Research Article |
Corresponding author: Maria Gabriela Cuezzo ( gcuezzo@webmail.unt.edu.ar ) Academic editor: Rosana Rocha
© 2017 Maria Gabriela Cuezzo, Meire Silva Pena.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Gabriela Cuezzo M, Pena MS (2017) Minaselates, a new genus and new species of Epiphragmophoridae from Brazil (Gastropoda: Stylommatophora: Helicoidea). Zoologia 34: 1-12. https://doi.org/10.3897/zoologia.34.e13230
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We describe a new genus and a new species in the family Epiphragmophoridae, Minaselates paradoxa sp. n. The new species was found at the National Park Cavernas do Peruaçu, in northern portion of the state of Minas Gerais, Brazil. Minaselates paradoxa sp. n. is classified in Epiphragmophoridae based on the fact that it shares the following diagnostic features of the family: a dart apparatus with a single dart sac, and two unequal mucous glands at the terminal genitalia. Minaselates gen. n. differs from Epiphragmophora Doering, 1874 by having a granulose protoconch, shell spire with blunt apex, complex microsculpture on the teleoconch and closed umbilicus fused with the shell wall. Also, significant differences between the two genera are the presence of a long and thin kidney that extends more than half the length of the pulmonary cavity, the presence of a flagellar caecum, and a smooth jaw in Minaselates gen. n. The finding of this new species and genus is particularly significant to refine the definition of the family, since Epiphragmophoridae has been traditionally diagnosed using the same characters of Epiphragmophora. Dinotropis Pilsbry & Cockerell, 1937, the other valid genus in the family, is monospecific and is only known by the morphology of the shell. In many ways it is similar to Epiphragmophora. A cladistics analysis was made in the present study which supports Minaselates gen. n. as a different entity and as sister group of the Epiphragmophora within Epiphragmophoridae.
Cerrado, Pleurodontidae , Pulmonata , South America, Taxonomy
Epiphragmophoridae is a Pulmonate land snail family exclusively distributed in South America. It is composed of the genera Epiphragmophora Doering, 1874 and Dinotropis Pilsbry & Cockerell, 1937. Epiphragmophora is currently composed of 63 species distributed in Peru, Bolivia, Argentina, Paraguay and southern Brazil with a single extra occurrence in Colombia (
Epiphragmophoridae is currently diagnosed by the same synapomorphies of Epiphragmophora because Dinotropis is only known by its original description, which is entirely based on shell characters. Based on a cladistic hypothesis (
Epiphragmophora was classified by
During a field trip to the National Park Cavernas do Peruaçu in northern Minas Gerais, Brazil, to collect gastropods, a striking group of land snails was found. Analysis of the specimens collected revealed that they represent a new species of the family Epiphragmophoridae. The objective of the present work is to describe the new species in a new genus of Epiphragmophoridae and to discuss its position among the South American Helicoidea.
Snails were collected at the National Park Cavernas do Peruaçu (14°56’S, 44°36’W) located in the state of Minas Gerais, Brazil. This conservation unit was created in 1999 with 143,353.84 ha (http://www.icmbio.gov.br), to protect limestone caverns. The calcareous massif is covered by rare and typical deciduous (Caatinga) or semi-deciduous forests called Seasonal Dry Tropical Forest (SDTF) (
Live specimens and dry shells of Minaselates paradoxa sp. n. were collected from rocky outcrops in dry deciduous forests of the National Park. Dry shells adhered to rocks or to leaves were abundant but live snails were scarce. The collected specimens were drowned in water for relaxation previous to fixation in ethanol 96%. Their shells were then photographed and measured using the software ImageJ 1.49 (Fig.
Intitutional abbreviations used in the text: IBN, Instituto de Biodiversidad Neotropical, Tucumán, Argentina; MLP-Ma, Museo de La Plata, Buenos Aires, Argentina; MNRJ, Museu Nacional Rio de Janeiro, Universidade Federal do Rio de Janeiro, Brazil; MCN, Museu Ciências Naturais, Pontifícia Universidade de Minas Gerais, Belo Horizonte, Brazil.
