Abstracts of some papers by Heike Reise
I work in the Malacology Department in the Senckenberg Museum für Naturkunde Görlitz.
Email: Heike.Reise@senckenberg.de
Tel.: (+49) (3581) 47605410
Fax: (+49) (3581) 47605499 (please announce by phone or email)
Address for post:
Heike Reise, Senckenberg Museum für Naturkunde Görlitz, Am Museum 1, 02826 Görlitz, GERMANY
H. Reise & M. Hanelt 1991. Ökofaunistik und populationsdynamische Aspekte der Kleinsäuger eines Basaltberges in der Oberlausitz. In: M. Stubbe, D. Heidecke & A. Stubbe: Populationsökologie von Kleinsäugerarten. Wissenschaftliche Beiträge der Martin-Luther-Universität Halle-Wittenberg 1990/34: 299–309.
Ecofaunistic and population dynamic aspects of the small mammal community of a basaltic mountain. Species composition, abundance and population dynamics were studied on a basaltic mountain near Görlitz by snap-trapping in 1986 and 1989. Presence of Clethrionomys glareolus and Apodemus agrarius are discussed. C. glareolus (51,4%), Apodemus flavicollis (27,1%) and Sorex araneus (12,6%) are the most frequent species. In some species few individuals only were caught, mostly on the treeless xerothermous slope. A competitive superiority of C. glareolus to A. agrarius in the area’s forest habitats is supposed.
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H. Reise & T. Backeljau 1994. Deroceras panormitanum (Lessona & Pollonera, 1882), sensu Giusti, 1986 in Ostsachsen (Gastropoda, Stylommatophora, Agriolimacidae). Abhandlungen und Berichte des Naturkundemuseums Görlitz 68(1): 71–76.
The slug species Deroceras panormitanum (Lessona & Pollonera) was found in the town of Görlitz. This represents the first report of this species in Eastern Saxonia and the most eastern population in Germany. Although the slugs were found at only one single site, it seems as is this population already exists there since at least 1991. The animals and the locality are described.
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H. Reise 1995. Mating behaviour of Deroceras rodnae Grossu & Lupu, 1965 and D. praecox Wiktor, 1966 (Pulmonata, Agriolimacidae). Journal of Molluscan Studies 61: 325330.
The mating behaviour of Deroceras rodnae Grossu & Lupu, 1965 and D. praecox Wiktor, 1966 was observed under laboratory conditions. Both species differ by the shape and functioning of the sarcobelum, the timing and duration of courtship and the colour of the sperm mass. Particularly the different timing of the courtship behaviour seems to act as an effective prezygotic isolation mechanism preventing interspecific crosses. Usually, sperm masses are exchanged mutually. In three homospecific pairings, unilateral sperm transfer was observed followed by a characteristic behaviour of the donor slug. This observation is discussed in light of the hermaphrodite’s dilemma and sperm trading models.
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H. Reise 1997. Deroceras juranuma Mendelian colour morph of D. rodnae (Gastropoda: Agriolimacidae). Journal of Zoology 241: 103115.
The terrestrial slug Deroceras juranum Wüthrich, 1993 (Gastropoda: Agriolimacidae) has been described as an endemic species of the Swiss Jura Mountains (Switzerland) where it occurs at high altitudes. The species description was mainly based on its characteristic blackish-violet body colour. Breeding experiments and comparative studies of the genital anatomy and mating behaviour indicate that D. juranum is nothing but a colour morph of the usually cream-coloured D. rodnae Grossu & Lupu, 1965. The body colour seems to be coded by a single locus with two alleles, of which violet is dominant over cream. Topographical colour variation has also been observed in arionid slugs. Its possible biological (i.e. selective?) meaning is discussed. The slugs are able to reproduce uniparentally with their offspring showing recombination.
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H. Reise, J.M.C. Hutchinson, R.F. Forsyth & T. Forsyth 2000. The ecology and rapid spread of the terrestrial slug Boettgerilla
pallens in Europe with reference to its recent discovery in North America.
The Veliger 43:313318.
The terrestrial slug Boettgerilla pallens Simroth, 1912, is reported
from two sites on Vancouver Island, British Columbia, the first records
for this Palaearctic species in America. This paper describes how to recognise
the species, and summarises European studies of its ecology. It is unusually
worm-like in appearance, lives mostly underground, and occurs in a very
wide range of habitats. This century the species has spread remarkably
far and fast across Europe from the Caucasus. This is demonstrated by a
table of first occurrences in each country, and by three case studies of
spread within Great Britain, Belgium and north-west Austria. We predict
that it will spread rapidly in North America, and may already occur more
widely, but there is no evidence that it will become an important pest.
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R.G. Forsyth, J.M.C. Hutchinson & H. Reise 2001. Aegopinella nitidula (Draparnaud, 1805) (Gastropoda: Zonitidae) in British Columbiafirst confirmed North American record. American Malacological Bulletin 16:6569.
The European land snail, Aegopinella nitidula (Draparnaud, 1805), is reported for the first time from British Columbia, from three sites in the city of Vancouver. These new records are the only documentation of the species in North America, except for two old records that are probably erroneous and have been ignored in recent literature. Comparisons are made between A. nitidula and similar native and introduced species. Information about its ecology in Europe is summarised.
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H. Reise & J.M.C. Hutchinson 2001. Morphological variation in terrestrial slug Deroceras turcicum (Simroth, 1894), and a northern extension of its range in Central Europe. Folia Malacologica 9:6371.
We report seven new localities of the terrestrial slug Deroceras turcicum (Simroth), including the first occurrences in the Czech Republic and Slovakia. These demonstrate that the distribution of D. turcicum extends much further north than known previously and it might even be rather common there. We discuss possible reasons for its belated discovery. D. turcicum might have often been confused not only with the common D. reticulatum (O.F. Müll.) but also with the syntopical D. rodnae Grossu et Lupu and D. praecox Wiktor. We describe its variability in size, coloration, genital morphology, and caecum length, and consider characters for its reliable discrimination from other species. Two colour morphs (white and violet) are described for the first time. Intriguingly, the coloration of D. turcicum often matches that of congeners found at the same site, which suggests that colour has some selective value. Since external appearance often gives no reason to suspect that two species are present at a site, dissection of several specimens is advisable.
