Non-native terrestrial slugs from Sinaloa, Mexico: Deroceras laeve (O. F. Müller, 1774) and Sarasinula plebeia (P. Fischer, 1868) (Mollusca, Gastropoda)

Abstract

This is the first record of two non-native terrestrial slug species from Sinaloa, Mexico. Deroceras laeve and Sarasinula plebeia were collected between 2019 and 2022 in Concordia and Mazatlan Municipalities (north-western Mexico). The external morphology and anatomic features of the dissected specimens coincide with the descriptions of each species, whose identities were also confirmed by their partial COI sequences. The ample occurrence of S. plebeia suggests that this species has an established population, while D. laeve was found as isolated individuals, likely associated with plant nurseries.

Full text

Biodiversity Data Journal 10: e87666
doi: 10.3897/BDJ.10.e87666
Research Article
Non-native terrestrial slugs from Sinaloa, Mexico:
Deroceras laeve (O. F. Müller, 1774) and
Sarasinula plebeia (P. Fischer, 1868)
(Mollusca, Gastropoda)
Laura Regina Alvarez-Cerrillo , Beatriz Yáñez-Rivera , Victoria Araiza-Gómez
‡ Centro de Investigación en Alimentación y Desarrollo, A. C. (CIAD), Unidad Mazatlán en Acuicultura y Manejo Ambiental,
Av. Sábalo-Cerritos s/n, Estero del Yugo, C.P. 82100, Mazatlan, Mexico
§ Instituto Politécnico Nacional, Escuela Nacional de Ciencias Biológicas, Departamento de Zoología, Prolongación de Carpio
y Plan de Ayala s/n, Col. Casco de Santo Tomás, 11340, Mexico City, Mexico
Corresponding author: Victoria Araiza-Gómez (varaiza9693@gmail.com)
Academic editor: Alexander M. Weigand
Received: 11 Jun 2022 | Accepted: 26 Jul 2022 | Published: 04 Aug 2022
Citation: Alvarez-Cerrillo LR, Yáñez-Rivera B, Araiza-Gómez V (2022) Non-native terrestrial slugs from Sinaloa,
Mexico: Deroceras laeve (O. F. Müller, 1774) and Sarasinula plebeia (P. Fischer, 1868) (Mollusca, Gastropoda).
Biodiversity Data Journal 10: e87666. https://doi.org/10.3897/BDJ.10.e87666
Abstract
This is the first record of two non-native terrestrial slug species from Sinaloa, Mexico.
Deroceras laeve and Sarasinula plebeia were collected between 2019 and 2022 in
Concordia and Mazatlan Municipalities (north-western Mexico). The external morphology
and anatomic features of the dissected specimens coincide with the descriptions of each
species, whose identities were also confirmed by their partial COI sequences. The ample
occurrence of S. plebeia suggests that this species has an established population, while D.
laeve was found as isolated individuals, likely associated with plant nurseries.
Keywords
introduced, pest slug, COI, Agriolimacidae, Veronicellidae
‡ ‡ §
© Alvarez-Cerrillo L et al. 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.

