Philadelphia;
BMSM Bailey-Matthews National Shell Museum, Sanibel;
CMNH Carnegie Museum of Natural History, Pittsburgh;
FLMNH Florida Museum of Natural History, Gainesville;
FMNH Field Museum of Natural History, Chicago;
LMD Löbbecke Museum Dusseldorf, Dusseldorf;
MCZ Museum of Comparative Zoology, Cambridge, MA;
MNHN Muséum national d’Histoire naturelle, Paris;
MNRJ Museu Nacional do Rio de Janeiro, Rio de Janeiro;
MZSP Museu de Zoologia da Universidade de São Paulo;
NBCN Naturalis Biodiversity Center,  e Netherlands, Leiden;
RBINS Royal Belgian Institute of Natural Sciences, Brussels;
SANBI South African National Biodiversity Institute, Cape Town;
USNM United States National Museum – Smithsonian Institu-
tion, Washington D.C.
RESULTS
Family SUBULINIDAE Fischer & Crosse, 1877
Subfamily OBELISCINAE  iele, 1931
Genus Obeliscus Beck, 1837
Obeliscus agassizi Pilsbry, 1906
Obeliscus agassizi Pilsbry, 1906: 249, pl.36,  g.76.— Clench&
Turner 1962: 7.— Salgado& Coelho 2003: 155.— Simone 2006:
187,  g.686.
TYPES.— Lectotype. ANSP 59334 • Paralectotypes. ANSP 465075;
FMNH 140941.
TYPE LOCALITY.— Brazil (collected by J.G. Anthony, during the
Agassiz expedition), no speci c locality.
O
THER
LOCALITIES
.— Brazil, Alagoas and Pernambuco states,
Biological Reserve of Pedra Talhada (present study).
M
ATERIAL
EXAMINED
.—
Brazil • Pedra Talhada Biological Reserve,
municipalities of Quebrangulo Alagoas state and Lagoa do Ouro,
Pernambuco state; MZUSP 133507 • MZUSP 133635.
HABITAT.— In leaf-litter.
MORPHOLOGICAL DESCRIPTION
Shell (Fig.1A-H)
Shell turriform-conic, minutely perforate, not decollated,
dextral, tapering; periostracum thin, yellowish, opaque in
fully developed individuals, glossy in young individuals.
Columella nearly straight, slightly concave (Fig.1C). Em-
bryonic whorls smooth, shining (Fig.1A). Protoconch with
c.3whorls. Boundary between protoconch and teleoconch
well de ned. Last whorls plicate below suture, nearly smooth
elsewhere (Fig.1B). Spire of c.12whorls, convex, very regu-
larly increasing (Fig.2A). Sutures well marked, a little inclined
(Fig.1B, E). Teleoconch of c.10whorls, smooth, except for
growth lines (Fig.2D). Aperture somewhat oblique, ovate
(Fig.1C); inner lip simple, outer lip thin.
Head-foot (Figs1D-F; 2B, C)
Head protruded. Head-foot almost transparent in living
animals, color pale cream, with  ne tessellation (Figs1D-F,
2B). Foot broad, sole oval, edges rounded in  xed animals.
Genital opening visible as a rounded area, without tessella-
tion in  xed animals, and as a small  ssure near head in living
animals (Fig.2B).
Mantle organs (Fig.2C)
Mantle edge simple, thick. Rectum narrow, walls thick, run-
ning along right edge of pulmonary cavity. Ureter running
parallel to rectum, both structures covered by  ne membrane.
Pulmonary cavity very long and narrow, occupying the two
last whorls. Kidney and pericardium restricted to middle and
left sides, occupying c.½  of pallial roof. Pulmonary vein
running along entire pallial cavity, oblique in anterior ,
running along left edge in remaining regions.
Circulatory and excretory systems (Fig.2C)
Pericardium located at columellar margin of posterior end
of pallial cavity, c. ⅓ of the kidney length. Auricle small,
triangular, located in middle length of kidney left margin.
Ventricle near same size of auricle. Kidney occupying c.
of pulmonary cavity length, located along middle and left
regions of transition pallial cavity and visceral mass. Kidney
rather bilobed, with smaller lobe just anterior to auricle.
Pulmonary vein running along columellar margin of pallial
cavity, straight in posterior  portion of the mantle, curved
in anterior  portion, converging to pneumostome region
along with renal tube and rectum.
Visceral mass (Figs2C, E)
Digestive gland brownish, located along inferior region of
seventh visceral whorl, covering part of stomach at sixth
whorl and  lling entire visceral whorls posterior to stomach,
including nepionic whorls (Fig.2E). Rectum visible at eighth
visceral whorl.
Genital system (Figs3A-F)
Gonad located at the sixth visceral whorl, covered by the diges-
tive gland (Fig.3E: go). Albumen gland elongated, sheltering
spermoduct under its columellar face. Carrefour lying at tran-

163
Rediscovery of Obeliscus agassizi
ZOOSYSTEMA • 2020 • 42 (12)
FIG. 1. — Obeliscus agassizi Pilsbry, 1906 shell and living specimen: A, SEM of apex in profi le; B, same, middle region of teleoconch; C, same, last whorl, frontal
view, asterisk showing epiphragm; D-F, living crawling specimen, shell length c. 6 mm; G, detail of suture of protoconch, SEM; H, detail of epiphragm insertion
on body whorl. Scale bars: A-C, 1 mm; G, 0.15 mm; H, 0.1 mm.
A
B
C
D
E
F
GH

164 ZOOSYSTEMA • 2020 • 42 (12)
D’ávila S. et al.