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H. Reise, M. Benke & J.M.C. Hutchinson 2002. A sinistral specimen of the terrestrial slug Arion lusitanicus (Gastropoda: Pulmonata: Arionidae). Malakologische
Abhandlungen Staatliches Museum für Tierkunde Dresden 20:247252.
A juvenile sinistral specimen of Arion lusitanicus was found near Münster (Germany, Nordrhein-Westfalen) and raised in isolation. Its entire external and internal morphology is a mirror image of normal dextral slugs. In mating experiments with dextral individuals the partners showed clear courtship behaviour but were never successful. This was probably caused by the opposite position of their genital pores, which might make copulation impossible for mirror-image slugs. Reports of sinistral terrestrial slugs are very rare. The authors review earlier cases, and discuss the possible genetic basis of sinistrality in slugs.
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H. Reise, J.M.C. Hutchinson, R.G. Forsyth & T.J. Forsyth 2005. First records of the terrestrial slug Deroceras turcicum (Simroth, 1894) in Poland. Folia Malacologica 13:177179.
Deroceras turcicum (Simroth) is reported from six woodland sites around Wałbrzych in southwest Poland.
This extension of the species range to Poland was expected given the number of reports from adjacent areas of the Czech Republic. We collate these reports as well as local records of Deroceras praecox Wiktor, 1966, which is found in similar habitats. We briefly discuss the difficulty of distinguishing D. turcicum from Deroceras reticulatum (O. F. Müll.), with which it may also co-occur.
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H. Reise, J.M.C. Hutchinson & D.G. Robinson 2006. Two introduced pest slugs: Tandonia budapestensis new to the Americas, and Deroceras panormitanum new to the Eastern USA. The Veliger 48:110115.
This paper reports new findings in North America of two pest slugs from Europe. Tandonia budapestensis, previously unknown from America, was found in Washington DC and near Philadelphia. Deroceras panormitanum, unreported from the Eastern United States and from Eastern North America outside of greenhouses, was found in Washington DC. We describe how to recognize these species and briefly summarize knowledge of their distribution and ecology.
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H. Reise, S. Visser & J.M.C. Hutchinson 2007. Mating behaviour in the terrestrial slug Deroceras gorgonium: is extreme morphology associated with extreme behaviour? Animal Biology 57:197215.
Mating in Deroceras consists of an investigation phase (precourtship), then a long courtship involving mutual stroking with the extruded sarcobelum, then sperm exchange (copulation). The penial gland, if present, everts over the partners skin during copulation: this is hypothesised to apply a secretion manipulating the partner to use received sperm. Deroceras gorgonium has a particularly large penial gland, divided into many finger-like branches. We studied D. gorgonium mating behaviour in the hope of further indications of the glands function. Precourtship and courtship together last longer than in other Deroceras (c. 6 h to >9 h); precourtship is highly variable, often with many bouts of different behaviours, including seemingly inactive phases. During most of the courtship partners remain apart waving their particularly long, pointed sarcobela; only at a later stage do the tips of these contact the partner. This waving alternates with circling for half a turn. For the first time in Deroceras we observed the sarcobelum transferring a secretion. The copulation is amongst the fastest: genital eversion and sperm exchange occur within 1 s, and slugs separate 1825 s later. The penial gland is everted immediately after sperm exchange, but, surprisingly, is often spread underneath the partner rather than over its back and, if on top, is not always fully spread over the partners body. We discuss these observations with respect to penial gland morphology and in the light of possible sexual conflicts. The long courtship and distant sarcobelum waving might reflect attempts to transfer, but not receive, secretion, and the circling might serve size assessment.
doi link from Ingenta
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The genus Deroceras Rafinesque, 1820 (the largest genus of terrestrial slugs) shows a high diversity of penis morphologies and mating behaviors. The function of most of the appending external and internal penis structures, some of them truly bizarre, is largely unknown. This paper reviews mating behavior and reproduction, based on data on 16 species from the literature and unpublished observations. I analyze patterns common to all Deroceras species, and differences among species. The general mating pattern consists of a long courtship with mutual stroking with a sarcobelum, a sudden penis eversion and external sperm exchange (copulation). I distinguish also precourtship and withdrawal phases. Sperm exchange is usually very quick but, in a few species, occupies a considerable proportion of the total mating duration. Mutual sperm exchange is the rule. Species differences involve the durations of certain mating phases, presence and nature of initial trail following, nature and intensity of stroking (including the degree of contact with the sarcobelum), aggressiveness of courtship behavior, and the timing of the penial gland eversion. I hypothesize that the radiation of mating behaviors and associated structures may have been driven by an arms race resulting from conflicting interests of mating partners over sperm donation and use. This could also have increased the rate of speciation in Deroceras. There are indications for the presence of sperm competition and conflicting interests between mating partners: individuals mate repeatedly, can store and digest sperm, and simultaneously use sperm from different mating partners for egg fertilization. Some details of mating behavior also suggest conflict. The timing of the penial gland eversion after sperm exchange suggests a manipulation akin to the role of love darts in helicid snails. Finally, some recommendations for studying mating behavior in Deroceras are given.
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J.M.C. Hutchinson & H. Reise 2009. Mating behaviour clarifies the taxonomy of slug species defined by genital anatomy: the Deroceras rodnae complex in the Sächsische Schweiz and elsewhere. Mollusca. 27:183200.