For the cladistic analysis, a matrix of 35 characters from the general anatomy plus shell morphology was generated for 24 terminal taxa (Appendix 1), following characters and codifications of
Shell measurements. Abbreviations (AD) dorsal view area, (Aap) apertural area, (Abw) body whorl area, (AL) lateral view area, (Dap) apertural diameter, (DM) major diameter, (Dm) minor diameter, (H) total shell height, (Hap) height of aperture, (Hbw) body whorl height, (Pbw) body whorl perimeter, (PD) dorsal view shell perimeter, (PL) lateral view shell perimeter, (PS) parietal space.
Supra superfamily classification follows
Minaselates gen. n. is distinguished by the following characters: 1) shell globose with blunt apex; 2) protoconch sculptured with granules; 3) teleoconch sculptured with complex microstructures; 4) umbilicus imperforate, parietal wall fused with columellar zone of peristome; 5) wavy spiral lines below the periphery and over ventral teleoconch surface; 6) genitalia with a dart apparatus composed by a single dart sac and two unequal mucous glands, one globose and the other oval; 7) presence of a flagellar caecum; 8) bursa copulatrix duct short, no longer than the sac.
Minaselates paradoxa sp. n. by original designation.
Shell globose, with 4 to 5 convex whorls. Spire conic with blunt apex. Protoconch granulose. Teleoconch sculptured. Wavy spiral grooves at the ventral teleoconch surface. Aperture subcircular with thin peristome. Umbilicus closed. Presence of spiral brownish bands more pronounced in the body whorl. Kidney long and thin, more than half the lung roof length. Genitalia with a dart apparatus and two unequal mucous glands.
Minaselates is a compound name formed by Minas in honor to the Brazilian state where the species was found, and selates, a noun in the genitive singular, that derives from the Greek meaning “snail” (
Minaselates gen. n. is classified in Epiphragmophoridae because it has a dart apparatus and two unequal mucous glands at the terminal genitalia. These structures are diagnostic of Epiphragmophoridae (Helicoidea) and their morphology serve to differentiate this family from the remaining helicoidean groups. Dinotropis differs from Minaselates in its depressed shell with an acute peripheral keel and open umbilicus. Minaselates differs from Epiphragmophora in its general shell shape with blunt apex, granulose protoconch and complex sculpture of the teleoconch surface. The wavy spiral grooves at the ventral teleoconch surface in Minaselates are lacking in both, Epiphragmophora and Dinotropis. The presence of a long and thin kidney in Minaselates is very different to the kidney shape in Epiphragmophora, which is triangular and shorter.
Shell globular, with three spiral continuous pigmented bands, the middle, equatorial band thinner. Protoconch granulose. Dorsal side of teleoconch with axial lines bearing triangular lamellae, ventral teleoconch with wavy, concentric, spiral grooves. Imperforate umbilicus. Jaw smooth. Kidney triangular, long and thin, of about 60 to 70% the length of the lung roof. Vas deferens insertion in lower portion of flagellar caecum. Strong, short muscular penial retractor inserting at proximal epiphallus.
The species name derives from the Greek paradoxos meaning “strange, contrary to expectation” (
Shell (Figs
Brazil, Minas Gerais: Itacarambi, National Park Cavernas do Peruaçu, Vale dos Sonhos (523m, X = 0599645, Y = 8343426), M.S. Pena, A. Suhett, D.C. Souza leg., December 2010, (MNRJ 34.580), Holotype (ethanol preserved specimen).