Introduction
Non-native slug species can be responsible for the displacement of native species, crop
damage and habitat destruction. Several ways of introduction into new ecosystems have
been hypothesised, for example, commercial trade of plants or passive transportation of
eggs and juveniles or adults attached to birds (Aubry et al. 2006, Barbato et al. 2017). The
most updated assessment of non-native slugs in Mexico was performed by Naranjo-García
and Castillo-Rodríguez (2017), who reported thirteen species: Arion circumscriptus G.
Johnston, 1828; A. intermedius Normand, 1852; Boettgerilla pallens Simroth, 1912;
Deroceras invadens Reise, Hutchinson, Schunack & Schlitt, 2011; D. laeve (O. F. Müller,
1774); D. reticulatum (O. F. Müller, 1774); Ambigolimax valentianus (A. Férussac, 1821)
(listed as Lehmannia valentiana); Limacus flavus (Linnaeus, 1758); Limax maximus
Linnaeus, 1758; Milax gagates (Draparnaud, 1801); Phyllocaulis gayi (P. Fischer, 1871)
and Sarasinula plebeia (P. Fischer, 1868). Recently, Arion vulgaris Moquin-Tandon, 1855
was recorded by Araiza-Gómez et al. (2021).
The non-native slugs previously reported in literature for the State of Sinaloa are A.
valentianus (Naranjo-García and Castillo-Rodríguez 2017), P. gayi and S. dubia (Naranjo-
García et al. 2007). However, in the same reports, the authors argued the need to confirm
the occurrence of P. gayi due to the absence of records since 1925. The knowledge of
native slugs from Sinaloa is limited. Citizen observations in iNaturalist (https://
www.naturalista.mx/projects/moluscos-de-Mexico), suggest the presence of Leidyula
moreleti (Crosse & Fischer, 1872). The occurrence of the species was confirmed by us
using the external morphological characteristics of the species given the limitations of the
platform.
The native origin of D. laeve remains uncertain; its actual distribution seems to be
cosmopolitan (Gittenberger et al. 2018). It has been hypothesised that the species is
Palearctic in origin and subsequently spread throughout Europe (Wiktor 2000). However,
several previous records suggest a Holarctic origin of the species (Sysoev and Schileyko
2009), capable of colonising a wide range of habitats due to its huge ecological plasticity.
The genus Deroceras (Agriolimacidae) includes at least 123 species ( Wiktor 2000). In
Mexico, D. laeve was first reported from the State of Veracruz (south-eastern Mexico)
(Strebel and Pfeffer 1880), with recent records from Chihuahua and Durango (northern
Mexico) (Araiza-Gómez et al. 2017).
Sarasinula plebeia was described from New Caledonia (Oceania). Darrigran et al. (2020)
commented that its origin in South America is unknown. Thomé (1993) explains the
presence of this species in Rio de Janeiro, Brazil concerning changes in taxonomy and
nomenclature due to the native species Vaginula behni Semper, 1885, which was later
synonymised with S. plebeia. Recent reports as a non-native species were documented in
North and Central America (Daglio et al. 2020). The first record of S. plebeia in Mexico was
from Chiapas and Veracruz (Andrews and Dundee 1987), with most records occurring in
the centre and south of the country and some records from the north of Mexico. In the
2Alvarez-Cerrillo L et al

current contribution, we present new distributional records of D. laeve and S. plebeia from
Sinaloa. We support these records with anatomic and molecular data.
Material and methods
Manual collection for living slugs were carried out between August 2019 and April 2022 in
different urban and rural locations of six municipalities in the State of Sinaloa (north-
western Mexico), but these species were only found in two: Concordia and Mazatlan (Table
1). In total, 122 specimens of Sarasinula plebeia and three of Deroceras laeve were
collected.
Species Municipality Locality Altitude (m a.s.l.) Coordinates n
Deroceras laeve Mazatlan *Portomolino 11 23.26263, -106.406 1
**Siqueros 28 23.339309, -106.239243 2
Sarasinula plebeia Mazatlan *Marivento 11 23.278083, -106.429989 112
*Cerro del Vigia 38 23.190944, -106.425833 7
Concordia **Panuco 656 23.428808, -105.896592 3
The specimens were relaxed in a jar with water until fully stretched and died (~ 12 h).
Some specimens were photographed alive using a digital camera (Lumix DMC–FS3,
Panasonic). Once dead, the cleaning of mucus was performed in a sieve under running
cold water. The slugs were then fixed on 90% ethanol.
All specimens were examined for external morphology and 25 slugs of S. plebeia and two
of D. laeve were dissected. Two specimens of S. plebeia and one of D. laeve (which was
damaged during processing), were selected for molecular analysis.
DNA extraction of the tissue of the foot muscle was performed using the Blood and Tissue
kit according to the manufacturer's specifications (QIAGEN, California, USA). The integrity
and quality of the DNA was verified on an agarose gel by electrophoresis. The COI gene
was amplified by PCR using COIF and COIR (López et al. 2019). The reaction mixture
consisted of 50 ng of DNA, 0.6 μl of each 10 mM primer, 1.5 μl of 10x buffer, 0.6 μl of 10
mM dNTPs, 1.5 μl of 50 mM MgCl and 0.2 μl of Taq Polymerase (Invitrogen) in a total
volume of 25 µl. The amplification conditions were initial denaturation at 94°C for 5 min,
followed by 35 cycles at 94°C for 30 s, 50°C for 30 s, 72°C for 30 s and a final extension at
72°C for 7 min.
The PCR products were purified with the GFX™ PCR DNA and GelBand Purification Kit
(GE Healthcare, Buckinghamshire, UK), then sequenced by Macrogen Inc., Korea. The
nucleotide sequences were compared with the sequences deposited in the gene bank
2
Table 1.
Table 1. Slugs collected in two municipalities of Sinaloa State, Mexico. Urban (*), agricultural/rural
(**).
Non-native terrestrial slugs from Sinaloa, Mexico: Deroceras laeve (O. ... 3

