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Systematics, phylogeny and zoogeography of Bulimulinae (Mollusca)
Abstract
In this publication a revision is given of the genera of the subfamily Bulimulinae (Gastropoda, Pulmonata, Bulimulidae). The morphological variation of the shell, radula, pallial organs and genitalia is analized and 21 character transition series are recognized. In the systematical part the following data are presented for each genus : description of shell and anatomy, distribution, ecology, bibliography and a list of taxa. The number of (sub) genera is reduced from 80 to 43 ( + two nomina inquirenda). A new synonymy is : Paracochlea Hylton Scott, 1967 = Eudioptus Albers, 1860. The following new species names are introduced : Bostryx sophieae, Drymaeus (Drymaeus) marcapatensis, Drymaeus (Drymaeus) sophieae, Drymaeus (Mesembrinus) pseudobesus. Berendtia digueti Mabille is designated type species of Teneritia Mabille ; Helix zoographica d'Orbigny is designated type species of Hamadryas Albers. Based on the transition series mentioned above, hypotheses of phylogenetic relationships are presented for the genus groups, using the methods of Hennig. The relationships between the five subfamilies of the Bulimulidae are also investigated but remain tentative. In the zoogeographical section the various theories are reviewed and their relevance for the distribution of the Bulimulinae is treated, using the hypotheses of phylogenetic relationships and Croizat's vicariance theory.
... One of them is Antidrymaeus L. Germain, 1907, originally erected solely for a few sinistral species. Breure (1979) critically revised the genus Drymaeus s.l. based on data later presented by Breure & Eskens (1981). ...
... based on data later presented by Breure & Eskens (1981). Although anatomical data are known from a number of species, Breure (1979) only tentatively recognised 2 subgenera within Drymaeus, Drymaeus s.s. and Mesembrinus Albers, 1850. Later, Schileyko (1999 considered Antidrymaeus L. Germain, 1907 of D. (Mesembrinus). ...
... Although we have some evidence that species treated in the current paper might all belong to Antidrymaeus, we keep them tentatively within their current classifications until further research provides more evidence. Species are allocated to Mesembrinus or Drymaeus following Breure (1979), based on whether the peristome is expanded and on characters of the mandible and radula. ...
Phylogenetic analyses confirm the monophyly of a group of Drymaeus s.l. species that are characterised by tones of bluish or greenish colours of their bodies. This group is now considered as Antidrymaeus L. Germain, 1907, which was recently elevated from synonymy to generic status. Spe- cies of this group have been found from Florida to Brazil and Peru, but further fieldwork may show that more species belong to this genus than hitherto known. The following new taxa are described: Antidry- maeus multifasciatus nevisensis Robinson & Breure and Antidrymaeus multifasciatus luciae Robinson & Breure. The genus Antidrymaeus now includes A. dormani (W.G. Binney, 1857), A. gereti (C.M.F. Ancey, 1901), A. immaculatus (Reeve, 1850), A. inusitatus (Fulton, 1900), A. multifasciatus christopheri (Pilsbry, 1899), A. multifasciatus laticinctus (Guppy, 1868), A. multifasciatus luciae Robinson & Breure, 2024, A. multifasciatus multifasciatus (Lamarck, 1822), A. multifasciatus nevisensis Robinson & Breure, 2024, A. pinchoti (Pilsbry, 1930), A. stramineus (Guilding, 1824), and A. sulphureus (L. Pfeiffer, 1857). Additionally, several species of Drymaeus Albers, 1850 and Mesembrinus Albers, 1850 that also have bluish or greenish colours in their body are treated here. The hypothesis is posed that these also might be members of Antidrymaeus; however, this can only be corroborated through future phylogenetic studies.
NOTE: This article was chosen "editors' choice" and will be freely available from the AfM website (https://www.schweizerbart.de/content/papers/download/106020) till the next issue in March 2025.
... The family Bulimulidae Tryon, 1867 represents a diverse group of land snails occurring in tropical and subtropical regions of North, Central, and South America (Albers, 1850;Breure, 1979). This family comprises three subfamilies, Bostrycinae Breure, 2012, with 15 genera, Bulimulinae Tryon, 1867, and Peltellinae Gray, 1855 (two genera), (MolluscaBase, 2020). ...
... Accordingly, species of the subgenus Mesembrinus are characterized by shells with unexpanded, simple outer lips and those from the subgenus Drymaeus are characterized by shells with expanded or reflected lips. Breure (1979) added other traits for the differentiation of the two subgenera (i.e.: numbers of plates in the jaw and morphology of the radular teeth). The subgenus Mesembrinus is characterized by a peristome usually simple, a jaw with more than 20 ...