Brazil, Minas Gerais: Itacarambi, National Park Cavernas do Peruaçu, Nossa Senhora Aparecida Farm (532 m, X = 0589284, Y = 8328970), M.S. Pena, A. Suhett, D.C. Souza leg., December 2010, (IBN 21-S, MLP-Ma 14216), dry shells, (IBN 861), ethanol preserved specimens. Paratypes. Brazil, Minas Gerais: Itacarambi, National Park Cavernas do Peruaçu, Brejal, Peruaçu River side (663 m, X = 0579404, Y = 8332170), (MCN 192), dry shells. Brazil, Minas Gerais: Itacarambi, Natiomal Park Cavernas do Peruaçu, Janelão Cave (714 m, X = 0581514, Y = 8329046) M.S. Pena, A. Suhett, D.C. Souza leg., December 2010, (MCN 208).
Thus far known only from National Park Cavernas do Peruaçu, northern region of Minas Gerais, Brazil.
Minaselates differs from all known species of Epiphragmophora by having a granulose protoconch, the shell spire with a blunt apex, and by the wavy, spiral grooves in the ventral region of the shell. The fused, imperforate umbilicus on the shells of this new species is not typical of Epiphragmophora. Most of species of the genus present an open, perspective umbilicus. The exceptions are E. argentina (Holmberg, 1909) and some specimens of E. variegata Hylton Scott, 1962. The presence of wavy spiral grooves at the base of the shell of M. paradoxa sp. n. is noteworthy, sharply contrasting with the condition found in all other species of Epiphragmophora, where it is absent, except for E. (Pylsbrya) farrisi (Pfeiffer, 1859), which has shallow spiral lines. Minaselates paradoxa sp. n. also differs from the species classified in Epiphragmophora in the shape and length of the kidney, and the smooth jaw. In the species of Epiphragmophora for which the anatomy has been studied, the kidney is no more than half the length of the pulmonary roof, while the jaw is ribbed. A noteworthy character present in Minaselates paradoxa sp. n. is the presence of a flagellar caecum, a structure not found in Epiphragmophora. In this new species, the vas deferens inserts in the lower portion of the caecum. The penial retractor muscle in Epiphragmophora is mostly long and thin, inserting in the epiphallus, while in Minaselates paradoxa sp. n. this muscle is stronger, inserting in the caecum. M. paradoxa sp. n. is isolated from the area of distribution of Epiphragmophora, whose area of highest species richness is in the western portion of South America.
Minaselates paradoxa sp. n. resembles some species of Pleurodontidae by the presence of complex structures in the terminal genitalia of the male, such as the flagellar caecum. It is also similar to some Pleurodontidae in its long and thin kidney, the crowded granules in the surface of the shell protoconch, the globular general shape of the shell, the complex microsculpture on the teleoconch and in its smooth jaw. The presence of a flagellar caecum is noteworthy in Minaselates, this structure being absent in Epiphragmophoridae, while it is characteristic of some Pleurodontidae such as Polydontes Montfort, 1810 and Pleurodonte incerta (Férussac, 1823) (
In Minaselates, however, the vas deferens inserts in the flagellar pouch while in Polydontes the vas deferens inserts above this caecum. At first glance, the shell of this new species is very similar to some species of Pleurodonte (Pleurodontidae), except for the presence of concentric sculpture below the periphery, and in the basal area. It also differs in the morphology of the terminal genitalia that has a dart complex and a different flagellum shape. In Pleurodonte, the vas deferens is twisted around the epiphallus, descending to the peni-oviducal angle, while in M. paradoxa sp. n. the vas deferens runs straight, parallel to the penis complex, without looping around the epiphallus. Most of the Helicoidean families have a ribbed jaw or ‘odontognath jaw’, except for the Sagdidae with a ‘stegognath jaw’ and the Sphicterochilidae and Cepoliinae with a smooth jaw (‘oxygnath jaw’) (
Minaselates paradoxa sp. n. is distributed within a National Park area where Cerrado and Caatinga are the dominant biomes. These are considered high diversity hotspot areas. Within these hotspots areas specimens were collected in typical deciduous or semi-deciduous forests called Seasonally Dry Tropical Forests (SDTF) (
Shell dimensions in mm or mm2 (n = 14). DM major diameter, Dm minor diameter, AD dorsal area, PD dorsal perimenter, H total height, HBw body whorl height, AL lateral area, PL lateral perimeter, ABw body whorl area, PBw body whorl perimeter, Dap diameter of the aperture, Hap height of the aperture, EP length parietal space, AAp apertural area, Pap apertural perimeter, DP penultimate whorl diameter, DPr protoconch diameter (see Fig.
