Biodiversity Data Journal :
Taxonomy & Inventories
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Corresponding author: Bruno Vila (bruno.vila@univ-amu.fr)
Academic editor: Ivailo Dedov
Received: 09 Dec 2022 | Accepted: 07 Jun 2023 | Published: 27 Jun 2023
© 2023 Louis Aureglia, Jean-Baptiste Guy, Magali Deschamps-Cottin, Bruno Vila
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:
Aureglia L, Guy J-B, Deschamps-Cottin M, Vila B (2023) First observations of Rumina paivae Lowe, 1861 (Mollusca, Gastropoda) in the south-east of France, based on taxonomic clarifications. Biodiversity Data Journal 11: e98627. https://doi.org/10.3897/BDJ.11.e98627
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Amongst the genus Rumina, R. paivae was decribed from North Africa for the first time by Lowe in 1861 on the basis of a limited number of samples. During the 19th and 20th centuries, it was described several times, under different names and different ranks leading to a taxonomic imbroglio before being forgotten. In 2002, Mienis rehabilitated R. paivae, but Prevot et al. (2013, 2014) considered it as a large phenotype of R. decollata Linnaeus (1758) on the basis of genetic and anatomical studies.
In this study, we present morphological and anatomical comparisons and differences between two groups of shells collected in France and considered as R. decollata. Using seven morphological characters related to the size and one to the microscopic sculptures of the shell and two related to the eggs and the colours of the morphs, we attribute these two groups to two morphologically described species: R. paivae and R. decollata. We propose a way to easily distinguish them from each other. With regard to their distribution, morphology and genetics, we discuss their relative systematic position. Moreover, in this study, we report for the first time R. paivae, a given north African taxa, in the south-ast of France, in Marseille.
Rumina paivae, morphology, anatomy, eggs, French Mediterranean area
The genus Rumina includes land snails with an elongated shell and a truncated apex. Of the family Subulinidae, it is the only genus adapted to dry, open and anthropogenised environments with a Mediterranean distribution (
Taxonomically, R. decollata was first described as Helix decollata (
The diagnoses given by Lowe discriminate R. decollata from R. paivae on the basis of conchyliological characters and microscopic sculptures of the shell. Thus, R. paivae, with much larger and wider shells, has more pronounced microscopic sculptures at the sutures than R. decollata (
More recently, within the genus Rumina,
In view of this bibliography, we collected Rumina specimens from the Bouches-du-Rhône (south-east France) that we distinguished, according to strongly contrasting conchyliological differences, within two respective groups, L1 and L2. First, considering all specimens together, we tested whether one or two groups could really be distinguished on the basis of their morphological measurement. Second, on the basis of morphology and anatomy and in comparison with the literature, we associated the group(s) observed with taxa already described. Finally, we discussed the systematic position of the groups as taxa and investigated whether the morpho-anatomical data would allow us to define the first occurrence of a new taxa in France.
Sixteen living individuals and 541 adult Rumina shells were collected in urban areas in parks, gardens and wastelands in Aix-en-Provence, Marseille, Trets and Salon-de-Provence (Bouches-du-Rhône, France). Within these municipalities, our research was concentrated in semi-open environments, under shrubs and hedges (see Material in Taxon treatments). To compare our two groups, we selected only adult individuals with a thickened lip at the peristome (
We measured seven characters of the shell on each of the 541 specimens collected: maximum height (MH), maximum width at the base (BW), maximum width at the apex (AW), maximum diagonal length of the aperture (DLO), maximum horizontal width of the aperture (HLO), height of the first body whorl (HW) and number of whorls (NW) (
The genitalia were studied on 16 individuals, eight individuals from group L1 and eight individuals from group L2 by measuring the length (PL) and width of the penis (PW), the length (VL) and width of the vagina (VW) (Fig.