(GenBank) of the National Center for Biotechnology Information (http://
www.ncbi.nlm.nih.gov), with the BLASTn algorithm. The sequences obtained were
deposited in GenBank (Access number: ON678123–25).
A taxonomic assignment was carried out using a phylogenetic inference analysis by
Maximum Likelihood (ML) with the algorithm implemented in PhyML v. 3.0 (Guindon et al.
2010). The nucleotide substitution model that best fits the data was determined with SMS:
Smart Model Selection in PhyML (Lefort et al. 2017). The evolutionary selected model by
the Akaike Information Criterion for each dataset (Suppl. material 1) was the HKY85 model
with gamma distribution, shape = 0.173 for Deroceras and shape = 0.566 for Sarasinula.
To estimate the reliability of each node, a bootstrap procedure was performed with 1000
pseudoreplicates. We employed as outgroup Limax maximus ( KF894386, KM612139) for
the genus Deroceras and Onchidium (MN528062, KX179520) for the genus Sarasinula.
Results
Family Agriolimacidae
Deroceras laeve (O. F. Müller, 1774) (Fig. 1)
Morphology: Live pigmentation grey and dark brown (Fig. 1a), ocular tentacles almost
black, with concentric striations in the mantle characteristic for the genus. Dorsum with
Figure 1.
Deroceras laeve: a live specimen; b genitalia (Bc: bursa copulatrix, Ov: oviduct, Pe: penis); c
external appearance of a preserved specimen.
4Alvarez-Cerrillo L et al

tubercles, distally with barely noticeable keel. Body length of preserved specimens, 18-22
mm. The bursa copulatrix on the oviduct and a barely visible penis were observed (Fig. 1
b). The aphallic or phallic form was unconfirmed, possibly related to the size of the
specimen and preservation state.
Molecular markers: A fragment of 717 bp was obtained. After editing, a fragment of 661
bp was used for the BLAST analysis and another of 310 bp for the phylogenetic analysis.
The sequence obtained in this study had 98.22% nucleotide identity of similarity with
sequence KX959495.1 registered in GenBank as D. laeve. The sequence clustered closer
in the phylogeny with specimens from Mexico (KX959494, KX959501) and Canada
(MG421943) (Fig. 2).
Remarks: The main differences between D. laeve and other species of the genus are
related to the mantle coverage, pigmentation and reproductive features (Araiza-Gómez et
al. 2017). Deroceras laeve and D. invadens share the mantle length that covers almost half
the body and have similar sizes (~ 24 mm and ~ 28 mm, respectively). However, these two
species differ in pigmentation; D. invadens has a pale greyish body with a few spots on the
creamy-brown mantle. The differences with D. reticulatum are the cover of the mantle that
reaches one third of the body length (~ 35 mm) and pale brown pigmentation, almost white
with dark spots, with the sole cream-coloured. The pigmentation patterns in D. laeve
include light brown, grey to almost black, some with a speckled mantle, the sole cream-
coloured and others black delineated (Araiza-Gómez et al. 2017).
Regarding reproductive features, D. laeve can be found in two forms: the phallic form with
a long slender penis and the aphallic form with greatly reduced or missing male
Figure 2.
Maximum Likelihood (ML) phylogenetic tree reconstruction of Deroceras using 310 bp of the
COI. Numbers on branches indicate ML bootstrap values. GenBank access numbers of
sequences employed in supplementary material (Suppl. material 1).
Non-native terrestrial slugs from Sinaloa, Mexico: Deroceras laeve (O. ... 5