... The shell was mounted, without any preparation, on a stub covered with adhesive tape of carbon and imaged in a Hitachi low vacuum tabletop scanning electron microscope, model TM3000. The identification of D. (Mesembrinus) interpunctus was based on the species original description (Albers, 1850) as well as on Breure (1979), , and Simone (2006). ...
Background
Herein, we attempted to obtain detailed data on the distribution of the species of Drymaeus ( Mesembrinus ) in Brazil, using biodiversity databases, malacological collections and literature as sources of occurrence records. We provided a catalogue of species, along with distribution maps. We also estimated the suitable distribution of Drymaeus ( Mesembrinus ) interpunctus using the maximum entropy approach. A detailed description of the anatomy of the soft parts of this species was provided, with new data on the pallial system.
Materials and Methods
For each species we provided information on the compiled data associated with museum collections and the literature. Distribution maps including geographic boundaries, Brazilian biomes and altitude were made with QGIS software 3.16.10 Hannover. For niche modelling, nineteen bioclimatic variables and a topographic variable were used as predictors. The models were performed with MaxEnt version 3.3.3k.
Results
Most of the species are represented by scarce material in malacological collections; for some species, these records correspond to type material, indicating that they have not been recollected. Most of the species were represented by shells making anatomical comparison and DNA analysis difficult, limiting our ability to provide new criteria for species delimitation. Our results allowed us to expand the known distribution area for three species, Drymaeus dutaillyi , D. gereti and D. oreades , with new occurrence records in Brazil. The MaxEnt model showed a thin area of high suitability to D. ( M. ) interpunctus in the Southeastern Brazil, corresponding to the Atlantic Forest. Minimum temperature of the coldest month and mean temperature of coldest quarter were the variables that most influenced the development of the model.
Discussion
Drymaeus interpunctus was described based on specimens collected in Brazil without mention to the exact localities. Herein the new records from databases allowed to expand the known geographic distribution for this species and to infer its potential distribution. Although the type locality of D. interpunctus is in Brazil, the anatomy of the soft parts of specimens from this country was not previously described. The anatomy of the reproductive system of the specimens analyzed herein mostly corresponds to a previous description for specimens from Paraguay, except for the absence of penial sheath and the relative length of the bursa copulatrix duct. The results of niche modeling showed a thin area of high suitability for D. interpunctus and a vast area of moderate suitability, indicating that this species present a niche breadth that favors its occurrence in a range of different biomes, including less suitable areas.
Conclusion
The small number of records obtained for most of the species and their restricted ranges associated with habitat destruction may indicate that they are of conservation concern.
... With 26 genera, this family of terrestrial gastropods is very diverse. Bulimulids can be found associated with a wide variety of environments, such as herbaceous vegetation, dry leaf litter, rocky walls, and arboreal habits; with an altitudinal distribution range from sea level to 5,200 m (Breure, 1979). The family is highly adaptable to a wide variety of climatic conditions and habitats from near-desert to temperate humid forest climates (Coppois, 1995). ...
... In addition, these snails have the ability to colonize new areas and a wide variety of ecologic niches with different types of vegetation, temperature, and humidity conditions (Cabrera et al., 2021). In particular, the genus Bulimulus has a native neotropical distribution (Breure, 1979). In relation to ecological and life history research, the only few available publications from South America involve studies on the life cycles, growth, and reproduction of Bulimulus tenuissimus (D'Orbigny, 1835) from Brazil Meireles et al., 2010;Silva et al., 2008;Silva et al., 2009;Silva et al., 2013). ...
Bulimulus bonariensis is considered a species of relevance to agribusiness, having been declared a pest with indirect damage because of its negative effects on several crops such as soybeans, chickpeas, and corn in central and northern Argentina. The objective of this work was to analyze the growth pattern of a population born under laboratory conditions, to explore population aspects such as survival and mortality, to estimate the age and size at gonadal maturity and first reproduction, and to contribute to the knowledge of the reproductive biology of this gastropod. From the clutches obtained, the basic biologic parameters were calculated and the individuals hatched under laboratory conditions counted and measured every two weeks. The clutches contained an average of 44 eggs, which took about 13.7 days to hatch at a birth rate of 41.82%. The growth pattern in the five clutches was analyzed individually, and the logistic model used was the one with the highest degree of fit to that observed growth pattern, followed by the Gompertz model, and finally the von Bertalanffy model. In addition, the models were applied to the 102 specimens analyzed together as a cohort, where the best fitting model was also proved to be the logistic growth model. A concave type III survival curve was obtained from the horizontal life table. The cohort was reduced by 48% during the first 50 days after birth. Beyond one month of hatching, life expectancy gradually increased and remained high between 65–302 days of life. After day 330, life expectancy decreased and only 13.72% exceeded one year of birth, with an average length of 16.68 mm. The last specimen died after 23 months at a total length of 20.24 mm, and the life expectancy was estimated at almost three years. In addition, it was inferred that gonadal maturity, when these gastropods reach 12 mm of total shell length, is reached after 200 days of life. Therefore, the individuals that are born are able to reproduce for the first time a year after birth, when they have the approximate size of 16.68 mm.