Character | Mean | SD | Min | Max | Holotype |
---|---|---|---|---|---|
DM | 28.526 | 0.765 | 27.103 | 29.714 | 28.934 |
Dm | 25.582 | 0.669 | 24.182 | 26.374 | 26.058 |
AD | 557.304 | 30.211 | 502.938 | 596.790 | 566.908 |
PD | 85.289 | 2.375 | 80.699 | 88.531 | 86.135 |
H | 18.663 | 1.005 | 17.183 | 20.370 | 19.375 |
HBw | 15.068 | 0.878 | 13.904 | 16.351 | 15.904 |
AL | 375.374 | 30.818 | 324.118 | 411.659 | 386.807 |
PL | 74.999 | 3.084 | 69.055 | 78.120 | 76.548 |
ABw | 343.840 | 28.940 | 299.433 | 379.724 | 355.669 |
PBw | 74.081 | 2.727 | 69.154 | 77.544 | 74.445 |
Dap | 14.886 | 1.091 | 12.997 | 16.888 | 15.004 |
Hap | 14.937 | 0.598 | 13.647 | 15.763 | 15.158 |
PS | 11.382 | 0.610 | 10.245 | 12.335 | 11.158 |
AAp | 190.094 | 14.988 | 158.663 | 208.360 | 208.360 |
PAp | 52.650 | 2.173 | 49.201 | 56.690 | 53.222 |
DP | 16.605 | 0.874 | 14.900 | 17.782 | 17.782 |
DPr | 3.050 | 0.264 | 2.651 | 3.550 | 2.873 |
Shell morphology: (8) Protoconch in dorsal view, scale bar = 4mm; (9) lateral view of the protoconch and first whorls, scale bar = 4mm; (10) general view of the teleoconch microsculpture, scale bar = 100μ; (11) body whorl microsculpture consisting on axial rows of triangular lamella separated by wrinkles; (12) perpendicular view of the lamella, scale bar = 5μ; (13) detail of a triangular lamellae showing its central axis, scale bar = 2 μm.
Digestive system: (14) Dorso-lateral view of the radula, arrows points to central tooth in two transverse rows of teeth; (15) detail on a dorso-lateral view of the lateral teeth;(16) detail of lateral teeth close to margin in dorsal view (17) margin of the radula showing curve shaped marginal teeth; (18) dorsal view of the smooth, fragile jaw; (19) detail of the dorsal surface of the jaw showing transversal shallow grooves. Scale bars = 10 μm.
Pallial system: (20) photograph of the ventral zone of the pulmonary roof, note the long kidney, with respect to the total length of the lung; (21) line drawing of the same region detailing the limits between kidney and ureter and the shallow veins crossing the pulmonary roof. Scale bar: 21 = 5mm. Abbreviations (k) kidney, (mc) mantle collar, (pv) pericardic vein, (r) rectum, (su) secondary ureter.
Genital system: (22, 23) line drawing and photograph of the complete dissected out reproductive system (ag) albumen gland, (bc) bursa copulatrix, (ds) dart sac, (ec) flagellar caecum, (fl) flagellum, (go) genital opening, (hd) hermaphroditic duct, (mg1) mucous gland 1, (mg2) mucous gland 2, (p) penis, (pr) penial retractor muscle, (s) spermoviduct, (vd) vas deferens; (24) detail of the terminal genitalia (pl) penial plates, (ps) penial sheath, (v) verge; (25) vas deferens and insertion of mucous gland ducts into dart sac (mgd) mucous gland duct; (26) inner sculpture of the penial complex showing the verge inside the penis. Scale bars: 22 and 25 = 5 mm.