FIG. 2. — Obeliscus agassizi Pilsbry, 1906
anatomical drawings: A, outline of adult shell, part of read-foot also shown; B, soft parts last whorls, specimen just
extracted from shell; C, whole specimen view, pallial cavity extracted and extended to right; D, detail of shell fi rst whorls showing delicate sculpture; E, detail of
visceral mass. Abbreviations: see Material and methods. Scale bars: 2 mm.
AB
rt
ur
ur
mb
he
ki
ag
rt
pn
pn mb
cv
cv
pu
dg
ft
gp
rt
da
ur ki
cm
vd fo
ft
ve
pu
dg
st
rt
da
at
au
C
E
D

165
Rediscovery of Obeliscus agassizi
ZOOSYSTEMA • 2020 • 42 (12)
FIG. 3. — Obeliscus agassizi Pilsbry, 1906
anatomical drawings: A, outline of adult shell, part of read-foot also shown; B, soft parts last whorls, specimen just
extracted from shell; C, whole specimen view, pallial cavity extracted and extended to right; D, detail of shell fi rst whorls showing delicate sculpture; E, detail of
visceral mass. Abbreviations: see Material and methods. Scale bars: 2 mm.
ABC
D
E
F
ag
hd
so
pt
al
bc
vd
di pm
fo
om
fp
fo
vd
ey
om
pe
es
sg
so pt
vd
pe
pm
hd
dg
go
pe
om
gm
fo
ar
bc
vd
al
ag
sr
sr
ag
hd
gg
at

166 ZOOSYSTEMA • 2020 • 42 (12)
D’ávila S. et al.
sition between spermoviduct and albumen gland (Fig.3B).
Seminal receptacle as wide sinuous tube, lying at superior half
of albumen gland (Fig.3B: sr). Spermoviduct long, occupy-
ing  rst three shell whorls, running along columellar muscle
(Fig.3A, E). Autospermoduct situated at columellar face of
spermoviduct (Fig.3A: at). Allospermoviduct situated at the
opposite face, visible as a separated duct until reach the free-
oviduct (Fig.3A, E: al). Prostate with well de ned acini, oc-
cupying columellar surface of spermoviduct (Fig.3A, E: pt).
Bursa copulatrix about as wide as its duct (Fig.3E: bc), lying
in transition between spermoviduct and free oviduct; length
c.½ of that of spermovioduct. Free oviduct c.½ length of bursa
copulatrix duct. Vagina c.½ free oviduct length. Atrial retractor
muscle well developed. Vas deferens with greater diameter at
level of free oviduct, adhered to free oviduct wall at its more
convoluted and thicker portion through  ne muscle bundles.
Vas deferens meeting gland with unidenti ed function after
leaving prostate, running along entire free oviduct; vas def-
erens convoluted in median portion of free oviduct, showing
greater diameter until basis of free oviduct, joining to penis.
Penis long, club-shaped (Fig.3A, C: pe). At the meeting point
with the penial complex, the vas deferens forms a discrete bursa
which lies on the phallus wall. Penial complex passing under
right ommatophore and under esophagus. Penial retractor
muscle (Fig.3D: pm) originated in diaphragm region close
to columellar muscle, inserted terminally in the epiphallus.
ANNOTATED CHECKLIST OF SPECIES
OFOBELISCUS BECK, 1837
Family SUBULINIDAE Fischer & Crosse, 1877
Subfamily OBELISCINAE  iele, 1931
Genus Obeliscus Beck, 1837
Obeliscus abbotti Vanatta, 1918
Obeliscus abbotti Vanatta, 1918: 238,  g.3.
MUSEUM DEPOSITS.— Lectotype. Haiti • 1shell; near  e Generals
Cave, 1mile west of Jeremie; 1918; Abbott W.L. leg.; ANSP 118244.
Puerto Rico • 2shells; USNM 252558.
TYPE LOCALITY.— Haiti: General’s Cave, west of Jeremie. Found
in debris.
OTHER LOCALITIES.— Puerto Rico.
ILLUSTRATION.— Vanatta (1918:  g.3).
R
EFERENCES
.—  e only bibliographic reference is the original
description of the species.
Obeliscus agassizi Pilsbry, 1906
Obeliscus agassizi Pilsbry, 1906: 249, pl.36,  g.76.— Clench&
Turner 1962: 7.— Salgado& Coelho 2003: 155.— Simone 2006:
187,  g.686.
MUSEUM DEPOSITS.— Lectotype. Brazil • 1shell; Anthony J.G.
leg.; Agassiz expedition; ANSP 59334.
Paralectotypes. Brazil • 4shells; Anthony J.G. leg.; Agassiz expedi-
tion; ANSP 465075. Peru • 4shells; Chosica, North side of valley,
Dept. Lima; X.1964; Hocking P. leg.; FMNH 140941.
TYPE LOCALITY.— Brazil (collected by J.G. Anthony, during the
Agassiz exped.), no speci c locality.
O
THER
LOCALITIES
.— Brazil: Alagoas and Pernambuco states,
Biological Reserve of Pedra Talhada (present study). Peru: Chosica,
Nside of valley, Dept Lima.
ILLUSTRATION.— Pilsbry (1906: pl.36;  g.76)
R
EFERENCES
.— Clench& Turner (1962); Salgado& Coelho (2003);
Simone& Salvador (2016).
Obeliscus bacterionides (d’Orbigny, 1835)
Helix bacterionides Orbigny, 1835: 9.
