Research Article |
Corresponding author: Luiz Ricardo L. Simone ( lrlsimone@usp.br ) Academic editor: Frank Köhler
© 2020 Luiz Ricardo L. Simone, Daniel Caracanhas Cavallari, Rodrigo Brincalepe Salvador.
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:
Simone LRL, Cavallari DC, Salvador RB (2020) A new troglobite species of Habeastrum Simone, 2019 from Brazil, and support for classification in Diplommatinidae (Mollusca, Caenogastropoda). Zoosystematics and Evolution 96(2): 639-647. https://doi.org/10.3897/zse.96.53880
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The genus Habeastrum Simone, 2019 was recently described based on empty shells, counting with two troglobite species. Conchological features allowed a preliminary classification in the caenogastropod family Diplommatinidae, but this family allocation was left open to future studies. Herein, we present a detailed anatomical study of newly acquired specimens, confirming the classification in Diplommatinidae. These new specimens, from Minas Gerais state, SE Brazil, belong to a new troglobite species described herein, Habeastrum strangei sp. nov. The present records extend the genus distribution ca. 1,100 km east-northeast.
Cyclophoroidea, Gastropoda, Habeastrum strangei sp. nov., Minas Gerais, troglofauna
The genus Habeastrum Simone, 2019, has been recently described to allocate two unique and previously unknown species collected in caves in Mato Grosso do Sul state, central Brazil. It presently includes two terrestrial troglobite species: the type species H. parafusum Simone, 2019 and H. omphalium Simone, 2019.
The original description of the genus was based solely on empty shells (the operculum was also unknown) and the following conchological features allowed a tentative classification in Diplommatinidae (
Such specimens were recently sent to us by two different sources: Prof. Dr. Maria Elina Bichuette, a researcher from the Universidade Federal de São Carlos (UFSCar, Brazil) specialized in Brazilian troglofauna, and the environmental consulting company Carste Ciência e Meio Ambiente (CARSTE, Brazil). These new specimens were collected in caves in various different municipalities in Minas Gerais state, southeastern Brazil. We present here an anatomical study of the specimens, confirming the diplommatinid affinity of the genus. Furthermore, they are deemed to belong to a new species, described herein as Habeastrum strangei sp. nov.
The new material from Minas Gerais contained both whole specimens (preserved in 70% ethanol) and empty shells. It was donated by Dr. Bichuette to the malacological collection of the Museu de Zoologia da Universidade de São Paulo (
Two specimens were randomly selected for DNA extraction, with two samples acquired from each. Unfortunately, we could not acquire good quality DNA for amplification and sequencing. Even though the specimens seemed fresh, we suspect they might have stayed too long in the field sample before being fixed in ethanol and/or not have been thoroughly fixed: the snails were deeply retracted into their shells and thus, the ethanol might not have reached them properly given the presence of the operculum. After these four trials, using two different extraction kits and protocols, we opted for not spending more specimens.
List of abbreviations used in the figures: aa, anterior aorta; an, anus; ap, aperture of pallial gonoduct; au, auricle; bg, buccal ganglion; ce, cerebral ganglion; cm, columellar muscle; cv, pulmonary (ctenidial) vein; dg, digestive gland; di, diaphragm septum; eg, esophageal gland; es, esophagus; ey, eye; fs, foot sole; ft, foot; go, gonad; in, intestine; jw, jaw; ki, kidney; m1 to m6, extrinsic and intrinsic odontophore muscles; mb, mantle border; mj, jaws and perioral muscles; mo, mouth; oa, opercular pad; oc, odontophore cartilage; od, odontophore; op, operculum; ot, oral tube; ov, pallial oviduct; pa, penis distal aperture; pc, pericardium; pd, penis duct; pe, penis; pl, pleural ganglion; pn, pedal ganglion; pt, prostate; py, pallial cavity (lung); rn, radular nucleus; rs, radular sac; rt, rectum; sn, snout; st, stomach; te cephalic tentacle; tg, integument; vd, vas deferens; ve, ventricle; vo, visceral oviduct.
Family Diplommatinidae L. Pfeiffer, 1856
Habeastrum parafusum Simone, 2019; Recent, Brazil.
Holotype
Shells of Habeastrum strangei sp. nov. A–E. specimens before shell crushing for anatomical study (
Brazil, Minas Gerais state, Presidente Olegário municipality, Lapa da Fazenda São Bernardo (18°16'37"S, 46°06'46"W).
Brazil. Minas Gerais. Pains municipality: Gruta dos Coralóides (20°20'33.4"S, 45°46'45.7"W),
The specific epithet is given in honor of Dr. Stephen V. Strange, a fictional character from Marvel Comics. Besides being an acknowledgement of one of the greatest Marvel characters, the name also alludes to the weirdness of this dextral diplommatinid species.
Spire conical and tall. Teleoconch sculpture consisting of strongly prosocline, markedly raised ribs, more widely spaced than in congeners.
Shell (Figs
Head-foot (Fig.
Anatomy of Habeastrum strangei sp. nov. A. head-foot, male, right-slightly anterior view; B. pallial cavity, ventral view and visceral mass partially uncoiled, some structures seen be translucency; C. head and haemocoel, ventral view, foot and columellar muscle removed; D. foregut, ventral view; E. same, left view; F. odontophore, dorsal view, superficial layer of structures removes, cartilages deflected. Scale bars: 250 μm.
Pallial organs (Figs
Visceral mass (Fig.
Circulatory and excretory systems (Fig.
Digestive system (Fig.
Radular sac very long, stored as 4–5 compact whorls located to the right of esophagus in the region preceding buccal mass (Fig.
Buccal cavity with pair of wide and low dorsal folds as continuation from jaw plates, gradually diminishing towards esophagus. Pair of buccal ganglia (Fig.
Genital system (Fig.
Central nervous system (Fig.
In the caves in Presidente Olegário municipality, this species was found in epigeal environments and in all cave zones (entrance, twilight, and dark zones). Most specimens collected consisted of empty shells, but the good preservation considering their fragility suggests they were not subjected to transport. No precise information was recorded from other municipalities.
As discussed in the original description of Habeastrum (
Even though the conchological variation in Diplommatinidae is well described and illustrated in the literature, the same is not true for their anatomical features. Most works have focused on other cyclophoroidean families (
Overall, the shape of the snout, the arrangement of the pallial cavity, reproductive system, and the hypoathroid central nervous system of Habeastrum (Figs
The only other Latin American diplommatinid species with a detailed anatomy is Adelopoma paulistanum Martins & Simone, 2014, from São Paulo state, SE Brazil. Habeastrum strangei sp. nov. shares some features with it, such as: the shape of the snout; the duplication of the eyes; the attributes of the pallial cavity (lacking gill and osphradium, with kidney and pericardium disposed in a similar way); the characters of the jaws and odontophore; and the relatively large and similarly-arranged nerve ring. By contrast, Habeastrum strangei sp. nov. differs from A. paulistanum by: the dextral coiling; the broader width of the pallial genital structures; the presence of esophageal glands; the lack of salivary glands; the stomach bulging posteriorly; and the presence of a penis. The latter is a plesiomorphic state in Diplommatinidae, as some species are aphallic (
There is more information available on the radulae of other Neotropical diplommatinids than any other anatomical feature (
Besides the now three species of Habeastrum, there are only six other diplommatinids reported from Brazil, belonging to the genera Adelopoma and Habeas Simone, 2013 (
We are deeply grateful to Lina Bichuette and her team (UFSCar) and the people from CARSTE for the collection and donation of the specimens (collection under the auspices of ICMBIO); to Lara Guimarães (