Collections of Deroceras from the uplands south of Dresden (along the Czech-German border) revealed two similar species differing in mating behaviour. The ranges interdigitate, but the species never occurred together. Species A has a wide sarcobelum held leftwards over its head and the everted penes are fully visible from above. It most resembles Deroceras praecox, occurring 100 km further east, whose anatomy and courtship behaviour are nevertheless consistently distinct; if these are different species, species A appears endemic to the Sächsische Schweiz. In species B, courtship and copulation take longer. Its sarcobelum is narrower, with a much enlarged base, and is directed forward or to the right. Most distinct is that the penes evert downwards and coil round each other for an additional revolution; their eversion is hidden from above except for the hand-like penial gland. This species is conspecific with Swiss, German and Austrian populations of Deroceras rodnae, but distinct from eastern populations, which more closely resemble D. praecox and species A. Unpublished molecular analyses support this division. Nevertheless, identifying non-mating animals by genital anatomy can be difficult: the base of the sarcobelum provides the best character. Western populations of D. rodnae should be termed Deroceras juranum Wüthrich, 1993.
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M. Benke, H. Reise, K. Montagne-Wajer & J.M. Koene 2010. Cutaneous application of an accessory-gland secretion after sperm exchange in a terrestrial slug (Mollusca: Pulmonata). Zoology 113:118124.
Competition for fertilisation in hermaphroditic animals seems to have led to many odd behaviours and complex morphologies involved in the transfer of accessory-gland products to the partner. Terrestrial slugs of the genus Deroceras show remarkably elaborate and interspecifically diverse penis morphologies and mating behaviours. Most species have an appending penial gland, which in Deroceras panormitanum consists of a few long fingers that are everted after sperm exchange and laid onto the partners back. To investigate whether this gland transfers a secretion onto the partners skin, we killed slugs at different mating stages and studied their penial glands and skin histologically. Two types of secretion granules appeared at a very early stage of courtship, and the penial gland was already filled 15 min into the courtship. At copulation, the gland everted this secretion onto the partners body, where it remained for at least 50 min. No lysis of skin tissue or other effects on the skin were observed. The slugs tried to lick the received secretion off their own body, and some droplets were observed to be shed with the body mucus. Our results indicate the external application of a glandular substance that could function as either a pheromone or allohormone. The behaviours of the recipients suggest sexual conflict, although mutual interest cannot be ruled out.
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H. Reise, J.M.C.Hutchinson, S. Schunack & B. Schlitt 2011. Deroceras panormitanum and congeners from Malta and Sicily, with a redescription of the widespread pest slug as Deroceras invadens n. sp. Folia Malacologica 19:201223.
The name Deroceras panormitanum is generally applied to a terrestrial slug that has spread worldwide and can be a pest; earlier this tramp species had been called Deroceras caruanae. Neither name is appropriate. The taxonomic descriptions apply to a species from Sicily and Malta. This true D. panormitanum and the tramp species are distinct in morphology and mating behaviour. For instance, the penial caecum of D. panormitanum is more pointed, everting faster at copulation. The size of the penial lobe varies considerably in preserved specimens but is always prominent at copulation. D. panormitanum is distinct from the Maltese endemic Deroceras golcheri, but a phylogeny based on COI mtDNA implies that they are more closely related than is the tramp species. D. golcheri has a still closer counterpart on Sicily, but we leave the taxonomy of this species X unresolved. In interspecific crosses, D. panormitanum may transfer sperm to the partners sarcobelum whereas the partner fails to evert its penis (D. golcheri) or to transfer sperm (the tramp species). Names previously applied to the tramp species originally referred to D. panormitanum or are otherwise invalid, so it is here formally redescribed as D. invadens. Deroceras giustianum Wiktor, 1998 is synonymised with D. panormitanum.
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E. Dreijers, H. Reise & J.M.C. Hutchinson 2013. Mating of the slugs Arion lusitanicus auct. non Mabille and A. rufus (L.): different genitalia and mating behaviours are incomplete barriers to interspecific sperm exchange. Journal of Molluscan Studies 79:5163.
The large slug known as Arion lusitanicus (or A. vulgaris) is an important pest that is spreading through much of Europe. Arion rufus disappears at sites where A. lusitanicus has established strong populations. The finding of morphological intermediates suggests that A. lusitanicus hybridizes with A. rufus, but interspecific mating had not been proven. Considering the marked differences in their genitalia, it has been hard to envisage how mixed couples might transfer sperm. Arion lusitanicus and A. rufus were collected from pure populations near Görlitz, Germany, and used for laboratory mating trials involving either two individuals of A. rufus (henceforth RR), two of A. lusitanicus (LL), or one of each species (mixed). Matings were video recorded and some couples were killed during or after copulation to study spermatophore transfer and genital anatomy during mating. Three mixed pairs copulated. However, mixed pairs were significantly less likely to copulate than either RR or LL pairs (7% vs 52% and 36%). At each stage of mating, the probability of proceeding further was lower in mixed pairs than predicted from rates in RR and LL pairs, but this effect was strongest for yin-yang formation and initiating copulation. One problem was that A. lusitanicus tried to circle after yin-yang formation, whereas A. rufus remained stationary. In this respect, and in the repositioning of its everted oviduct, it was A. lusitanicus that compromised. LL copulations lasted over twice as long as RR copulations, but spermatophore formation took similar times, permitting reciprocal spermatophore exchange in mixed couples even though their copulations ended much earlier than in LL pairs. Our observations of mating behaviour of intraspecific pairs largely agree with previous descriptions of A. rufus, but we discuss some discrepancies with the fuller descriptions available for A. lusitanicus.
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H. Reise 2013. Wie seziert man Nacktschnecken?
Mitteilungen der Deutschen Malakozoologischen Gesellschaft 88:2932.
The determination of terrestrial slugs usually demands dissection. However, the literature recommends very different and partly contradictory methods how to access important internal structures. This article explains which structures are important for slug determination and discusses advantages and disadvantages of different methods. The method described by Wiktor is explained and illustrated. My own recommendations are added.