Bulimus bacterionides – Pfei er 1852: 117, pl.35,  gs 15, 16.
Obeliscus bacterionides – Pilsbry 1906: 250, pl.36,  gs 71, 72.
M
USEUM
DEPOSITS
.— Syntypes. Bolivia • 6shells; Rio Grande; col-
lection d’Orbigny 1818-1838; MNHN-IM-2000-28012 • 2shells;
USNM 117487 • Beni Dept., Cercado Province, 6km South Trini-
dad; 28.XII.1982;  ompson F. leg.; FLMNH 040622• same data
as for preceding; FLMNH 040623 • Santa Cruz Dept., Chiquitos
Province, 20km ENE of Santiago de los Chiquitos; 30.XI.1982;
 ompson F. leg.; FLMNH 210580.
TYPE LOCALITY.— Bolivia: Rio Grande.
OTHER LOCALITIES.— Bolivia: no speci c locality (Pfei er, 1852);
Beni Department, Cercado Province, 6km South Trinidad (FLMNH
040622; 040623); Santa Cruz, Chiquitos Province, 20km North
East of Santiago de los Chiquitos (FLMNH 210580).
ILLUSTRATION.— Pfei er (1852: pl.35,  gs 15, 16); Pilsbry (1906:
pl.36;  g.71, 72).
IMAGING. — MNHN-IM-2000-28012, syntypes.
REFERENCES.— Pfei er (1852); Pilsbry (1906); Haas (1949).
Obeliscus blandi (Pfei er, 1852)
Bulimus blandi Pfei er, 1852: 85.
Obeliscus blandi – Pilsbry 1906: 257, pl.36,  gs. 83, 84.
TYPE LOCALITY.— Colombia: Baranquilla, in the Andes.
OTHER LOCALITIES.— Species known only from the type locality.
ILLUSTRATION.— Pilsbry (1906: pl.36,  gs83, 84).
REFERENCES.— Pilsbry (1906).
Obeliscus boitata Simone& Salvador, 2016
Obeliscus boitata Simone& Salvador, 2016: 25, 29 ( gs77-79).
M
USEUM
DEPOSITS
.— Holotype. Brazil • 1; Minas Gerais, Nanuque;
17°51’S, 40°23’W; 120m a.s.l.; very close to the borders with the
states of Bahia and Espírito Santo, very small fragment of Atlantic
Forest; MZSP 106169.

167
Rediscovery of Obeliscus agassizi
ZOOSYSTEMA • 2020 • 42 (12)
Paratypes. Brazil • 39shells; same collection data as for holotype;
MZSP 106170.
TYPE LOCALITY.— Brazil: Minas Gerais, Nanuque.
OTHER LOCALITIES.— Species known only from the type locality.
R
EFERENCES
.—  e only bibliographic reference is the original
description of the species.
Obeliscus carphodes (Pfei er, 1852)
Bulimus carphodes Pfei er, 1852: 108, pl.34,  g.2.
Obeliscus carphodes – Pilsbry 1906: 246, pl.34,  g.53.— Salgado&
Coelho 2003: 155.— Simone 2006: 187,  g.687.
M
USEUM
DEPOSITS
.— Brazil • 1shell; A.D. Brown Collection;
ANSP 59330 • 2shells; 1872; Heukelon F. van leg.; ZMA.MOLL
32528 • 1shell; RBINS 539204 • 1shell; RBINS 499929 • 1shell;
Rio Grande do “Sal” [Rio Grande do Sul], Albuquerque;  ayer
Exp. to Brazil; MCZ:Mala:119453.
TYPE LOCALITY.— Brazil: no speci c locality.
OTHER LOCALITIES.— Brazil (ANSP 59330; ZMA.MOLL 32528;
RBINS 539204; RBINS 499929; MCZ:Mala:119453); Rio Grande
do “Sal” [Rio Grande do Sul], Albuquerque (MCZ- 119453).
IMAGING.— ZMA.MOLL 32528.
I
LLUSTRATION
.— Pfei er (1852: pl.34,  g.2); Pilsbry (1906:
pl.34.,  g.53).
Obeliscus columella (Philippi, 1844)
Bulimus columella Philippi, 1844: 158 (pl.2,  g.7).
Obeliscus columella – Pilsbry 1906: 248, pl.34.,  gs49-50.— Sim-
one 2006: 187,  g.688.
MUSEUM DEPOSITS.— Brazil • 1shell; USNM 20276.
TYPE LOCALITY.— Brazil.
OTHER LOCALITIES.— Species known only from the type locality.
ILLUSTRATION.— Pilsbry (1906, pl.34,  gs49, 50).
REFERENCES.— Pilsbry (1906).
Obeliscus latispira Pilsbry, 1944
Obeliscus latispira Pilsbry, 1944: 127, pl.11,  g.18.— Clench&
Turner 1962: 84.
MUSEUM DEPOSITS.— Paratypes. Peru • 1shell; Junin Dept.,
Huacapistana; 1800m; 1942; Weyrauch W. leg.; ANSP 451843 •
Junin Dept., Huacapistana, river Tarma; FLMNH 110894 • same
collection data as for preceding; FLMNH 203092.
TYPE LOCALITY.— Peru: Huacapistana.
OTHER LOCALITIES.— Peru: Dept. Junin, River Tarma, Huacap-
istana (FLMNH 110894; 203092).
ILLUSTRATION.— Pilsbry (1944: pl.11,  g.18).
REFERENCES.— Clench& Turner (1962).