The individuals reared having laid eggs, we carried out measurements: the diameter (ED) and the weight (EW) of eggs. The measurements were made with the digital measurement software (cellSens Entry 3.1) with a precision of 0.01 mm and the weight of the eggs with a balance (KERN EMB 100-3) with a precision of 0.001 g. Finally, when possible, we observed the body and foot colour of each living individual to associate them with a morph for each Rumina population. All the material (shells, genitalia and eggs) was preserved and deposited in the zoology collections of the University of Aix-Marseille.
In order to test whether one or two groups could be distinguished within the Rumina we collected, we carried out a cluster analysis to calculate the similarity between the individuals collected according the seven morphological criteria measured. For that, we followed the approach led by
Finally, to put our observations into context, we compared our observations of the two groups to the morphological data of width at the base of the shells (BW) of R. decollata collected in France (Supplementary material S1 of
All adult specimens in group L1 were assembled on the basis of their small size.
All adult specimens in group L2 were assembled on the basis of their large size.
The analysis separated the 541 shells into two distinct groups on the basis of seven morphological characters. Cluster 1 contains 351 shells and cluster 2 contains 190 shells (Fig.
The morphological measurements MH, BW, AW, DLO, HLO and HW of L2 are significantly larger than those of L1 (GLM, p < 0.001, Fig.
Characters studied (mean ± se) for groups L1 and L2: the seven morphological characters of the shell, the four characters of the genitalia and the two morphological characters of the eggs accompanied by observations of microscopic sculpture of the shells, the internal structure of the genitalia and colour morphs with number of individuals (n).
Groups (mean ± standard error) | |||
L1 | L2 | ||
Shells | n | 330 | 211 |
MH (mm) | 24.12 (± 4.34) | 37.54 (± 5.39) | |
BW (mm) | 9.04 (± 1.23) | 16.42 (± 1.28) | |
AW (mm) | 5.50 (± 0.96) | 9.53 (± 1.55) | |
DLO (mm) | 8.12 (± 1.40) | 15.26 (± 1.59) | |
HLO (mm) | 5.05 (± 1.22) | 9.69 (± 2.10) | |
HW (mm) | 10.77 (± 2.39) | 20.36 (± 3.74) | |
NW | 4.22 (± 0.82) | 3.50 (± 0.58) | |
Genitalia | n | 8 | 8 |
PL (mm) | 6.66 (± 2.14) | 7.62 (± 2.03) | |
PW (mm) | 1.33 (± 0.38) | 1.79 (± 0.41) | |
VL (mm) | 8.15 (± 1.35) | 7.89 (± 1.34) | |
VW (mm) | 1.64 (± 0.61) | 1.67 (± 0.39) | |
Eggs | n | 68 | 55 |
ED (mm) | 2.82 (± 0.13) | 3.47 (± 0.17) | |
EW (g) | 0.019 (± 0.006) | 0.033 (± 0.005) | |
Shell sculpture | growth streaks | little marked | very marked |
Genitalia structure | penis | prominent papillae | prominent papillae |
vagina | crenellated lamellae | crenellated lamellae | |
Colour morph | foot | black / light brown | white |
body | black / light grey with dorsal line | olive grey |
Estimated marginal means (± se) of the morpho-anatomical criteria between groups L1 and L2, based on GLM models for: A-G morphological values of the seven characters measured H-I morphological values of the eggs; J-M four anatomical values of the genitalia. GLM: ***, p < 0.001; *, p < 0.05; ns, p > 0.05.
The measurements of the genitalia, PL, PW, VL and VW, show no significant difference between the two groups (GLM, p > 0.05) (Fig.
Measurements of the eggs (taken 3 days after laying) show that their diameter (ED) (GLM, p < 0.001) and weight (EW) (GLM, p < 0.001) are significantly higher in L2 than in L1 (Fig.
We observed in L1 two colour morphs: the light grey morph with a black medio-dorsal line (Salon-de-Provence, Trets) and the black morph (Aix-en-Provence, Marseille, Trets). In L2, only one colour morph was observed: the olive-grey morph (Marseille) (Fig.