Die Bestimmung von Nacktschnecken erfordert in der Regel eine Sektion des Tieres. In der Literatur werden sehr verschiedene und teilweise widersprüchliche Methoden für den Zugang zu wichtigen inneren Strukturen empfohlen. Dieser Beitrag erklärt, welche Strukturen für die Nacktschnecken-Determination wichtig sind and diskutiert Vor- und Nachteile verschiedener Methoden. Die von Wiktor beschriebene Methode wird im Detail erläutert und durch eigene Empfehlungen ergänzt.
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J.M.C. Hutchinson & H. Reise 2013. A persisting population of an introduced slug, Milax nigricans, in Dunkirk, France.
Mitteilungen der Deutschen Malakozoologischen Gesellschaft 89:3538.
A population of Milax nigricans was found on February 19th 2011 outside a block of flats in Dunkirk, France. It was persisting 21 months later. This is the second finding in the department of Nord. We review earlier occurrences of the species outside its normal range in the Mediterranean and illustrate the most important identification characters.
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A.M. Păpureanu, H. Reise, & A. Varga 2014. First records of the invasive slug Arion lusitanicus auct. non Mabille (Gastropoda: Pulmonata: Arionidae) in Romania.
Malacologica Bohemoslovaca 13:611.
The invasive pest slug Arion lusitanicus (also known as Arion vulgaris) has spread throughout much of Europe within the last 60 years, becoming one of the worst agricultural and horticultural pest species. It has now been detected in Romania for the first time. There are dense populations at two sites in Braşov County (2013) and another occurrence in Mureş County (2012). We review occurrences in neighbouring countries and discuss evidence that A. lusitanicus might have arrived in Romania earlier. Further research on the species spread in Romania and on its impact on the native fauna and flora is necessary.
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J.M.C. Hutchinson, H. Reise & D.G. Robinson 2014. A biography of an invasive terrestrial slug: the spread, distribution and habitat of Deroceras invadens.
Neobiota 23: 1764.
The article reviews distribution records of Deroceras invadens (previously called D. panormitanum and D. caruanae), adding significant unpublished records from the authors own collecting, museum samples, and interceptions on goods arriving in the U.S.A. By 1940 D. invadens had already arrived in Britain, Denmark, California, Australia and probably New Zealand; it has turned up in many further places since, including remote oceanic islands, but scarcely around the eastern Mediterranean (Egypt and Crete are the exceptions), nor in Asia. Throughout much of the Americas its presence seems to have been previously overlooked, probably often being mistaken for D. laeve. New national records include Mexico, Costa Rica, and Ecuador, with evidence from interceptions of its presence in Panama, Peru, and Kenya. The range appears limited by cold winters and dry summers; this would explain why its intrusion into eastern Europe and southern Spain has been rather slow and incomplete. At a finer geographic scale, the occurrence of the congener D. reticulatum provides a convenient comparison to control for sampling effort; D. invadens is often about half as frequently encountered and sometimes predominates. Deroceras invadens is most commonly found in synanthropic habitats, particularly gardens and under rubbish, but also in greenhouses, and sometimes arable land and pasture. It may spread into natural habitats, as in Britain, South Africa, Australia and Tenerife. Many identifications have been checked in the light of recent taxonomic revision, revealing that the sibling species D. panormitanum s.s. has spread much less extensively. A number of published or online records, especially in Australia, have turned out to be misidentifications of D. laeve.
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J.M.C. Hutchinson & H. Reise 2015. Mating in Ariunculus isselii, an arionid slug without a spermatophore. Journal of Molluscan Studies 81: 247258.
Ariunculus is a genus of terrestrial slugs often treated as a subgenus of Arion. A survey of the literature reveals doubts about many of the described species: only A. speziae and A. isselii are now generally recognized as belonging to Ariunculus. Ariunculus isselii is widespread on Sardinia. We describe its external appearance, mating behaviour and genital anatomy, and interpret the functioning of the genitalia based on specimens killed during copulation or shortly afterwards. Instead of the distinct epiphallus of Arion, there is only a small ampulla, because sperm are pumped out piecemeal to the recipient rather than transferred in a spermatophore. We speculate (in a framework of sexual selection theory) that this may explain why mating lasts 12 h after the initial genital eversion, much longer than in most Arion. The absence of a spermatophore provides grounds for keeping Ariunculus separate from Arion. Similarly to some Arion species, the papilla is inserted into the partners partially everted bursa trunk; we discuss four different hypotheses for the function of this arrangemet. Also as in some Arion species, the oviduct is everted during mating so as to apply a ligula. We provide information on egg laying in captivity and report that the species can self-fertilize.
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A. Ludwig, H. Reise & J.M.C. Hutchinson 2015. Die Nacktschneckenfauna in Gärten der Stadt Görlitz (Sachsen,
Deutschland). Berichte der Naturforschenden Gesellschaft der Oberlausitz 23: 4357.
The slug fauna of gardens in the town of Görlitz was investigated by hand searching 14 house gardens, 13 allotments and four courtyards. The 13 species that were found were, in decreasing order of occurrence, Arion lusitanicus auct. non Mabille, Deroceras reticulatum, Arion distinctus, Deroceras invadens, Arion fasciatus, Limax maximus, Boettgerilla pallens, Deroceras sturanyi, Arion rufus, Arion silvaticus, Lehmannia valentiana, Arion circumscriptus und Deroceras laeve. More species of slug occurred in gardens rated as untidier (p = 0.02). The discovery of L. valentiana is the first outdoors record for Görlitz. The diverse state of establishment of the other four invasive species is discussed in relation to earlier faunistic data. Our investigation provides a baseline for long-term monitoring of the development of the slug fauna in Görlitz.
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J.M.C. Hutchinson & H. Reise 2015. An invasion from Germany; Deroceras invadens (Pulmonata, Agriolimacidae) and other synanthropic slugs in the southwest corner of Poland. Folia Malacologica 23: 301307.