Obeliscus moderatus Pilsbry, 1933
Obeliscus moderatus Pilsbry, 1933: 152.— Clench& Turner 1962: 99.
MUSEUM DEPOSITS.— Dominican Republic • Barahora Province,
Sierra de Bahoruco, 4km North Polo; 12.IX.1983;  ompson
F. leg.; FLMNH 045777 • Barahora Province, Sierra de Baho-
ruco, 4km North Polo; 19.III.1974;  ompson F. leg.; FLMNH
045778 •Barahora Province, Sierra de Bahoruco, 5km North Polo;
19.III.1974;  ompson F. leg.; FLMNH 210623 • Barahora Prov-
ince, 7km NNE of Polo; 19.III.1974;  ompson F. leg.; FLMNH
210624 •Estrelleta Province; 9km North East of La Descubierta;
02.VI.1974;  ompson F. leg.; FLMNH 210625 • Barahona Prov-
ince; 2km NNE of Polo; 19.III.1974;  ompson F. leg.; FLMNH
210626 • Estrelleta Province, 6km NW of Postrer Rio; 02.VII.1974;
 ompson F. leg.; FLMNH 210627 • Independencia Province, 4km
North East of El Aguate; 30.VI.1974;  ompson F. leg.; FLMNH
210628 • Barahona Province, 14km South of Cabral, 7km NNE of
Polo; 18.VI.1974;  ompson F. leg.; FLMNH 210629 • Estrelleta
Province, 3km North East of El Aguacate; 30.VI.1974;  ompson
F. leg.; FLMNH 210630.
T
YPE
LOCALITY
.— Dominican Republic: Santo Domingo - Del
Monte plantation, 5-6min W of Barahona.
OTHER LOCALITIES.— Dominican Republic: Barahona Province,
Sierra de Bahoruco, 4km North of Polo (FLMNH 045777; 045778);
Barahona Province, 5km North East of Polo (FLMNH 210623);
Barahona Province, 7km North East of Polo (FLMNH 210624);
Estrelleta Province, 9km North East of La Descubierta (FLMNH
210625); Barahona Province, 9km North East of Polo (FLMNH
210626); Estrelleta Province, 6km NW of Postrer Rio (FLMNH
210627); Independencia Province, 4km North East of El Aguacate
(FLMNH 210628); Barahona Province, 14km of Cabral, 7km
North East of Polo (FLMNH 210629); Estrelleta Province, 3km
North East of El Aguacate (FLMNH 210630).
REFERENCES.— Clench& Turner (1962).
Obeliscus obeliscus (Moricand, 1833)
Helix (Cochlicella) obeliscus Moricand, 1833: 540 (pl.1,  g.4).
Helix obeliscus – Moricand 1836: 424.
Bulimus obeliscus – Pfei er 1852: 107 (pl.34,  g.1).
Obeliscus obeliscus – Pilsbry 1906: 245 (pl.34,  gs. 44-48, 51).—
Salgado& Coelho 2003: 155.— Simone 2006: 187 ( g.689).
MUSEUM DEPOSITS.— Syntypes. Brazil • 3shells; MCZ Malacol-
ogy 27337.
Brazil • 1shell; donor Brown, A.D.; ANSP 3064 • 3shells; donor
Phillips, J.S.; ANSP 58374 • 16shells; donor Philips, J.S. ANSP
58375 • 2shells; provenance Swift Collection Moricand; ANSP
58376 • 2shells; donor Webb, W. F.; ANSP 59329 • 1shell; Bahia;
provenance Naide, M.& B. Meyer; Naide Collection; ANSP 449608
• 1shell; Espírito Santo, Aracruz, Serra do Galo; Oct. 1992; prov-
enance I. Wistar Morris III, Hideo Katori Collection; ANSP 426505
• Pernambuco, Tapacura Dam; dry shell without animal; BMSM
96940 • Bahia, Canavieiras; under dead branches and leaves, dry
shell without animal; BMSM 96939 • 1shell; CMNH 62.14868
•Dautzenberg, P.; FLMNH 110900 • Espírito Santo; Foster M. leg.;
FLMNH 118822 • 2shells; FMNH 36160 • 1shell; FMNH 36161
• 3shells; FMNH 100631 • 1shell; RBINS 357518 • 1shell. RBINS
538334. • 1shell; RBINS 369515 • 1shell; RBINS 566776 • 1shell;
RBINS 480428 • 1shell; RBINS 575417 • 1shell; RBINS 438330
• 7shells; MCZ Malacology 27323 • 1shell; near Campos, Serra do

168 ZOOSYSTEMA • 2020 • 42 (12)
D’ávila S. et al.
Rio Preta [Serra do Rio Preto]; MCZ Malacology 27336 • 44shells;
MCZ Malacology 27344 • Rio Grande do Sal [Rio Grande do Sul],
Albuquerque; MCZ Malacology 119454 • 1shell; MCZ Malacology
135694 • Bahia, Canavieiras, Comunidade Puxin; 31.X.2011; shell,
specimen in ethanol 70%, slides, stubs for SEM, soft parts; MNRJ:
Mollusca: 30581 • 1; Sergipe; MNRJ: Mollusca: 15039 • 1shell; Bahia,
Porto Seguro, Parque Nacional do Pau Brasil; 17.VI.2009; specimen
in ethanol 70%; MNRJ: Mollusca: 30281 • Rio de Janeiro, Santa
Maria Madalena; 16.II.2008; Salgado N.C. leg.; MNRJ: Mollusca:
32851 • 12shells; Rio de Janeiro. Santa Maria Madalena, Rancho do
Lelê; -20.004588 -41.934722; 428elev.; 16.III.2008; Salgado N.C.