reproductive organs (Wiktor 2000, Araiza-Gómez et al. 2017). The variant of this
species
was unconfirmed due to the preservation condition. Deroceras reticulatum has a penial
gland on the proximal part of its penis with a flagellum having a variable number of bulbous
branches. Deroceras invadens shows two side pockets, the penial lobe and caecum of
roughly equal width, but with a longer penial lobe, both having rounded tips; gland fingers
long mid-way between the pockets and the retractor muscle attaches between the lobe and
caecum.
Habitat: The three specimens were collected in gardens and from plants recently
purchased at plant nurseries. The first individual was collected from Spathiphyllum spp. in
an urban house. The other two were collected from Euphorbia pulcherrima in an
agricultural/rural area. Globally, D. laeve inhabits an extremely wide range of habitats
(Dedov et al. 2020).
Family Veronicellidae
Sarasinula plebeia (P. Fischer, 1868) (Fig. 3)
Morphology: Live pigmentation brown (Fig. 3a), with scattered small punctuations in the
thickened notum (Fig. 3b); light grey after preservation. Body length 45.26 ± 10.92 mm
Figure 3.
Sarasinula plebeia: a live specimen; b punctuations on the notum of the specimen; c male
reproductive system (Pa: papilla of the digitiform gland, Pe: penis, Tu: digitiform tubules, Vd:
vas deferens); d hyponotum (Pf: female genital pore).
6Alvarez-Cerrillo L et al

(min = 20.75 mm, max = 57.95 mm, n = 122). Penis short, smooth, without annular
protrusion, bilaterally symmetrical, with enlarged glands, even the digitiform gland (Fig. 3c)
with an elongated form; with four to six tubules subequal in length, but some individuals
showed one shorter than the rest.
Molecular markers: A fragment of 800 bp was obtained. After editing, a fragment of 621
bp was used for the BLAST analysis and another of 470 bp for phylogenetic analysis,
matching the length of the sequences in GenBank. The sequences (n = 2) had 100% of
similarity with sequence JX532107.1 registered in GenBank and 99.83% with sequences
MZ598573.1, KM489367.1, both identified as S. plebeia. Mexican sequences clustered
closer in the phylogeny with specimens from Okinawa, Japan (Fig. 4).
Remarks: The morphology of the penis is the main feature to differentiate S. plebeia and
S. dubia. The penis is club-shaped in S. plebeia and tapering distally in S. dubia;
Figure 4.
Maximum Likelihood (ML) phylogenetic tree reconstruction of Sarasinula using 470 bp of the
COI. Numbers on branches indicate ML bootstrap values. Tip label is the accession number of
GenBank and the country of origin.
Non-native terrestrial slugs from Sinaloa, Mexico: Deroceras laeve (O. ... 7

identification of veronicellid slugs is valid when characters of sexual anatomy and penial
gland are taken into consideration (Thomé 1989). We observed a slight variation in
digitiform tubules (four to six).
Habitat: Abundant in gardens of urban houses. Only three specimens were found in
natural vegetation next to a tributary of the Panuco River (near an abandoned mine).
Globally appears limited to tropical environments.
Specimens of S. plebeia were collected in all months, except for June and July between
2019 and 2022 (Fig. 5). In these months, the mean temperature increases with relatively
less humidity than in August, the hottest month in summer. This probably limits the activity
of slugs.
Discussion
The list of species of non-native terrestrial slugs from Mexico, provided by Naranjo-García
and Castillo-Rodríguez (2017), already includes both species reported herein. However,
their data include limited coverage of localities in northern Mexico. The occurrence of
Deroceras laeve and Sarasinula plebeia in the State of Sinaloa are the first records for this
region. The introduction of terrestrial gastropods is related to horticulture, agriculture and
ornamental plants (parks and gardens), in at least 20% of the cases. The vectors of the
40% of documented introductions are unknown (Darrigran et al. 2020). Our findings
indicate that both species could have been introduced in this area via nursery plants,
occurring mostly in gardens, but the presence of S. plebeia in natural vegetation suggests
that this species is already invading natural habitats.
Figure 5.
Temporal distribution of S. plebeia during the sampling along the years 2019–2022 (n = 122).
8Alvarez-Cerrillo L et al