Ten towns in the southwest corner of Poland within 57 km of the German border were surveyed for slugs. Deroceras invadens was found only in the four westernmost towns. In Zgorzelec, which was surveyed more intensively than the other towns, it occurs widely but only sporadically. Although D. invadens had been known in the adjacent German town of Görlitz since 1991, hitherto the only Polish record was from Wrocław, 140 km to the east. This pattern suggests a new colonisation across the border. Amongst the other species, one surprise was the rarity of Deroceras reticulatum in several towns, and another that Arion fuscus was common in one town but not found in any of the others. Records of Deroceras praecox extend its known range in Poland further west. Contrary to expectations from the literature, Deroceras sturanyi was adult in spring, and reproducing at a small size typical of Deroceras laeve.
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J.M.C. Hutchinson, H. Reise & G. Skujienė 2017. Life cycles and adult sizes of five co-occurring species of Arion slugs. Journal of Molluscan Studies 83: 88105.
Five species of Arion slugs were collected repeatedly at a woodland site in southern England and all individuals weighed. Selected samples of these were dissected so as to weigh components of the reproductive tract. The relative weights of the gonad, spermoviduct and albumen gland provided the basis to categorize individuals into adult, subadult or immature classes, or as juvenile if the sum of these weights was below a threshold. This procedure was validated by raising A. subfuscus in captivity and killing at a range of known ages before and after egg laying. In the other species, organ weights from individuals observed to have laid eggs or mated also helped to calibrate the divisions. Such data from two species demonstrated that, following the production of an egg clutch, the albumen gland took days gradually to regrow. There was little evidence of much variation in life cycle from year to year and the broad patterns, although not precise timings, agreed with studies elsewhere. No species produced more than one generation per year and in all there was a season (brief in A. subfuscus) when adults were absent. The life cycles were predominantly annual, although in some species a minority of individuals might take 18 months to mature. The time of year at which individuals matured into adults varied between species: A. intermedius in August and September, A. distinctus mostly in December and January, A. circumscriptus mostly January to April, A. subfuscus April to early October and A. rufus July to September. The largest two species thus dominated in summer, but at other times the species overlapped considerably in size. In four species, individuals maturing later in the season did so at a smaller size; the possible exception was A. intermedius, in which maturation was highly synchronized. The coefficients of variation in adult size were compared against a collection of such data from other terrestrial molluscs. The smallest species, A. intermedius, had disproportionately large hatchlings.
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J.E. MAUNDER, R.G. NOSEWORTHY, J.M.C. HUTCHINSON & H. REISE 2017. Terrestrial molluscs of the Province of Newfoundland and Labrador, Canada. Part 1: Boettgerillidae. Checklist 13: 277284.
The family Boettgerillidae, represented by the Eurasian slug Boettgerilla pallens Simroth, 1912, is first recorded for Newfoundland and Labrador, Canada
—a range extension of almost exactly 5000 km within the Americas. Compiled, within an appendix, to provide a national perspective for the Newfoundland and Labrador record, are 13 previously
unpublished B. pallens records from British Columbia, Canada. Incidentally recorded is the second eastern Canadian outdoor occurrence of the European slug
Deroceras invadens. This paper is the first in a series that will treat all of the terrestrial molluscs of Newfoundland and Labrador.
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T. von Proschwitz, H. Reise, B. Schlitt & K. Breugelmans 2017. Records of the slugs Ariolimax columbianus (Ariolimacidae) and Prophysaon foliolatum (Arionidae) imported into Sweden. Folia Malacologica 25: 267271.
Two North American slug species are reported from three locations in southern Sweden: three records (in 2005, 2014 and 2015) of Ariolimax columbianus (Gould) and one 2005 record of Prophysaon foliolatum (Gould). The record of P. foliolatum is the first for Europe. In all cases it is highly probable or proven that the slugs had been accidentally imported with ornamental plants from North America, particularly salal, Gaultheria shallon Pursh. Salal is collected on a massive scale from the rainforests of the Pacific Northwest, particularly in Canada and Washington State, and exported as green filler for floral arrangements. The harvest, storage and transport conditions seem perfectly suitable for these slugs, which are not synanthropic but live in natural habitats where salal thrives. We point out that these apparently uncontrolled imports might open the door to the introduction of new, non-synanthropic species.
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H. Reise, B. Schlitt & J.M.C. Hutchinson 2018. Bericht über die 33. Regionaltagung des Arbeitskreises Ost der DMG in Ostritz bei Görlitz vom 23. bis 25. September 2016: Mollusken des Rotsteins bei Sohland, Sachsen. Mitteilungen der deutschen malakozoologischen Gesellschaft 98:3544.
Das 33. Regionaltreffen der DMG-Arbeitsgruppe Ost fand vom 23.-25.9.2016 im Internationalen
Begegnungszentrum St. Marienthal in Ostritz, Ostsachsen statt. Es kamen 35 Teilnehmer und Begleitpersonen.