leg.; mata secundária; MNRJ: Mollusca: 13382 • Espírito Santo,
Paraisópolis, Sooretana; 19.IV.2006; Salgado N.C. leg.; MNRJ:
Mollusca: 31954 • 1shell; Bahia, Porto Seguro; 28.II.2005; MNRJ:
Mollusca: 31956. • 1; Bahia, Campo Formoso; IX.2002; specimen
preserved in ethanol 70%; MNRJ: Mollusca: 13219 • 4; Pernambuco;
São Lourenço; 16.VI.2001; Salgado N.C. leg.; specimens preserved
in ethanol 70%; MNRJ: Mollusca: 30399 • 8shells; Ceará, Camocin;
10.VII.1996; Salgado N.C. leg.; MNRJ: Mollusca: 30393 • 1; To-
cantins, Barrolandia, Mata da SEPLAC; 20.II.1985; specimen in
ethanol 70%; MNRJ: Mollusca: 18936 • 5; Espírito Santo, Sul de
Meaípe; 1.IX.1993; specimens preserved in ethanol 70%; MNRJ:
Mollusca: 18937 • 1shell; Bahia, Cajaiba; IV.1987; MNRJ: Mol-
lusca: 7758 • 1; Bahia, Belmonte, Barrolandia; 20.II.1985; Costa C.J.
F.leg.; specimens preserved in ethanol 70%; MNRJ: Mollusca: 31949
• 3shells; Bahia; V.1983; MNRJ: Mollusca: 42127 • 5shells; Rio de
Janeiro, Areia Branca; II.1977; MNRJ: Mollusca: 41611 • 10shells;
Bahia, Itajiba, Fazenda Pedra Branca; 23.I.1974; Costa C.J.F. leg.
MNRJ: Mollusca: 30296 • 1; Bahia, Itajiba, Fazenda Pedra Branca;
17.I.1975; Costa C.J. F. leg., specimens preserved in ethanol 70%;
MNRJ: Mollusca: 32347 • 3; Bahia, Itajiba, Fazenda Pedra Branca;
13.I.1972; specimens preserved in ethanol 70% MNRJ: Mollusca:
11364 •
Espírito Santo, Baixo Guandu; IX.1971; specimens preserved
in ethanol 70%; MNRJ: Mollusca: 11366 • 2shells; Bahia, Itajiba,
Pedra Branca; 05.I.1970; Costa C.J.F. leg.; MNRJ: Mollusca: 32346
• 1shell; Bahia, Matas do Novo Mundo; I.1970; MNRJ: Mollusca:
41354 • 1shell; Bahia, Santa Cruz Cabrália; Dec. 1970; Costa C.J.F.
leg.; MNRJ: Mollusca: 35423 • 2shells; Sergipe; Aug. 1962; Guerra
Jr O. leg.; MNRJ: Mollusca: 6702 • 5shells; Espírito Santo, Cupido,
Sooretama; 9.V.1960; Lopes H.S. leg.; MNRJ: Mollusca: 56453 •
9shells; Bahia, Fazenda Primavera; 1951; MNRJ: Mollusca: 53371 •
5shells; Espírito Santo, Rio Itaúnas; Faz. Boa Esperança; 22.VIII.1950;
MNRJ: Mollusca: 53370 • 5shells; Espírito Santo, Fazenda do Cu-
pido; 2.III.1947; MNRJ: Mollusca: 750 • 7shells; Espírito Santo,
Fazenda do Cupido; 2.III.1947; MNRJ: Mollusca: 756 • 8shells;
Bahia, Itapetininga; MNRJ: Mollusca: 42716 • 3shells; Alagoas, Ma-
ceió, Paripueira; Lopes H.S. leg.; MNRJ: Mollusca: 56062 • 1shell;
Espírito Santo; Salgado N.C. leg.; MNRJ: Mollusca: 31655 • shell;
RMNH.MOL.271447 (NBCN) • 1

shell; RMNH.MOL.271446
(NBCN) • 1shell; ZMA.MOLL.32527 (NBCN) • 2shells; USNM
57834 • 2shells; USNM 307450 • 2shells; USNM 20275. Colombia
• 1shell; USNM 251168. Ecuador • 1shell; USNM 426531. Brazil •
1 shell; USNM 316229. Mexico • 2shell; USNM 316353.Venezuela
• 1shell; MCZ Malacology 27324.
TYPE LOCALITY.— Brazil, Bahia, Caravelhas.