Consensus tree obtained from two trees under implied weight approach highlighting the position of Minaselates gen. n. Only taxa with described anatomy were included in the matrix. Numbers below branches are node numbers. Symmetric resampling values (left number) and jackknife values (right number) are located above branches. Only values above 50% are illustrated.
The main goal of the analysis performed was to evaluate and support the description of a new genus. For this, only the species of Epiphragmophora with known complete anatomical and shell morphology information were used. The morphological data set analyzed under the implied weights approach resulted in two most parsimonious trees. The resulting strict consensus tree is illustrated (Fig.
Epiphragmophora (node 29) was also recovered as a monophyletic genus in all optimal trees, supported by the following synapomorphies: Body whorl surface with axial ribs regularly distributed (0:3); umbilicus overlapped but not solded to body whorl (1: 1); protoconch smooth (8:0), and Penial retractor muscle inserting in medial zone of epiphallus (21:1). Within Epiphragmophora, the clade [E. variegata [E. hemiclausa [E. tucumanensis+E. argentina]]] (node 27) has the highest SR and jackknife support and resulted monophyletic in all trees obtained.
Minaselates gen. n. is the sister group of Epiphragmophora in both optimal trees. This relationship is supported by the following synapomorphies: Presence of mucous glands in terminal genitalia (10: 1); presence of a dart apparatus (11:1); medium to short length of the bursa copulatrix duct (22:01) and a finger-like short to medium flagellum (25:1).
Thanks to the Instituto Chico Mendes de Conservação da Biodiversidade for the working permits and support (SISBIO-ICMBIO 19133-1 de 08/04/2009) and Fundo de Incentivo à Pesquisa (FIP), Pontifícia Universidade Católica de Minas Gerais for financial support to MSP. MGC is a researcher of the Argentine National Council for Scientific Research (CONICET). Financial support to MGC has been received through PIP 0055 (CONICET). We also thank the anonymous reviewers and the editor for their valuable comments.
Character matrix used in cladistic analysis. Only taxa with described anatomy were included in the matrix. Character codification according to
Characters | |||||||||||||||||||||||||||||||||||
1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | 10 | 1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | 20 | 1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | 30 | 1 | 2 | 3 | 4 | ||
Pleurodonte | 4 | 0 | 0 | 1 | 0 | 0 | 0 | 1 | [01] | 1 | 0 | 0 | – | – | – | – | – | – | – | 0 | 1 | 2 | 2 | 1 | 0 | 2 | 0 | 0 | [01] | – | 0 | 0 | 0 | 0 | 0 |