Impacts related to biological invasions have a direct effect on biodiversity loss (Bellard
et
al. 2021, Dueñas et al. 2021). In addition, introducing invasive slugs, mainly the Sarasinula
species, is related to dry-bean and maize crop damage in Central America (Rueda et al.
2002). Although it appears that their effect as agricultural pests has been reduced by
improving management by farmers, it remains as a vector for parasites for rodents and
other mammals and humans (Nurinsiyah and Hausdorf 2018). Spatio-temporal occurrence
of S. plebeia indicates an actual expansion process and their genetic convergence with
Japanese and South American individuals demonstrate their invasive potential (Hirano et
al. 2022).
The widespread distribution of both species could be true. In Pakistan, Havlác (2004)
discussed the role of gardening activities in the population establishment of D. laeve. This
species is the most widely distributed in the country, although the spread of three species
of the genus (D. laeve, D. invadens, D. reticulatum) is a fact in Mexico (Araiza-Gómez et
al. 2017). Deroceras laeve still have a limited distribution in the region and is restricted to
winter conditions and in plant species commonly used in gardening. This species has been
registered as a pest in crops, such as cabbages, maize, soybean, amongst others, being a
pest in agriculture and horticulture worldwide (Byers and Calvin 1994, Gittenberger et al.
2018). The genetic convergence of individuals from Sinaloa is related to other Mexican
localities, such as Mexico City and the States of Oaxaca, Puebla, Queretaro and San Luis
Potosi (GenBank access: KX959492-99, KX959500-01) and from Canada.
The establishment of non-native species requires favourable local conditions and
temperature and humidity seem to be crucial, for the development of the population and
further expansion into surrounding natural habitats (Dedov et al. 2020). The reproductive
characteristics of non-native species play a crucial role in their invasive potential. Both
species here considered are hermaphrodites with a self-fertilisation strategy (Rueda et al.
2002, Clemente et al. 2007). The development of D. laeve has two phases, a juvenile
stage of pre-oviposition and a mature stage of oviposition during which the slugs lay their
eggs; self-fertilisation is its normal breeding system; isolated individuals can produce fertile
eggs. The reproduction occurs either in autumn or spring, once during their lifetime (Faberi
et al. 2006). The individuals of S. plebeia can function as both male and female during their
lifetime and self-fertilisation may occur in isolation (Rueda et al. 2002). The reproductive
maturity occurs at ~ 2.5 months of age. An organism can produce one to four clutches per
year with approximately 30 eggs. Reproduction is generally high during the rainy season
(Naranjo-García et al. 2007).
Biodiversity inventories require reliable species identification, but in terms of biological
invasions, the correct species assignment is essential. In this regard, the high number of
synonyms is related to the use of few morphological characters for species delimitation or
superficial revision, based on external features and pigmentation patterns (Maceira F. 2003
). The taxonomy of the slugs requires analysing their internal anatomy of male genitalia
and the use of molecular markers (Hirano et al. 2022), a crucial requirement to understand
the dispersion of invasive species. For instance, considering that S. plebeia is one of the
two species with the widest distribution (Gomes and Thomé 2004), further studies on this
Non-native terrestrial slugs from Sinaloa, Mexico: Deroceras laeve (O. ... 9

species could use molecular data to identify clades associated with introduction routes and
spreading.
The main contribution of this paper consists of new records of two non-native species
supported by molecular data, as a step towards the better understanding on terrestrial
slugs invasions.
Acknowledgements
LRAC wants to give special thanks to collector volunteers: Yazmín Segura, Eber Barraza,
Albert M. van der Heiden, Maleny Lizárraga, Diana Teresa, Citlallic Pimentel, Martha
Chapa and Guillermo Otero. We thank Laura Márquez and Nelly López (IB-UNAM) for
sequencing service, Miguel Betancourt-Lozano and Samuel Gómez for the helpful
comments on the manuscript.
Author contributions
All authors have reviewed and agreed upon the content of the manuscript and have met
the requirements for authorship.
Conflicts of interest
There are no conflicts of interest to declare and the study did not involve human subjects.
References
• Andrews KL, Dundee L (1987) Las babosas veronicellidos de Centroamérica con
énfasis en Sarasinula plebeia (=Vaginulus plebeius). Ceiba 28: 163‑172. URL: http://
hdl.handle.net/11036/3909
• Araiza-Gómez V, Naranjo-García E, Zúñiga G (2017) The exotic slugs of the genus
Deroceras (Agriolimacidae) in Mexico: Morphological and molecular characterization,
and new data on their distribution. American Malacological Bulletin 35: 126‑133. https://
doi.org/10.4003/006.035.0205
• Araiza-Gómez V, Naranjo-García E, Zúñiga G (2021) Occurrence in Mexico of two
European invasive slug species: Arion vulgaris Moquin-Tandon, 1855 and Arion
intermedius (Norman, 1852). BioInvasions Records 10 (1): 10‑20. https://doi.org/
10.3391/bir.2021.10.1.02
• Aubry S, Labaune C, Magnin F, Roche P, Kiss L (2006) Active and passive dispersal of
an invading land snail in Mediterranean France. Journal of Animal Ecology 75 (3):
802‑813. https://doi.org/10.1111/j.1365-2656.2006.01100.x
• Barbato D, Benocci A, Caruso T, Manganelli G (2017) The role of dispersal and local
environment in urban land snail assemblages: an example of three cities in Central Italy.
Urban Ecosystems 20 (4): 919‑931. https://doi.org/10.1007/s11252-017-0643-8
10 Alvarez-Cerrillo L et al