Die Sammelexkursion mit einem Schwerpunkt auf Landgastropoden führte am 24. September an drei
Sammelstellen im Ostlausitzer Hügelland: das Naturschutzgebiet des Basaltbergmassivs Rotstein bei Sohland
am Rotstein, ein Teich und Erlenwald bei Sohland und der Basaltberg Eichler bei Rennersdorf. Weitere Aufsammlungen
wurden im Umfeld der Klosters St. Marienthal vorgenommen. Insgesamt wurden 81 Molluskenarten
gesammelt, davon 53 Arten am Rotstein, dem Schwerpunktgebiet der Tagungsexkursion. Zehn Arten wurden
zum ersten Mal auf dem Rotstein nachgewiesen. Insgesamt sind nun 61 Arten für den Rotstein nachgewiesen,
was den besonderen Stellenwert des Gebietes für die Molluskenfauna der Oberlausitz unterstreicht. Zu den
faunistisch besonders interessanten Arten gehören u. a. Vertigo alpestris, Euomphalia strigella, Truncatellina
cylindrica und Vitrea subrimata. Letztere ist gleichzeitig der Erstnachweis für die Oberlausitz, ebenso wie das
Leergehäuse einer Nesovitrea petronella aus dem Erlenwald am Teich bei Sohland. Weitere besonders bemerkenswerte
Funde sind Vitrinobrachium breve vom Eichler und Limacus flavus aus Ostritz, beide in der sächsischen
Oberlausitz erstmalig außerhalb der Stadt Görlitz. Leider wurde der invasive Arion lusitanicus auch im
Kerngebiet des Rotstein-Waldes angetroffen, was belegt, dass die Art nun in natürliche Lebensräume eindringt,
wo sie gegenwärtig neben einheimischen A. rufus agg. vorkommt.
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H. Reise, Z. Arslangündoğdu, B. Schlitt, J.M.C. Hutchinson, E. Hızal & E. Bacack 2018. First records of the terrestrial slug Arion ater s.l. (Linnaeus, 1758) (Pulmonata: Arionidae) from Turkey. Folia Malacologica 26: 213–220. 301307.
A strong population of the terrestrial slug Arion ater s.l. is reported from the European and
Asian parts of Istanbul, Turkey. This is the first confirmed report of this large, conspicuous taxon from
Turkey and from Asia. Our samples from five synanthropic sites indicate that it is already well established.
Partial sequences of the mitochondrial COI gene (cytochrome c oxidase subunit I) place the Turkish slugs
in a small clade shared with a few specimens from western France, perhaps indicating the origin of the
Istanbul population. The next closest haplotypes (9% difference) fall within the clade identified as Arion
ater s.s. This fits with the genital morphology of the Turkish slugs, which is most similar to the ater-form
of A. ater s.l. Our discovery also puts a new light on the recent report of the highly invasive pest slug Arion
lusitanicus auct. non Mabille, 1868 (often called Arion vulgaris Moquin-Tandon, 1855) in Isparta, which was
identified only on the basis of external morphology. As reliable morphological distinction of these two
species requires examination of the genital anatomy, the specimen from Isparta should be reinvestigated.
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J.M.C. Hutchinson, B. Schlitt & H. Reise 2019. Monacha claustralis (Rossmässler 1834), a hygromiid snail new to Germany. Mitteilungen der deutschen malakozoologischen Gesellschaft 100: 1722.
Colonies originally identified as Monacha cartusiana have often turned out to be M. claustralis. This was the case with a sample collected in 2016 from near Jena, and thus the first record of M. claustralis from Germany. We compare these animals’ genital anatomy against several distinguishing characters advocated in the literature. A partial-COI sequence indicates a close genetic relationship with Polish colonies near Gdansk and Kielce. Although originally from western Turkey and adjacent parts of the Balkan Peninsula, M. claustralis is liable to have established itself at other sites in Germany, maybe sometimes hidden in mixed colonies with M. cartusiana, from which it is indistinguishable using external characters.
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H. Reise, A.-K. Schwarzer, J.M.C. Hutchinson & B. Schlitt 2020. Genital morphology differentiates three subspecies of the terrestrial slug Arion ater (Linnæus, 1758) s.l. and reveals a continuum of intermediates with the invasive A. vulgaris Moquin-Tandon, 1855. Folia Malacologica 28: 134.
The terrestrial slug Arion vulgaris Moquin-Tandon (= A. lusitanicus auct. non Mabille) is an important agricultural pest that has invaded much of Europe. Previous work has demonstrated hybridisation with A. ater (Linnæus) s.l. We describe the genital anatomy of morphological intermediates found in eastern Saxony (Germany), comparing them with the parent species. We recommend a standard method of genital dissection and consider a set of five genital characters. The intermediates demonstrate the homology of the ligula-bearing organs of the two species; the ancestral position of the ligula in the dilated part of the oviduct has moved to the upper atrium in A. ater s.l. Furthermore we differentiate three morphotypes of A. ater s.l. in eastern Saxony, associated with different mitochondrial DNA sequences. One is A. ater ater, previously unrecognised from this part of Germany and the Czech Republic. The two other morphotypes correspond to the predominantly British and the Continental subspecies previously recognised on genetic and morphological grounds. We designate as the lectotype of Limax rufus Linnæus, 1758 a non-surviving specimen from Almondbury, England, described by Lister; thus the predominantly British subspecies becomes A. ater rufus. The appropriate name for the Continental form is A. ater ruber (Garsault, 1764).
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J.M.C. Hutchinson, B. Schlitt, T. Kořínková, H. Reise & G.M. Barker 2020. Genetic evidence illuminates the origin and global spread of the slug Deroceras invadens. Journal of Molluscan Studies 86: 306–322.
The terrestrial slug Deroceras invadens has spread across much of the world over the last century. What is to learn from mitochondrial sequences (partial COI) about the species’ origin, colonisations, and diversity? Samples from 317 localities covering most of its range yielded 87 haplotypes. Higher diversity, the predominance of private haplotypes, and geographic structuring all indicate a native range centred on southern Italy, including eastern Sicily. In contrast, central Italy is dominated by one haplotype, although accompanied by both close and more distant relatives; the lack of geographic structuring suggests recent expansion from a restricted distribution within that region. Beyond the Alps, two haplotypes predominate, accompanied by very similar variants; such star-shaped genealogies characterise recent population growth. Also some rarer haplotypes have been independently introduced. Generally haplotypes are well mixed here, often co-occurring at a locality. In North America and Australasia, some frequent haplotypes were likely directly introduced from Italy, because they were not found elsewhere in Europe. The rarity or absence in these continents of one or other of the two dominant European haplotypes suggests that import inspections have restricted repeated introductions. A skyline plot detects the recent demographic expansion but also indicates an earlier population decline in the native area. This may explain why the one-dimensional summary statistics FS and R2 did not signal population growth. A review of 41 other studies that used DNA to analyse invasions of terrestrial molluscs documents considerable diversity in methodology. Studies using genetic data to date recent invasions probably should adjust standard substitution rates upwards.