O
THER
LOCALITIES
.— Brazil: (HIST.106; 103; 105; 341); Tocantins,
Barrolândia (MNRJ: Mollusca: 18936); Ceará, Camocin (MNRJ:
Mollusca: 30393); Pernambuco, São Lourenço (MNRJ: Mollusca:
30399); Pernambuco, Tapacura Dam (BMNSM 96940); Alagoas,
Maceió, Paripueira (MNRJ: Mollusca: 56062); Sergipe (MNRJ:
Mollusca: 6702); Sergipe (MNRJ: Mollusca: 30581); Bahia (Pilsbry
1906); Bahia (ANSP 449608; Malacology 135694); Bahia, Bahia
(MNRJ: Mollusca: 42127); Bahia, Canasvieiras (BMSM 96939;
MZUSP 98140); Bahia, Canasvieiras, Comunidade Puxin (MNRJ:
Mollusca: 30581); Bahia, Itambé, Serra Campo Formoso ou Olho
d’Água (MZUSP 86775); Bahia, Porto Seguro (MNRJ: Mollusca:
31956); Bahia, Cajaíba (MNRJ: Mollusca: 7758); Bahia, Belmonte,
Barrolandia, Mata do Seplac (MNRJ: Mollusca: 31949); Bahia, Ita-
jiba, Fazenda Pedra Branca (MNRJ: Mollusca: 30296; 32347; 11364;
32346); Bahia, Matas do Mundo Novo (MNRJ: Mollusca: 41354);
Bahia, Santa Cruz Cabrália (MNRJ: Mollusca: 35423); Bahia, Fa-
zenda Primavera (MNRJ: Mollusca: 53371); Espírito Santo (ANSP
426505; FLMNH 118822; MNRJ: Mollusca: 31655); Espírito Santo,
Paraisópolis, Sooretana (MNRJ: Mollusca: 31954); Espírito Santo,
Meaípe (MNRJ: Mollusca: 18937); Espírito Santo, Baixo Guandu
(MNRJ: Mollusca: 11366); Espírito Santo, Rio Itaúnas, Fazenda Boa
Esperança (MNRJ: Mollusca: 53370); Espírito Santo, Fazenda do
Cupido (MNRJ: Mollusca: 750; 756); Espírito Santo, Itapetininga
(MNRJ: Mollusca: 42716); Rio de Janeiro, Santa Maria Madalena
(MNRJ: Mollusca: 32851; 13382); Rio de Janeiro, Macaé, Areia
Branca (MNRJ: Mollusca: 41611); near Campos, Serra do Rio Preto
(Malacology 27336); Rio Grande do Sul, Taguara (Pilsbry 1906);
Rio Grande do Sul, Albuquerque (Malacology 119454). Venezuela
(Malacology 27324). Colombia (USNM 251168). Ecuador (USNM
426531). Mexico (USNM 316229).
ILLUSTRATION.— Moricand (1833: pl.1;  g.4); Pfei er (1852:
pl.34,  g.1); Pilsbry (1906: pl.34,  gs44-48, 51).
REFERENCES.— Moricand (1836); Pfei er (1852); Pilsbry (1906);
Salgado& Coelho (2003).
Obeliscus pattalus Pilsbry, 1906
Obeliscus pattalus Pilsbry, 1906: 249, pl.36,  g. 70.— Clench&
Turner 1962: 115.— Salgado& Coelho 2003: 155.— Simone
2006: 188 ( g.690).
M
USEUM
DEPOSITS
.— Lectotype. Brazil • 1; 4.VI.1908; ANSP 59335.
Paralectotype. Brazil • 1; 21.VIII.2015; ANSP 465076.
TYPE LOCALITY.— Brazil, no speci c locality.
OTHER LOCALITIES.— Species known only from the type locality.
ILLUSTRATION.— Pilsbry (1906: pl.36,  g.70).
R
EFERENCES
.— Clench& Turner (1962); Salgado& Coelho (2003).
Obeliscus peregrinus (Pfei er, 1855)
Bulimus peregrinus Pfei er, 1855: 9.
Obeliscus peregrinus – Pilsbry 1906: 258.
TYPE LOCALITY.— Unknown.
OTHER LOCALITIES.— Unknown.
ILLUSTRATION.— Not  gured.
REFERENCES.— Pilsbry (1906).
Obeliscus planospirus (Pfei er, 1852)
Bulimus planospirus Pfei er, 1852: 108, pl.34,  g.3.
Obeliscus planospirus – Pilsbry 1906: 247, pl.34,  g.44.— Sal-
gado& Coelho 2003: 155.— Simone 2006: 187.
MUSEUM DEPOSITS.— Brazil • Espírito Santo, Rio Doce; ANSP
A9714 • 2shells; Espírito Santo, Rio Doce; 24.V.1910; ANSP
100535 • 1shell; CM: Mollusks 62.13330.

169
Rediscovery of Obeliscus agassizi
ZOOSYSTEMA • 2020 • 42 (12)
TYPE LOCALITY.— Unknown.
OTHER LOCALITIES.— Brazil: head waters of the Mucury river in
eastern Minas Gerais (Pilsbry, 1906).
Brazil: Espírito Santo, Rio Doce (ANSP100535).
I
LLUSTRATION
.— Pfei er (1852: pl.34,  g.3); Pilsbry (1906
pl.34;  g.44).
REFERENCES.— Pfei er (1852); Pilsbry (1906); Salgado& Coelho
(2003).
Obeliscus subuliformis (Moricand, 1836)
Helix subuliformis Moricand, 1836: 427, pl.2,  g.3.
Obeliscus subuliformis – Pilsbry 1906: 251, pl.36,  g.75.— Sal-
gado& Coelho 2003: 155.— Simone 2006: 188,  g.691.
MUSEUM DEPOSITS.— Paratypes. Brazil • 1shell; Bahia; Malacol-
ogy 177339 (MCZ) • 1; USNM 316229 • 1; USNM 117490 •
shell; RMNH.MOL.271464 (NBCN).
T
YPE
LOCALITY
.— Brazil: Bahia, São Gonçalves forest. Brazil: Bahia
(Malacology 177339,
MCZ).
OTHER LOCALITIES.— Species known only from the type locality.
ILLUSTRATION.— Pilsbry (1906: pl.36,  g.75).
REFERENCES.— Pilsbry (1906); Salgado& Coelho (2003).
Obeliscus sylvaticus (Spix& Wagner, 1827)
Bulimus sylvaticus Spix& Wagner, 1827: 11, pl.5,  g.4.