Finally, comparison (Fig.
Estimated marginal means (± se) of the base width (BW), based on GLM models according to R. decollata collected by Prevot included in the MOTUs A and Eb (
Comparing over 500 individuals, we find highly significant differences between groups L1 and L2 for all conchyliological characters. Group L2 corresponds exactly to the measurements (BW) provided in the diagnosis of R. paivae (
In contrast, our measurements of the anatomical characters of the genitalia show no significant differences between groups L1 and L2, except for the width of the penis. However, this slight difference does not enable us to make it a determining character. Similarly, we do not observe any differences in the internal structures of the genitalia of our two groups, whereas these differ between R. decollata and R. saharica (
With respect to egg diameter, those of group L1 match the diameter of R. decollata eggs (on average 2 mm) measured by
The observed body and foot colour morphs are characteristic for each of the groups: individuals from group L1 share the same morphs as French R. decollata from MOTUs A and Eb and those from group L2 share the same morph as R. paivae from MOTUs C, D and Ea (
Like the majority of plant and animal species, the species of the genus Rumina have been described solely on the basis of morphological characters and correspond to the practical concept of morphological species. Recent genetic studies by
The present study does not aim to discuss the notion of species within the genus Rumina since we do not have sufficient data (whether morphological, anatomical or genetic), but also and above all because the question of the notion of species is variable and difficult. The complementary elements that we can bring to the new data presented here (morphology and anatomy) concern observations relating to the biology of the species that may have repercussions on the notion of species. During our sampling, the two taxa were not found in sympatry on a local scale. This implies that the populations seem to be independent and autonomous. The absence of sympatry of the two taxa also leads us to rule out the existence of a large phenotype. However, at the same time, within the genus Rumina, there is the case of frequent self-fertilisation that can promote genotypic and phenotypic differentiation by fixation of alternative alleles at various nuclear gene loci (
In view of the uncertainties related to the division of R. decollata into MOTUs (
We observe that the two groups identified for the first time in France, in Marseille (in separate localities), are easily distinguished on the basis of morphological characters and micro-sculptures of the shells. As a result, in the context of this study, we relate the two statistically identified groups to the two species already described under the names R. decollata Linnaeus, 1758 and R. paivae Lowe, 1860.
To date, no occurrence of R. paivae has been reported in France nor in Europe. Knowing that the distribution area of this species is currently north African (Algeria, Morocco, Tunisia), we report here the first occurrence of R. paivae in the south-east of France, in Marseille.
We can formulate two hypotheses to explain the presence of this species in Marseille:
(1) R. paivae, of north African origin, may be a locally introduced species due to the numerous links and exchanges between these two areas. The presence of self-fertilisation within this genus could have favoured its establishment and the maintenance of populations from a small number of individuals. We know that R. decollata has been accidentally introduced in the United States, Argentina, Brazil, Uruguay, South Africa, China and Japan (
(2) The distribution of R. paivae may be underestimated around the Mediterranean Basin. The species may be more common, but might go unrecorded because of the polymorphism attributed to the taxon. A good example is provided in botany with Arundo micrantha Lam. Described with a north African distribution,
A morphological and genetic study, including a large number of specimens (body and shells) from North Africa and other countries around the Mediterranean where large forms exist, would undoubtedly shed new light on the taxonomic rank to be attributed to what we call in this publication R. paivae. Similarly, further research within the MOTUs of R. decollata would perhaps make it possible to identify distinctive characters, at least on living individuals.
We would like to thank L. Cavalli and O. Gerriet for their help and advice during the dissections, Q. Bremond and F. Blanc for giving us access to dissection material and the staff of the Museum of Natural History of Aix-en-Provence, Avignon, Marseille and Nice for having allowed us to consult their collections. We also thank F. Magnin for his advice and the proofreaders who enabled us to improve the quality of the manuscript.