• Bellard C, Bernery C, Leclerc C (2021) Looming extinctions due to invasive species:
Irreversible loss of ecological strategy and evolutionary history. Global Change Biology
27 (20): 4967‑4979. https://doi.org/10.1111/gcb.15771
• Byers R, Calvin D (1994) Economic injury levels to field corn from slug
(Stylommatophora: Agrolimacidae) feeding. Journal of Economic Entomology 87 (5):
1345‑1350. https://doi.org/10.1093/jee/87.5.1345
• Clemente N, Faberi A, López A, Manetti P, Álvarez H, et al. (2007) Biología de
Deroceras reticulatum y D. laeve, moluscos de cultivos en siembra directa. Revista de
Investigaciones Agropecuarias 36 (2): 129‑142. URL: https://www.redalyc.org/
articulo.oa?id=86436210
• Daglio ED, de Lucía M, Rodrigues Gomes S, Gutiérrez Gregoric DE (2020) First
records of the bean-slug Sarasinula plebeia (Gastropoda: Veronicellidae) in Argentina.
Papéis Avulsos de Zoologia 22: 853‑871. https://doi.org/10.11606/1807-0205/
2020.60.47
• Darrigran G, Agudo-Padrón I, Baez P, Belz C, Cardoso F, Carranza A, Collado G,
Correoso M, Cuezzo MG, Fabres A, Gutiérrez Gregoric DE, Letelier S, Ludwig S,
Mansur MC, Pastorino G, Penchaszadeh P, Peralta C, Rebolledo A, Rumi A, Santos S,
Thiengo S, Vidigal T, Damborenea C (2020) Non-native mollusks throughout South
America: emergent patterns in an understudied continent. Biological Invasions 22:
853‑871. https://doi.org/10.1007/s10530-019-02178-4
• Dedov IK, Schneppat UE, Reise H, Quang Vu M M (2020) First record of an
agriolimacid slug in Southeast Asia – Deroceras laeve (O. F. Müller, 1774) (Gastropoda:
Pulmonata) recently introduced to the Socialist Republic of Vietnam. Biodiversity Data
Journal 8: e59644. https://doi.org/10.3897/BDJ.8.e59644
• Dueñas M, Hemming D, Roberts A, Diaz-Soltero H (2021) The threat of invasive
species to IUCN-listed critically endangered species: A systematic review. Global
Ecology and Conservation 26: e01476. https://doi.org/10.1016/j.gecco.2021.e01476
• Faberi AJ, López AN, Manetti PL, Clemente NL, Álvarez Castillo HA (2006) Growth and
reproduction of the slug Deroceras laeve (Müller) (Pulmonata: Stylommatophora) under
controlled conditions. Spanish Journal of Agricultural Research 4 (4): 345‑350. https://
doi.org/10.5424/sjar/2006044-211
• Gittenberger E, Gyeltshen C, Leda P, Zangpo T, van Klinken RD (2018) The first record
of the cosmopolitan slug Deroceras laeve (O. F. Müller, 1774) (Gastropoda: Pulmonata:
Agriolimacidae) in Bhutan. Folia Malacologica 26: 89‑93. https://doi.org/10.12657/
folmal.026.006
• Gomes SR, Thomé JW (2004) Diversity and distribution of the Veronicellidae
(Gastropoda: Soleolifera) in the Oriental and Australian biogeographical regions.
Memoirs of the Queensland Museum 49 (2): 589‑601. URL: https://biostor.org/
reference/224707
• Guindon S, Dufayard JF, Lefort V, Anisimova M, Hordijk W, Gascuel O (2010) New
algorithms and methods to estimate maximum-likelihood phylogenies: Assessing the
performance of PhyML 3.0. Systematic Biology 59 (3): 307‑321. https://doi.org/10.1093/
sysbio/syq010
• Havlác JC (2004) A new record of Deroceras laeve (O. F. Müller, 1774) from Pakistan
(Gastropoda: Pulmonata: Agriolimacidae). Folia Malacologica 12: 181‑182. https://
doi.org/10.12657/folmal.012.016
Non-native terrestrial slugs from Sinaloa, Mexico: Deroceras laeve (O. ... 11