Obeliscus sylvaticus – Pilsbry 1906: 248, pl.34,  gs 54, 55.— Sal-
gado& Coelho 2003: 155.— Simone 2006: 188,  g.692.
M
USEUM
DEPOSITS
.— Brazil •
ZSM/MOL 110009 • 3; donor
Brown, A.D.; ANSP 3065 • 2; 4.VI.1908; donor Anthony, J.G.;
ANSP 59332 • 1; 4.VI.1908; donor Anthony, J.G.; ANSP 59333
• 4; Bahia, Bom Fim, Fazenda Ambroter; 22.XI.1907; Haseman
J.D. leg.; CMNH 62.16564 • 2; CMNH 62.14869 • 1; FMNH
103188 • 1shell; HIST.370 (RBINS) • 1shell; HIST.346 (RBINS)
• 1shell; HIST.110 (RBINS) • 1shell; MCZ 27322 • 1shell; MCZ
27321 • 3; USNM 117495 • 1; USNM 20275 • 1shell; ZMA.
MOLL.32567 (NBCN). French Guiana • 1; Cayenne; FMNH
36245 • 1; Cayenne; FMNH 36173 • 1; Cayenne; soft parts pre-
served in ethanol, dry shell; MB-A053789 (SANBI).
TYPE LOCALITY.— Brazil: Piauí, in woods.
OTHER LOCALITIES.— French Guiana: Cayenne (FMNH 36173;
MB-A053789).
ILLUSTRATION.— Spix& Wagner (1827: pl.5,  g.4); Pilsbry
(1906: pl.34;  gs54, 55).
REFERENCES.— Pilsbry (1906); Salgado& Coelho (2003).
Obeliscus virescens (Da Costa, 1898)
Stenogyra (Spiraxis) virescens Da Costa, 1898: 82, pl.6,  g.10.
Obeliscus virescens – Pilsbry 1906: 257, pl.36,  g.81.
TYPE LOCALITY.— Colombia: Valley of the Cauca River.
OTHER LOCALITIES.— Species known only from the type locality.
ILLUSTRATION.— Da Costa (1898: pl.6,  g.10); Pilsbry (1906:
pl.36,  g.81).
REFERENCES.— Pilsbry (1906).
DISCUSSION
HABITAT OF SPECIES OF OBELISCUS
Fragmentary information obtained from taxonomic literature
allows concluding that the species of the genus Obeliscus rather
dwell in leaf litter (Moricand 1836; Cousin 1887; Vanatta
1918). With reference to the habitat of Obeliscus sylvaticus,
for example, Moricand (1836: 10) said that this species is
very common in forests, under the trees, under dead and
humid leaves. According to Vanatta (1918: 238) Obeliscus
abbotti was found in debris. Some other species, as Obeliscus
bacterionides, have been found “under stones” (Pilsbry 1906).
Other obeliscine genera probably are leaf-litter-dwelling as well.
According to Cousin (1887: 239), Protobeliscus jousseaumei
(Cousin, 1887) was found buried in the humus or hide under
dead leafs. According to Hylton-Scott (1946: 366), Rectobelus
birabeni was collected in the leaf-litter.
DISTRIBUTION OF OBELISCUS IN BRAZIL
Information on the distribution of Obeliscus species from
Brazil is very scant. For some species as Obeliscus carphodes,
Obeliscus pattalus, Obeliscus agassizi and Obeliscus columella,
the type locality was given simply as Brazil (Pfei er 1852;
Pilsbry 1906).  e information available in the literature
on the distribution of the other species is also very limited.
Obeliscus obeliscus was recorded for Bahia and Rio Grande
do Sul states and Obeliscus sylvaticus for Piauí, Bahia and Rio
de Janeiro states. Obeliscus subuliformis and Obeliscus boitata
are known only for their type localities at Bahia and Minas
Gerais states, respectively (Spix& Wagner 1827; Moricand
1833; 1836; Pfei er 1852; Simone& Salvador 2016). Ob-
eliscus planospirus was described without reference to the type
locality and, later, Pilsbry (1906) added “head waters of the
Mucury river in eastern Minas Gerais, Brazil”.
In the present study, the research in collection databases
allowed to expand the recognized distribution of the species
O.planospirus, which is now recorded in Espírito Santo state,
O.obeliscus now recorded in Tocantins, Ceará, Pernambuco,
Alagoas, Sergipe, Espírito Santo and Rio de Janeiro states, as
well as O.sylvaticus now recorded in French Guiana, besides
Brazil.
LIFE HISTORY TRAITS OF OBELISCUS
Although there is no study on the life history of species of
Obeliscus, fragmentary data on reproduction can be found in
the taxonomic literature. Some information given by Moricand
(1836: 11, pl.2,  g.28) and rea rmed by Pilsbry (1906:
246) on O.obeliscus indicate that this species is ovoviviparous,

170 ZOOSYSTEMA • 2020 • 42 (12)
D’ávila S. et al.
releasing non-calci ed eggs formed by a thin and transpar-
ent membrane containing shelled embryos. About Obeliscus
swiftianus (Pfei er, 1852), Pilsbry (1906) a rmed: “…none
of the specimens seen contains eggs, such as are commonly
seen in Opeas. It is probably viviparous, like all Obelisci.”