... Futuras observagoes anatomicas, mais apuradas, poderao vir a demonstrar a presenga de estruturas como a fosseta aqui descrita em outras espdcies do genero Scutalus Albers 1850, revistas por Breure (1978Breure ( , 1979 e fomecer mais subsidies para a filogenia do grupo. E possfvel tambem prever-se o encontro em especies de outros generos de Bulimulidae. ...
... Although the taxonomy of the family has been contradictory from several points of view, especially in the morphology and anatomy of the shell (Breure 1979, among others), and that the current distribution of several of these species raises the possibility of the existence of species complexes (Borrero & Breure 2011: 50-51), the general geographic distribution of amphibulimids is as follows: Northern and Northwestern South America (Ecuador, Colombia, Venezuela and Brazil), Central America (Panama) and West Indies (Trinidad and Tobago). In Venezuela, according to the available technical literature (Tello 1975, Breure 1979, 2009, 2011Borrero & Breure 2011; Breure 2012 a -b; Agudo-Padrón 2021; Agudo-Padrón & Vera-Caripe 2022), the family Amphibulimidae is distributed throughout almost the entire national territory, with a total of 24 species taxonomically grouped in two (2) genera: Dryptus with three (3) species, and Plekocheilus with 21 species. ...
Introduction: Since 2014, the “AM Project” (Brazil) has been conducting taxonomic, bioecological and conservationist research on molluscs present in the Caribbean continental territory of Venezuela, with special emphasis on its land (terrestrial and arboreal) forest species (Agudo-
Padrón 2021, Agudo-Padrón & Vera-Caripe 2022). Up to date, these aspects of the natural history of this geographic region have/ had been poorly studied. Objective: Substantial update of this knowledge through the unprecedented curatorial review of the collection of molluscs deposited in the malacological collection of the “Museo de Biología de la Universidad Central de Venezuela – UCV”, based in city of Caracas. Methodology: Between October 2022 and February 2023, a
taxonomic review and examination of material from forest snails of the family Amphibulimidae deposited in the mollusc internal collection of the Museum was performed. Results and conclusions: Determination of fourteen (14) species, collected/ found in 16 geopolitical entities of the country, which were taxonomically grouped into one (1) family and two (2) genera.
... The family Bulimulidae Tryon, 1867 comprises about 543 valid species (MolluscaBase, 2022) native to the tropics and subtropics of South America, Australia, New Zealand, Tasmania, and Africa (Breure, 1979;Herbert & Mitchell, 2009). Especially in the Argentine Republic, this family is the richest and most phylogenetically diverse, as it includes five genera with 46 species (Cuezzo, Miranda & Ovando, 2013;Miranda, 2014;Miranda & Cuezzo, 2014). ...
Bulimulus bonariensis , a snail distributed over a large part of Argentina, has generated negative effects on the agribusiness. We accordingly conducted a seasonal sampling during 2018–2019 in order to characterize the gametic cycle and establish the size of this gastropod at gonadal maturity. Three reproductive stages were identified: the mature in autumn; the spawn, which begins in winter but occurs mainly in the spring, followed by the post-spawn in summer, where an absence of gonadal rest was evidenced. In the fall, in maturity resorption was observed in addition to abundant primary spermatocytes, secondary spermatocytes, and growing vitellogenic oocytes. At the same time, could correlate the degree of spermioviduct development with the stage of gonadal development. Moreover, we used a logistic regression to calculate the size at gonadal maturity, which was established at the total shell length of 12 mm. In addition, we found that at the beginning of reproductive development those gastropods are protogynous hermaphrodites, but after reaching gonadal maturity became simultaneous hermaphrodites. Finally, B. bonariensis also proved to be an iteroparous species. The information provided here will be essential for delineating and establishing population control strategies.