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Dedov, I.K., Schneppat, U.E, Reise, H. & Quang Vu, 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.
Several individuals of the terrestrial slug Deroceras laeve were collected in 2018 in the Hoàng Liên Son mountain range of northern Vietnam. The three specimens which were investigated anatomically were aphallic or hemiphallic. A partial COI sequence verified the species identity. This is the first discovery of D. laeve and also of the slug family Agriolimacidae on the Indochinese Peninsula. The collecting site is situated near a cable-car station and below a tourist complex on Fansipan mountain, both of which had just been built by a Swiss-Austrian company between 2013 and 2016. This and the fact that the species had not been found elsewhere in the surrounding area, although searched for thoroughly, indicate that D. laeve is most probably a recent introduction, potentially with building material from Austria or Switzerland.
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J.M.C. Hutchinson, H. Reise & B. Schlitt 2020. Mating behaviour and genital anatomy of Deroceras cecconii (Pollonera, 1896), a widespread but overlooked slug from Italy, now introduced to eastern Germany. Archiv für Molluskenkunde 149: 221236.
The terrestrial slug Deroceras cecconii was described from Vallombrosa, a forest in the hills above Florence, Italy. However, its validity has long been disbelieved. Fresh samples from the type locality agreed well with the original description and proved to be distinct in genital anatomy, mating behaviour and COI genetic sequence. Video recordings of the copulation showed that, besides the penial gland, the penis had 2 pockets that evert, the caecum and lobe. The caecum deposits sperm onto the partner’s penis but is short and not always apparent in the retracted state. Many other components of the mating behaviour are modifications of those observed in D. invadens, D. panormitanum and D. golcheri, but the molecular phylogeny, based on the mitochondrial COI gene, did not fully resolve their relationships. Genetically closest to D. cecconii was a slug from the island of Montecristo previously considered possibly to be D. golcheri. Deroceras golcheri and an undescribed close relative from Sicily are liable to be confused with D. cecconii anatomically, but D. cecconii is distinguished by the presence of a lobe and the concave saddle at the proximal end of the penis where the vas deferens inserts. The penis of D. cecconii is less similar to those of D. invadens and D. panormitanum, which both have a long caecum and, usually, lobe. Deroceras cecconii is common and widespread in much of Italy, although not the far south, occurring in a wide range of habitats. Also, a local but persistent population has been introduced to the town of Ostritz on the eastern border of Germany. The genetic uniformity amongst most populations raises the possibility that it might also be an introduction within part of its Italian range.
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Link to journal (paywalled): https://doi.org/10.1127/arch.moll/149/221-236
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J.M.C. Hutchinson, B. Schlitt & H. Reise 2021. One town’s invasion by the pest slug Arion vulgaris (Gastropoda: Arionidae): microsatellites reveal little introgression from Arion ater and limited gene flow between infraspecific races in both species. Biological Journal of the Linnean Society 134: 835850.
The terrestrial slug Arion vulgaris has recently spread across most of Europe, often causing the local extinction of resident populations of Arion ater s.l. The species hybridise, which leads to the prediction of massive introgression of A. ater genes into A. vulgaris. To test this, we utilised 16 microsatellite markers applied to samples of both species collected around Görlitz, Germany, during the invasion. Amongst A. vulgaris individuals with typical genitalia, an analysis using structure suggested that only 6% were appreciably admixed with local A. ater; admixture did not increase over the course of the invasion. Amongst the c. 4% of slugs with intermediate genitalia, microsatellites confirmed that often they were hybrids, their anatomy correlating with the estimated share of ancestry from each species. The microsatellites also distinguished the three subspecies of A. ater previously recognised on the basis of genital anatomy and mitochondrial DNA. The subspecies were not well mixed spatially, with A. a. ater in wilder places, and A. a. rufus never found in the Polish part of the town; nevertheless hybridisation between them was occurring. Unexpectedly, the microsatellites indicated three genetic races amongst A. vulgaris; these occurred in different districts and are only slowly mixing spatially and genetically.
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H. Reise, J. Simchen, C. Tluste & J.M.C. Hutchinson 2021. Ein vitales Vorkommen von Lucilla scintilla R. T. Lowe 1852 in Cottbus, Südost-Brandenburg (Gastropoda: Helicodiscidae). Mitteilungen der Deutschen Malakozoologischen Gesellschaft 106: 21–26.
A chance finding of a single Lucilla specimen in the town of Cottbus (southeast Brandenburg, Germany) initiated a targeted search at the same site. This revealed a population of living Lucilla scintilla, which is the first record from the state of Brandenburg that is firmly identified and supported by voucher specimens. The locality is a ruderal site on a railway area, consistent with earlier reports that the species seems to be rather synanthropic and thermophilic and prefers shady ground. Only sieving soil samples and the substrate attached to grass roots revealed more than single shells.
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J.M.C. Hutchinson, H. Reise & B. Schlitt 2022. Will the real Limax nyctelius please step forward: Lehmannia, Ambigolimax, or Malacolimax? No, Letourneuxia! Archiv für Molluskenkunde 151: 19–41.