E. argentina | 2 | [03] | 0 | 1 | 0 | 1 | 0 | 1 | 0 | 0 | 1 | 1 | 1 | 0 | 1 | 0 | 1 | 0 | 0 | 1 | 0 | 1 | 2 | 1 | 1 | 0 | 1 | 0 | 1 | 1 | 0 | 0 | 0 | 0 | 0 |
E. cryptomphala | 2 | [01] | [12] | 1 | 0 | 1 | 0 | 1 | 0 | 0 | 1 | 1 | 0 | 1 | 1 | 1 | 1 | 1 | 1 | 0 | 1 | 1 | 1 | 0 | 1 | 1 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 |
E. escoipensis | 0 | 1 | 1 | 0 | 0 | [01] | 0 | 1 | 0 | 2 | 1 | 1 | 0 | 1 | 1 | 1 | 0 | 1 | 1 | 0 | 0 | 1 | 1 | 0 | 0 | 1 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 |
E. hemiclausa | 2 | 1 | 0 | 1 | 0 | 1 | 0 | 1 | 0 | 2 | 1 | 1 | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 1 | 1 | 2 | 1 | 0 | 1 | 1 | 0 | 1 | 1 | 0 | 0 | 0 | 0 | 0 |
E. guevarai | 0 | 2 | 0 | 0 | 0 | 1 | 0 | 1 | 0 | 2 | 1 | 1 | 0 | 1 | 1 | 1 | 0 | 0 | 1 | 2 | 1 | 1 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 |
E. hieronymi | 0 | 2 | 0 | 0 | 0 | [01] | 0 | 0 | 0 | 0 | 1 | 1 | 0 | 1 | 1 | 1 | 0 | 1 | 1 | 1 | 1 | 2 | 0 | 0 | 1 | 1 | 1 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 |
E. jujuyensis | 2 | 1 | 1 | 1 | 1 | 1 | 0 | 1 | 0 | 0 | 1 | 1 | 0 | 1 | 1 | 0 | 1 | 0 | 0 | 2 | 1 | 1 | 0 | 0 | 1 | 1 | 0 | 0 | 1 | 1 | 1 | 1 | 0 | 0 | 0 |
E. parodizi | 3 | 1 | 0 | 1 | 0 | 1 | 0 | 1 | 0 | 0 | 1 | 1 | 0 | 1 | 1 | 1 | 0 | 1 | 1 | 2 | 0 | 0 | 1 | 0 | 0 | 1 | 0 | 0 | 1 | 0 | 1 | 0 | 0 | 0 | 0 |
E. puella | 4 | 2 | 0 | 2 | 0 | 0 | 1 | 0 | 0 | 0 | 1 | 1 | 0 | ? | ? | ? | 0 | 1 | ? | ? | ? | ? | 0 | ? | ? | 1 | ? | ? | 0 | 0 | 1 | ? | 0 | 0 | 0 |
E. puntana | 3 | 1 | 1 | 0 | 0 | 1 | 0 | 1 | 0 | 0 | 1 | 1 | 0 | 1 | 1 | 1 | 1 | 1 | 1 | 2 | 1 | 1 | 1 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
E. quirogai | 0 | 2 | 0 | 0 | 0 | [01] | 0 | 1 | 0 | 2 | 1 | 1 | 0 | 1 | 1 | 1 | 0 | 1 | 1 | 0 | 1 | 2 | 0 | 0 | 1 | 1 | 0 | 0 | 0 | 1 | 1 | 1 | 0 | 0 | 0 |
E. rhathymos | 2 | 3 | 0 | 1 | 0 | 1 | 0 | 1 | 0 | 0 | 1 | 1 | 0 | 1 | 1 | 1 | 0 | 1 | 1 | 1 | 0 | 1 | 1 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 |
E. saltana | 3 | 1 | 1 | 1 | 0 | 1 | 0 | 1 | 0 | 0 | 1 | 1 | 0 | 1 | 1 | 1 | 0 | 1 | 0 | 1 | 0 | 1 | 1 | 0 | 0 | 1 | 0 | 1 | 1 | 0 | 0 | 0 | 0 | 0 | 0 |
E. tomsici | 2 | 1 | 1 | 1 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 1 | 0 | 1 | 1 | 1 | 0 | 1 | 0 | 1 | 0 | 0 | 1 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 1 | 0 | 0 | 0 | 0 |
E. trenquelleonis | 0 | 1 | 0 | 0 | 0 | 1 | 0 | 1 | 0 | 0 | 1 | 1 | 0 | 1 | 1 | 1 | 0 | 1 | 1 | 2 | 1 | 1 | 0 | 2 | 1 | 1 | 0 | 0 | [01] | 0 | 1 | 0 | 0 | 0 | 0 |
E. trifasciata | 1 | 2 | 0 | 0 | 0 | 1 | 0 | 1 | 0 | 0 | 1 | 1 | 0 | 1 | 1 | 1 | 0 | 0 | 1 | 2 | 1 | 1 | 0 | 0 | 1 | 1 | 0 | 0 | 0 | 0 | 1 | 0 | 0 | 0 | 0 |
Character list used in the cladistics analysis of species of Epiphragmophora with known anatomy plus the new genus and species, Minaselates paradoxa sp. n.
