taxonID	type	description	language	source
03B387D81C179F1C258DEBFDFBDBB8DB.taxon	description	Preserved specimens were light-yellowish or cream with smooth (i. e. not prickly) skin (Fig. 3 A). Lateral bands were generally more visible on preserved individuals than on living specimens. Dark dots above the anterior foot fringe were also apparent in most preserved specimens, although they could be faint or even absent (Fig. 3 B). For anatomical confirmation of identity, it was already informative that fully developed genitalia were found in animals of this size, ruling out most other Arion species, which mature at a larger size. Other distinctive anatomical characters were the large and round bursa copulatrix with a short, thick peduncle, and the short and tubular oviductus (Fig. 3 C – E). In Hungary we collected A. intermedius specimens in October and they laid eggs at the beginning of November. The eggs were 2 – 3 mm in diameter, white, and were laid in clutches typically of about 10 eggs (Fig. 4 A). Freshly hatched specimens were about 3 mm long. Hatchlings and juveniles were similar in external appearance to adults, and the prickly appearance was visible from hatching (Fig. 4 B). The only known locality of A. intermedius in Hungary remains the Kámoni Arborétum, Szombathely (Fig. 5 A), outside but close to the part open to the public. We found the slugs at three locations within 50 m of each other on both sides of the Gyöngyös stream, which originates over the border in Austria. The slugs were found under the bark of decaying logs amongst deep leaf litter in woodland of mainly Alnus glutinosa and Acer platanoides (Fig. 6 A). The partial-COI sequences of two individuals from this locality were identical. The most similar haplotype in GenBank differs by just one nucleotide and had been found in A. intermedius from two distant sites in the eastern USA (GenBank EU 382743.1 and MG 813885.1).	en	Turóci, Ágnes, Hutchinson, John M. C., Schlitt, Bettina, Vidéki, Róbert, Farkas, Roland, Németh, Márk Z., Borostyán, Katalin, Páll-Gergely, Barna (2025): First records of Arion intermedius and Arion transsylvanus (Stylommatophora: Arionidae) in Hungary. Ecologica Montenegrina 83: 12-28, DOI: 10.37828/em.2025.83.2, URL: https://doi.org/10.37828/em.2025.83.2
03B387D81C1C9F0F258DEC9EFCBEB93D.taxon	description	The 455 - bp sequence was a close match (0.2 – 1.7 %) to the 14 A. transsylvanus 16 S haplotypes generated by Jordaens et al. (2010). In particular the Hungarian sequence differed by a substitution of just 1 bp from the 402 - bp haplotype AY 860806.1, from an animal collected in eastern Poland. In comparison, Jordaens et al. (2010) reported a difference of 15 % to the other species in the A. subfuscus species complex. The juvenile Hungarian specimen was 45 mm when fully extended. The colouration was a particularly vivid orange, especially on the back (Fig. 7). This is not diagnostic since Jordaens et al. (2010) documented wide and overlapping ranges of colouration in this species and its close relatives. One difference from their description of A. transsylvanus is that the foot fringe of our slug was light without dark stripes; this might reflect its juvenile state. The mucus was sticky, but not so much as that of Arion vulgaris, for instance. After preservation our specimen was approximately half as long as when living (25 mm after preservation). The body had become light brown with a pinkish tinge, and the dark markings showed up more conspicuously (Fig. 8). The single specimen proved to be too juvenile to conclude anything about species identity from the genital anatomy. The site at which we found A. transsylvanus in Hungary is on the plateau of the Bükk Mountains (Fig. 5 B, Table 1) at a relatively high altitude (above 800 m a. s. l. compared to the highest point of the country: 1014 m, Kékestető). It was found in a natural habitat in a beech forest. Although forestry activity takes place in the area, the locality is fairly undisturbed. The specimen was found under deep leaf litter in the stump of a beech tree (Fig. 6 B). After the molecular data revealed the presence of A. transsylvanus, we (RF & BPG) returned to the original site in May 2024 to collect more specimens. All four found were juveniles and the single example sequenced had the COI mtDNA sequence of A. fuscus.	en	Turóci, Ágnes, Hutchinson, John M. C., Schlitt, Bettina, Vidéki, Róbert, Farkas, Roland, Németh, Márk Z., Borostyán, Katalin, Páll-Gergely, Barna (2025): First records of Arion intermedius and Arion transsylvanus (Stylommatophora: Arionidae) in Hungary. Ecologica Montenegrina 83: 12-28, DOI: 10.37828/em.2025.83.2, URL: https://doi.org/10.37828/em.2025.83.2