In 1861 Bourguignat described Limax nyctelius, a species of terrestrial slug from Algeria. Although the surviving identifiable syntype specimen is Ambigolimax melitensis, the description and illustration unambiguously refer to the arionid Letourneuxia numidica, because of the position of the pneumostome. We designate the lectotype as the specimen that Bourguignat illustrated (now lost), requiring L. numidica to be known as Letourneuxia nyctelia. Unfortunately, the name nyctelia has meanwhile been used for 4 other species, occurring across 4 continents. Simroth and Pollonera referred to a North African species lacking a rectal caecum, so potentially a Malacolimax. Quick and others referred to a species of Ambigolimax presumed native in North Africa but known also from Scottish greenhouses, Elba, South Africa, Australia, and New Zealand; we name it A. waterstoni n. sp. Its long penis and lack of a penial appendage caused confusion with a Lehmannia s.s. species with a similar penis form from the Carpathian Mountains, extending also from Albania across Bulgaria. The structures inside the penis are very distinct from those of A. waterstoni. We name it L. carpatica n. sp. Lastly, the species invasive in Great Britain and California, and reported also from France, Greece, and Spain, including the Canary Islands, is named here Ambigolimax parvipenis n. sp. It also has no penial appendage, but the penis is very short. The length of the rectal caecum emerged as a useful identification character: usually it extends to, or beyond, the posterior tip of the visceral sac in Lehmannia s.s., whereas in Ambigolimax it stops well short, except that it often reaches almost to the tip in A. valentianus.
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U. Jueg, H. Reise, & H. Kelm 2022. Limacus maculatus (Kaleniczenko 1851) in Niedersachsen (Gastropoda: Limacidae). Mitteilungen der Deutschen Malakozoologischen Gesellschaft 107: 9–14.
A specimen of Limacus maculatus (Kaleniczenko 1851) was found in a garden in Streetz, Lüchow-Dannenberg. This is the first known occurrence in the German state of Niedersachsen and the third one in Germany. The locality and the accompanying mollusc fauna are described. We discuss current information about the distribution and spread of L. maculatus and indicate the differences to L. flavus (Linnaeus 1758).
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H. Reise 2022. Dr. Gisela Vater (3.1.1925 – 4.9.2021). Mitteilungen der Deutschen Malakozoologischen Gesellschaft 107: 61–65.
The malacologist Dr Gisela Vater died on the 4th September 2021. She was the first curator specifically of the mollusc collection at the Staatliches Museum für Naturkunde – Forschungsstelle – Görlitz (today the Senckenberg Museum of Natural History Görlitz). For many years, as well as curating the molluscs, she was also head of the department embracing all the collections and responsible for the exhibitions. Her interest in terrestrial slugs initiated the ongoing research focus on these gastropods in Görlitz as well as the collection of soft bodies.
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G. Kronenberg & H. Reise 2023. On the search for a conchological holy grail, the F.H.W. Martini (1729–1778) shell collection—a progress report Archiv für Molluskenkunde 152: 71–80.
The authors have attempted to locate the shell collection of the German conchologist Friedrich
Heinrich Wilhelm Martini (1729–1778). Martini often illustrated his own specimens in the Neues Systematisches
Conchylien-Cabinet, and thereby many later became referred to as types. For instance, this
is true for many species in the Stromboidea described by Röding in 1798. After Martini’s death, at least
part of his collection passed to Johann von Mattuschka, and then to the Nathusius family of Hundisburg,
near Magdeburg, Sachsen-Anhalt. In 1840 Louis Pfeiffer stated that the Martini collection was still there,
after which there is no firm information. Unfortunately, our detective efforts did not uncover the collection’s
current existence somewhere unrecognised, but an eyewitness report indicates that it might have
been destroyed in the early 1970s. However, there is no certainty yet, and we indicate a few tracks that
might still be followed.
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Link to journal (paywalled): https://doi.org/10.1127/arch.moll/152/071-080
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Á. Turóci, J.M.C. Hutchinson, B. Schlitt, H. Reise, M. Rapala & B. Páll-Gergely 2023. Five new introduced terrestrial slugs in Hungary. BioInvasions Records 12: 711–729.
Five terrestrial slug species are reported from Hungary, either for the first time or confirming earlier records that had been considered unreliable: Limacus maculatus, Deroceras invadens, Ambigolimax valentianus, Ambigolimax parvipenis, and Milax nigricans. In all cases identification was supported by anatomical examination and barcoding sequences of the COI mitochondrial gene. For M. nigricans we also sequenced individuals of this species from Italy and France so as to establish differences to sequences of M. gagates. Most records came from garden centres in Budapest and from two botanical gardens; these habitats may therefore be crucial in the spreading of non-indigenous species. A Facebook appeal was used to involve citizen scientists in searching for further sites for L. maculatus, but the Limacus populations reported proved all to be L. flavus. Studies over the last four years have added altogether seven slug species to the 26 hitherto recognised in the Hungarian fauna. This demonstrates the importance of continuing to check for additional introduced species.
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B.M.J. Williams, J.M.C. Hutchinson, H. Reise, O. Zauder & B. Schlitt 2024. Difficulties in distinguishing Monacha claustralis from M. cartusiana in Germany and Poland. Journal of Molluscan Studies 90: eyae030.
The two invasive snail species Monacha claustralis and M. cartusiana have been distinguished by their genital anatomy and mitochondrial haplotype. We assessed these distinctions in several populations from eastern Germany and northwestern Poland, as well as in reference populations (in part utilizing published illustrations). Absolute genital dimensions were unreliable characters because of their correlation with body size, which varies considerably within species. The ratio of epiphallus to vagina lengths was mostly effective in classifying animals from the reference populations, and it correlated closely with the second principal component of a multivariate analysis. It correlated also with the prominence of the vaginal sac, another identification character. However, even some individuals amongst the reference populations would have been misidentified by this ratio, and two German populations spanned a range of intermediate morphologies. These populations contained mitochondrial sequences of both species, but within-site variation in anatomy was not well explained by haplogroup; these seem to be populations of hybrids. The two Polish populations had anatomies of M. cartusiana, yet mitochondrial sequences of M. claustralis; surprisingly, samples from the same sites 11 years earlier were reported to have had M. claustralis anatomy. We warn that clear-cut identifications of Central European populations may increasingly often be unattainable and advocate investigation of whether the two taxa intergrade also in their native ranges.
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