Land-snail inventory suffers many gaps in knowledge, especially in countries of the Global South such as Brazil. Studying undersampled regions and providing data on species distributions are among the most pressing matters in Brazilian malacology. Here, we study a sample of land snails collected in the Central-West region of Brazil (“Midwest”) housed in the Coleção Malacológica, Departamento de Biologia, Faculdade de Filosofia, Ciências e Letras de Ribeirão Preto (University of São Paulo). Ten species from seven families were identified in the material: Alterorhinus constrictus, Anostoma rossi, Aperostoma redfieldi, Bulimulus corumbaensis, Cyclodontina gemellata, Cyclodontina sectilabris, Drymaeus poecilus, Megalobulimus aff. elongatus, Solaropsis fairchildi, and Solaropsis rosarium. The geographic range of four species and one family (Bothriembryontidae) are expanded. We reassess the taxonomic status of two species and consider Anostoma luetzelburgi Weber, 1925 to be a junior synonym of A. rossi Weber, 1925.
The expression ‘you need to know to conserve’ is a well-known cliche among biologists. Documenting the richness of a group of organisms is the first step towards understanding biodiversity and preparing efficient conservation plans. In this context, many efforts have been made to quantify the number of species on Earth and estimate the number of species still unknown to science. A few countries have complete and integrated databases estimating the approximate number of species recorded for their territory, particularly in the Global South. In Brazil, a country of continental dimensions, revealing the richness of the second most diverse clade of invertebrates (=Mollusca) has been a goal of taxonomists. Recently, in an unprecedented, collective, and integrated effort among Brazilian malacologists, it was possible to estimate how many valid species of molluscs are there in Brazil. In this effort, more than 30 mollusc experts joined together to update the Taxonomic Catalogue of the Brazilian Fauna (TCBF), a governmental website that allows a quick and real-time updating of all Metazoan. So far, more than 5,000 updates have been made in TCBF, indicating the presence of 3,552 valid species of molluscs in Brazil, distributed among the main clades as follows: Caudofoveata (10 spp.), Solenogastres (6 spp.), Polyplacophora (35 spp.), Scaphopoda (43 spp.), Cephalopoda (92 spp.), Bivalvia (629 spp.) and Gastropoda (2,737 spp.). The present study, in addition to demonstrating for the first time the richness of Brazilian molluscs, also presents the state of the art of this important phylum of invertebrates highlighting its most representative and neglected groups.
KEY WORDS:
Malacology; taxonomy; database; number of species; molluscan species; biodiversity; conservation
The superfamily Orthalicoidea comprises approximately 2,000 species of terrestrial gastropods, mostly concentrated in the Neotropics but also present in southern Africa and Oceania. We provide a multi-marker molecular phylogeny of this superfamily, reassessing its family- and genus-level classification. We exclude two families from the group, Odontostomidae and Vidaliellidae, transferring them to Rhytidoidea based on their phylogenetic relationships as recovered herein. Two new families are recognized herein as members of Orthalicoidea, Tomogeridae and Cyclodontinidae fam. nov. The family Megaspiridae and the subfamily Prestonellinae are paraphyletic but are retained herein for taxonomic stability. The subfamily Placostylinae is synonymized with Bothriembryontinae. The new genera Alterorhinus gen. nov. and Sanniostracus gen. nov. containing some Brazilian species are described here to better reflect the phylogeny. The fossil record and paleobiogeographic history of the group is explored under the new phylogenetic framework.
Leiostracus Albers, 1850 was established to allocate land snails from Central America and Northern South America. Currently, approximately 19 species are considered valid. However, for most of them, the internal morphology is unknown. Leiostracus obliquus was described as a species of Bulimus from the state of Bahia based on shell characters. Until now, knowledge of this species has been scarce. The discovery of ethanol-preserved specimens from MZSP of this species allowed us to characterize, for the first time, the internal anatomy of this species and update its distribution. The shell of L. obliquus has seven to eight whorls and a wide pale-pink disrupted band all over the teleoconch. The rachidian tooth is small rectangular, symmetric, with round edges, and without differentiated cusps. After comparing the anatomy and radular characters between the shells of L. obliquus and L. carnavalescus, we found remarkable similarities in the morphology and color pattern.
The genus Naesiotus Albers, 1850, which previously also has been considered a subgenus or a synonym of Bulimulus Leach, 1814, has provisionally been studied at the species level. The variation of the shell morphology of 60 species and the analysis of the anatomical data of 23 taxa do not justify a division of Naesiotus in different subgenera. Bulimulus gilderoyi Van Mol, 1972 is placed in the synonymy of Naesiotus cavagnaroi A. G. Smith, 1972. Lectotypes are designated for 10 taxa and new localities are mentioned for 5 species.
In one specimen of Drymaeus canaliculatus (PFEIFFER, 1845) a spermatophore was observed in the phallus complex. The structure of the spermatophore is described using light microscopy observations and histological methods. A hypothesis is presented describing the formation of spermatophores in the epiphallus and flagellum The process of spermatophore-formation is initiated in the most distal part of the epiphallus and the proximal part of the flagellum.
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