Corresponding author: Maria Gabriela Cuezzo (
We describe a new genus and a new species in the family
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
Snails were collected at the National Park Cavernas do Peruaçu (
Live specimens and dry shells of
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
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.
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
Shell (Figs
Brazil,
Brazil,
Thus far known only from National Park Cavernas do Peruaçu, northern region of Minas Gerais, Brazil.
In
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: (2) Dorsal view of Holotype, MNRJ 34.580; (3) lateral position of holotype shell and soft body; (4) dorsal, (5) ventral, and (6) lateral view of paratype, IBN 861; (7) live snail. Scale bar: 3–5 = 10 mm.
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
The main goal of the analysis performed was to evaluate and support the description of a new genus. For this, only the species of
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 | ||
|
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 |
|
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 |
|
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 |
|
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 |
|
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 |
|
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 |
|
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 |
|
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 |
|
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 |
|
4 | 2 | 0 | 2 | 0 | 0 | 1 | 0 | 0 | 0 | 1 | 1 | 0 | ? | ? | ? | 0 | 1 | ? | ? | ? | ? | 0 | ? | ? | 1 | ? | ? | 0 | 0 | 1 | ? | 0 | 0 | 0 |
|
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 |
|
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 |
|
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 |
|
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 |
|
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 |
|
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 |
|
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
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;
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.