03B387D81C1C9F0F258DEC9EFCBEB93D.taxon	discussion	Discussion The small size of Arion intermedius makes it is easy to overlook or to casually dismiss as a juvenile of another species. Nevertheless it is a species that can often be reliably identified on the basis of external characters, particularly the prickly appearance and the tiny dots along the foot fringe. However, these characters are not always apparent, and in particular preserved slugs often look smooth; in those cases, dissection or sequencing is needed for reliable identification. Within Hungary, the possibility of misidentification is perhaps greatest with Arion obesoductus Reischütz, 1973 formerly referred to as Arion alpinus Pollonera, 1887. Many populations from Central Europe and the Eastern Alps turned out to be A. obesoductus, and A. alpinus was synonymised with A. intermedius (Manganelli et al. 2010). Although A. obesoductus has not been recorded from Hungary, it is not improbable that it does or will occur. Arion obesoductus is known from Czechia (Dvořák et al. 2006), eastern Germany (Schniebs & Reise 1997) and Austria (Duda et al. 2019). More brownish colour morphs of A. intermedius resemble A. obesoductus, although the latter lacks the prickly appearance and dots along the foot sole. Internally, A. intermedius has a short, thin and tubular oviduct, whilst that of A. obesoductus is longer and wider. Also the bursa copulatrix differs: large and spherical with a short and wide peduncle in A. intermedius and elongated and thin without a prominent transition into its peduncle in A. obesoductus. Certain species of the subgenus Kobeltia (Arion hortensis and Arion distinctus) might also be confused with a dark-grey A. intermedius. If such slugs lack the diagnostic external characters of A. intermedius, Kobeltia species can be ruled out by their longer and thicker oviduct, which consists of more parts (A. distinctus 2 or 3 parts; A. hortensis always 3 parts) (Backeljau & De Bruyn 1990). Surprisingly, one of the biggest species of Arion, Arion ater can also superficially resemble A. intermedius in external appearance. Hatchlings of A. ater overlap A. intermedius in size and may also have straw-coloured bodies with dark tentacles. The proportions and, later in development, the prominence of the foot fringe give a different general impression, but those without experience of both species should not identify an A. intermedius lacking the prickly tubercles or the small dots without checking for maturing genitalia. However, A. ater is rare or extinct in Hungary (Turóci & Páll-Gergely in press). In other genera, juveniles of Krynickillus melanocephalus and certain colour morphs of some Deroceras species might look superficially similar to A. intermedius but these can be immediately ruled out by the position of the pneumostome relative to the mantle (posterior in Agriolimacidae, anterior in Arionidae) and the shape of the tail (pointed in Agriolimacidae, rounded in Arionidae). The life cycle of A. intermedius in the wild is one year. In southern England, individuals are highly synchronised in when they mature, in September (Hutchinson et al. 2017). The species was believed to reproduce only by selfing, based on genetic analyses (Foltz et al. 1982) and because of the lack of observations of spermatophores and mating (Davies 1977; Backeljau & De Bruyn 1990). Garrido et al. (1995) reported the presence of spermatophores for the first time, and the later discovery of some heterozygosity of allozymes implies that occasional outcrossing does occur (Reise et al. 2001; Jordaens et al. 2013). A revisit to the collecting site of A. intermedius (by RV) in December 2024 resulted in the rediscovery of one living specimen (not collected). The occurrence in winter corresponds with studies elsewhere in its range (Hutchinson et al. 2017), but also December 2024 was mild even though the area lies in the vicinity of the eastern slopes of the Alps. That month in Szombathely, on 20 days the minimum temperature fell below freezing but none were colder than − 5 ° C (AccuWeather). So possibly a population could persist over winter in that area.	en	Turóci, Ágnes, Hutchinson, John M. C., Schlitt, Bettina, Vidéki, Róbert, Farkas, Roland, Németh, Márk Z., Borostyán, Katalin, Páll-Gergely, Barna (2025): First records of Arion intermedius and Arion transsylvanus (Stylommatophora: Arionidae) in Hungary. Ecologica Montenegrina 83: 12-28, DOI: 10.37828/em.2025.83.2, URL: https://doi.org/10.37828/em.2025.83.2