Since this author mentioned only oviparity and viviparity,
discriminating viviparous species as those containing shelled
embryos in the spermoviduct, we believe that this species is
in fact ovoviviparous. Ovoviviparity and egg retention seem
to be widespread reproductive modes in subulinids (Pilsbry
1906; Baker 1927; Batts 1957; Araújo& Keller 1993; Schi-
leyko 1999; Carvalho etal. 2009; Pilate 2013) and may be
common to obeliscine snails (Moricand 1836; Pilsbry&
Vanatta 1899; Pilsbry 1899, 1906).
TAXONOMIC REMARKS
Schileyko (1999) estimated about 10species of Obeliscus.
In the present work, we listed 16species of Obeliscus, nine
of them recorded in Brazil. Among the species occurring in
Brazil, six are endemic to this country.
 e species of Obeliscus have been traditionally distinguished
by shell characters, mainly the general shell shape, number of
whorls proportionally to the shell size, shell ornamentation,
the shape of the protoconch, and aperture size and shape.
 e distinction between species is often subtle. Considering
that shell features may be not su cient as single taxonomic
characters, other evidences of species boundary, especially
anatomical characters, is needed.
Obeliscus agassizi and O.sylvaticus were found by Pilsbry
(1906) cohabiting in the same collecting site.  is fact, along
with the close resemblance in their shells, indicates the need
to provide a di erential diagnosis for these species, through
a more exhaustive anatomical characterization. Currently,
the two species can be di erentiated mainly by the shape of
the shell aperture. Obeliscus sylvaticus has a more slender shell
with a smaller aperture, which is also quadrangular in shape,
while the shell aperture of O.agassizi is somewhat oblique
and ovate in shape.  e strength of shell traits to delimit
species of Obeliscus should be evaluated in the future, with
the accumulation of information on the anatomy of the soft
parts and with the use of molecular markers.
Presently it is not possible to evaluate the taxonomic sig-
ni cance of the various morphological traits of the soft parts
for the Obeliscinae genera as there is information only on
four species from four genera.  e data on the anatomy of
the genital system of O.agassizi (present study), Neobeliscus
calcareus provided by Pilsbry (1899, 1906), Stenogyra ter-
ebraster provided by Schileyko (1999) and Rectobelus birabeni
provided by Hyton-Scott (1946), did not allow a comprehen-
sive comparative analysis, but some anatomical di erences
among the representatives of these four genera are worth of
mentioning. Obeliscus agassizi di ers from N.calcareus in the
site of insertion of the vas deferens into the penial complex,
the site of insertion of the bursa copulatrix duct into the free
oviduct, the site of insertion of the penial retractor muscle
and the size of the bursa copulatrix duct relative to the size
of the free oviduct. Obeliscus agassizi di ers from R.birabeni
in the site of insertion of the vas deferens into the penial
complex, the site of insertion of the penial retractor muscle
and the size of the bursa copulatrix duct relative to the size of
the free oviduct. Obeliscus agassizi di ers from S.terebraster
in the site of insertion of the bursa copulatrix into the free
oviduct and in the site of insertion of the vas deferens into
the penial complex.
Considering the anatomical characteristics common to
the genera Obeliscus, Neobeliscus, Stenogyra, some traits are
possible diagnostic characters for this subfamily.  is con-
cerns characters of the genital system, i.e., atrium and vagina
short; long, club-shaped penial complex and terminal penial
retractor muscle; of the circulatory and excretory systems,
i.e., venation faint; of the respiratory system, i.e., pulmonary
vein unbranched; and of the excretrary system, i.e., kidney
more than double the length of the pericardium, and kidney
length contained about 3½ times in that of the mantle.  e
genus Rectobelus Baker, 1927, share with the genera Obelis-
cus, Neobeliscus, Stenogyra the presence of short atrium and
vagina, and long, club-shaped penial complex.  is genus
must to be reexamined since there is no description of the
circulatory and excretory systems.  e penial complex of
R.birabeni is not long and club-shaped as observed for the
other three genera.
 is preliminary analysis gives an idea of potential diag-
nostic characters, but it is still necessary to evaluate if they
are consistent in examining a great number of species from
these and the other Obeliscinae genera, along with the intra
generic variability on these traits.
Author’s contributions
SD conceived and designed the study, performed anatomical
and taxonomic analysis, original illustrations and wrote this
manuscript. LRLS edited the plates and co-wrote this manu-
script. LFCO performed shell SEM analysis and revised this
manuscript. CL collected the studied specimens and revised
this manuscript. PM collected the studied specimens and
co-wrote this manuscript.
Competing interests
 e authors declare that there is no con ict of interests re-
garding the publication of this paper.
Acknowledgements
 e authors would like to thank the Post-graduate program
“PPGCB-Comportamento e Biologia Animal – UFJF”, Juiz
de Fora – Brazil, for the travel fund grant conceded to SD.
 e authors are very grateful to Philippe Bouchet for provid-
ing access to ethanol-preserved specimens of Obeliscus agas-
sizi deposited in the malacological collection of the Muséum
national d’Histoire naturelle. We also acknowledge the two
anonymous reviewers for their invaluable suggestions.  e
 eldwork was made possible through the ICMBio permit
(Sisbio #48925-2), requested by Eduardo Colley. We are also

171
Rediscovery of Obeliscus agassizi
ZOOSYSTEMA • 2020 • 42 (12)
grateful to the NGO “Nordesta” (Switzerland), and particu-
larly to its President Anita Studer, who covered the travel and
accommodation of two of the authors.  anks to Louis Nus-
baumer, Curator of the “Conservatoire et Jardin botaniques
de la ville de Genève”, (Switzerland) for his help on the  eld.
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Submitted on30 November 2018;
accepted on 11 June 2019;
published on 21 April 2020.