• Hirano T, Kagawa O, Fujimoto M, Saito T, Uchida S, Yamazaki D, Ito S, Shariar SM,
Sawahata T, Chiba S (2022) Species identification of introduced veronicellid slugs in
Japan. PeerJ 10: e13197. https://doi.org/10.7717/peerj.13197
• Lefort V, Longueville J, Gascuel O (2017) SMS: Smart Model Selection in PhyML.
Molecular Biology and Evolution 34 (9): 2422‑2424. https://doi.org/10.1093/molbev/
msx149
• López B, Zúñiga G, Mejía O (2019) Phylogeographic structure in the apparent absence
of barriers: a case study of the Mexican land snail Humboldtiana durangoensis
(Pulmonata: Humboldtianidae). Journal of Molluscan Studies 85: 244‑252. https://
doi.org/10.1093/mollus/eyz007
• Maceira F. D (2003) Las especies de la familia Veronicellidae (Mollusca, Soleolifera) en
Cuba. Revista de Biología Tropical 51 (3): 453‑461. URL: https://www.redalyc.org/
articulo.oa?id=44911879018
• Naranjo-García E, Thomé WJ, Castillejo JA (2007) Review of Veronicellidae from
Mexico (Gastropoda: Soleoliferea). Revista Mexicana de Biodiversidad 78: 41‑50. URL:
http://www.scielo.org.mx/scielo.php?script=sci_arttext&pid=S1870-
34532007000100004&lng=es&nrm=iso
• Naranjo-García E, Castillo-Rodríguez ZG (2017) First inventory of the introduced and
invasive mollusks in Mexico. The Nautilus 131 (2): 107‑126.
• Nurinsiyah AS, Hausdorf B (2018) Listing, impact assessment and prioritization of
introduced land snail and slug species in Indonesia. Journal of Molluscan Studies 85
(1): 92‑102. https://doi.org/10.1093/mollus/eyy062
• Rueda A, Caballero R, Kaminsky R, Andrews KL (2002) Vaginulidae in Central America,
with emphasis on the bean slug Sarasinula plebeia (Fischer). In: Barker GM (Ed.)
Molluscs as Crop Pests. CABI Publishing, New York. [ISBN 9780851993201]. https://
doi.org/10.1079/9780851993201.0115
• Strebel H, Pfeffer G (1880) Beitrag zur Kenntniss der Fauna mexikanischer Land-und
Süsswasser-conchylien. IV. Theil, 1-112; pls. 1-15 pp. [In German]. URL: https://
www.biodiversitylibrary.org/item/106422#page/7/mode/1up
• Sysoev A, Schileyko A (2009) Land snails and slugs of Russia and adjacent countries.
Pensoft Publishers, Sofia/Moscow, 455 pp. [ISBN 9546424749]
• Thomé JW (1989) Annotated and illustrated preliminary list of the Veronicellidae
(Mollusca: Gastropoda) of the Antilles, and Central and North America. Journal of
Medical and Applied Malacology 1: 11‑28.
• Thomé JW (1993) Estado atual da sistematica dos Veronicellidae (Mollusca:
Gastropoda) americanos, com comentários sobre sua importancia económica,
ambiental e na saúde. Biociencias 1: 61‑75.
• Wiktor A (2000) Agriolimacidae (Gastropoda: Pulmonata) a systematic monograph.
Annales Zoologici 49: 347‑590. URL: http://rcin.org.pl/Content/57364/
WA058_74192_P255-T49_Annal-Zool-Nr-4.pdf
12 Alvarez-Cerrillo L et al

Supplementary material
Suppl. material 1: GenBank accession numbers of sequences used in the
phylogenetic inference of Deroceras laeve (Fig. 2)
Authors: Araiza-Gómez Victoria, Alvarez-Cerrillo Laura Regina, Yáñez-Rivera Beatriz
Data type: GenBank accession numbers of sequences, localities
Brief description: In this table the species, number of sequences of GenBank and localities of
the specimens are listed. In total were used 143 sequences: 20 for Deroceras invadens, 101 for
D. laeve, 20 for D. reticulatum and two for outgroup Limax maximus.
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Non-native terrestrial slugs from Sinaloa, Mexico: Deroceras laeve (O. ... 13