0. Body whorl surface: with thin growth lines = 0; with thick growth ridges = 1; malleated with diagonal ribs = 2; with axial ribs regularly distributed = 3; pustules/granules to wrinkles = 4; triangular lamella in axial rows = 5. [additive].
1. Umbilicus: Fused with basal lip of peristome = 0; overlapped but not fused to body whorl = 1; perspective wide not overlapped by peristomal lip = 2; perspective narrow slightly overlapped = 3; wide partly overlapped = 4.
2. Shape of the aperture: sub circular = 0; oval horizontal = 1; sub quadrangular = 2.
3. Peristome: Thin expanded slightly reflexed = 0; thick wide reflexed = 1; thin highly expanded = 2.
4. Basal callus in peristome: absent = 0; present = 1.
5. Peripheral bands: absent = 0; present = 1.
6. Body whorl periphery: convex = 0; equatorially subcarinated = 1; supraequatorially subcarinated = 2; carinated = 3. [additive].
7. Aperture respect to body whorl: not descending = 0; descending = 1.
8. Protoconch sculpture: smooth = 0; granulose = 1.
9. Spire apex: pointed = 0; dull = 1; not evident = 2.
10. Mucous glands in terminal genitalia: absent = 0; present = 1.
11. Dart apparatus: absent = 0; present = 1.
12. Shape of dart sac: long finger-like usually with constriction = 0; short, cylindrical, no constriction = 1.
13. Dart sac insertion: in vagina = 0; in atrium = 1.
14. Dart sac papillae: absent = 0; present = 1.
15. Relation between ducts of both mucous glands: separated = 0; distally fused or contiguous = 1.
16. Position of left mucous gland duct: distal respect to the body of the gland = 0; equatorial respect to the body of the gland = 1.
17. Shape of right mucous gland: not sac-like = 0; sac-Like = 1.
18. Right mucous gland: not fused with atrium wall = 0; distally fused with atrium wall = 1.
19. Penis length respect to epiphallus: half epiphallus length = 0; as long as epiphallus = 1; longer than epiphallus length = 2.
20. Penial papillae (= verge): absent = 0; present = 1.
21. Penial retractor muscle: inserts in distal epiphallus = 0; inserts in medial zone of epiphallus = 1; inserts in proximal epiphallus = 2; inserts in proximal penis = 3.
22. Duct of bursa copulatrix: extremely short not longer than sac = 0; medium = 1; long = 2. [additive].
23. Vagina: short = 0; medium to long = 1; extremely long = 2. [additive].
24. Atrium:short = 0; medium to long = 1.
25. Flagellum: thin, long = 0; finger-like, short to medium = 1; Pleurodonte-like = 2; Labyrinthus-like = 3. [additive].
26. Penial muscular band: absent = 0; present = 1.
27. Penial sheath (penial tunica): simple = 0; double or multilayer = 1.
28. Microhabitat associated to: rocks = 0; tree trunks = 1.
29. Vas deferens: surrounding dart sac = 0; not surrounding dart sac = 1.
30. Epiphallus proximal portion: Not widen at point entrance vas deferens = 0; Widen at point of entrance of vas deferens = 1.
31. Penial retractor: not = 0; forming a loop around vas deferens before insertion in epiphallus = 1.
32. Flagellar caecum: absent = 0; present = 1.
33. Kidney length respect to pulmonary roof: not exceeding half of pulmonary roof length = 0; more than half the pulmonary roof = 1.
34. Jaw: ribbed = 0; smooth = 1.