03B387D81C1C9F0F258DEC9EFCBEB93D.taxon	description	Arion transsylvanus might also be confused with species besides those in the same complex. Certain colour morphs of Arion vulgaris juveniles can be similar to this species. Generally, A. vulgaris juveniles are more reddish than A. transsylvanus, which is usually yellowish with some orange tinge. The lineolation on the foot fringe is more prominent in A. vulgaris juveniles. It can be useful to consider the size of the tubercles; they are larger and the grooves between them are deeper in A. vulgaris, which results in a coarser skin surface compared with A. transsylvanus. The lateral bands are typically also stronger and wider in A. vulgaris. According to current knowledge, A. transsylvanus is predominantly a Carpathian species, mostly restricted to the Transylvanian region in Romania (eastern and southern Carpathian Mountains and the Apuseni Mountains). The species is the only known representative of the Arion subfuscus species complex in Transylvania (Jordaens et al. 2010). Besides the Romanian records, Jordaens et al. (2010) reported this species only from one site in eastern Poland. We are not aware of published records of the species since then, which may partly reflect a lack of confidence amongst malacologists in determining it anatomically and a lack of resources for DNA sequencing. However, Gural-Sverlova & Gural (2015) found no A. transsylvanus, only A. fuscus, in Ukraine, even though Ukraine bridges the gap between the Romanian and Polish records. Nevertheless, the absence of records between our finding in northern Hungary and the nearest locality in Romania (c. 200 km away) does not dissuade us from proposing that the species might be native in Hungary. One reason is that the Bükk Mountains are geographically part of the Western Carpathians. Furthermore, we found this species at a relatively high altitude (c. 800 m) in little-disturbed beech forest, which is consistent with occurrence data in Romania (Grossu 1970; Jordaens et al. 2010). Our collecting site was remote from settlements (along a hiking track c. 3 km from the village Répáshuta), which would be the most likely point of introduction. If it is indeed native in Hungary, A. transsylvanus would be the only such one of the nine slug species that we have either discovered as new to Hungary in the last six years or that were considered doubtful records until we confirmed them; the other eight species are very likely introductions. Besides A. intermedius and A. transsylvanus, these “ new ” species are: Ambigolimax parvipenis Hutchinson, Reise & Schlitt, 2022; Ambigolimax valentianus (A. Férussac, 1821); Deroceras invadens Reise, Hutchinson, Schunack & Schlitt, 2011; Krynickillus melanocephalus Kaleniczenko, 1851; Limacus maculatus (Kaleniczenko, 1851); Milax nigricans (Philippi, 1836); and Tandonia kusceri (H. Wagner, 1931). On the other hand, in Romania A. transsylvanus occurs also in synanthropic and other degraded habitats (Jordaens et al. 2010), so there is the potential that it has or will spread to such environments in countries adjacent to Romania, including Hungary. The discovery of these two species of Arion may provide a guide to predicting what sort of further slug species might turn up new to the Hungarian fauna. Most likely are synanthropic species that spread as a result of trade and usually turn up in habitats like gardens, perhaps facilitated by the milder winters. A prediction for such a species is Deroceras panormitanum (Lessona & Pollonera, 1882) s. s., which has recently been reported in France (Ventura et al. in press). But additionally perhaps further native species have been overlooked in natural habitats in Hungary; these species may already be known from neighbouring countries. For instance, the genus Lehmannia has a centre of diversity in Romania, but the species are particularly hard to find during daylight and their identification requires dissection. Acknowledgements Thanks to Fruzsina Matolcsi for helping to sequence our A. intermedius specimens, to Heike Reise for invaluable discussions and help, and to the referees Ben Rowson and Ivaylo Dedov for their most useful suggestions to improve the manuscript. Funding Declaration This study was supported by the Scientific Patronage Grant (Tudományos Mecenatúra Pályázat, MEC _ K 140853) funded by the National Research, Development and Innovation Office and the Ministry of Innovation and Technology to BPG. The funder had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript. The Senckenberg Museum of Natural History Görlitz paid for some of the sequencing. Authors’ Contribution ÁT & BPG initiated the concept. RV, ÁT, RF and BPG collected the specimens. BS, MZN & KB were responsible for the sequencing. RV constructed the maps. ÁT wrote the manuscript, and all authors contributed to the final version, especially JMCH who corrected and edited the drafts.	en	Turóci, Ágnes, Hutchinson, John M. C., Schlitt, Bettina, Vidéki, Róbert, Farkas, Roland, Németh, Márk Z., Borostyán, Katalin, Páll-Gergely, Barna (2025): First records of Arion intermedius and Arion transsylvanus (Stylommatophora: Arionidae) in Hungary. Ecologica Montenegrina 83: 12-28, DOI: 10.37828/em.2025.83.2, URL: https://doi.org/10.37828/em.2025.83.2
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