An annotated checklist of millipede fauna from Slovakia, with ecological and biogeographic characteristics

Abstract

Background

Two decades have passed since the publishing of the last checklist of the millipedes of Slovakia. During this time, several new faunistic records have been added and taxonomic revisions have occurred. The present updated checklist summarises data on all millipede species recorded in Slovakia, including altogether 93 species.

New information

For each species, general habitat characteristics, ecological classification and distributional pattern are provided. Ecological classification is presented for the first time for the millipede species occurring in Slovakia and is proposed as a tool for ecological studies and for the nature protection purposes. Special remarks are given to the species newly found for Slovakia, Geoglomerissubterranea Verhoeff, 1908, Brachyiuluslusitanus Verhoeff, 1898, Cylindroiulusbritannicus (Verhoeff, 1891), C.parisiorum (Brölemann & Verhoeff, 1896) and Polydesmusburzenlandicus Verhoeff, 1925, as well as to C.arborum Verhoeff, 1928, the species newly confirmed for Slovakia after more than 70 years.

Introduction

Until 1993, Slovakia was part of several political units (Hungarian, Austro-Hungarian Monarchy, Czechoslovakia, Slovak Republic, Hungary); therefore, historical data on Slovak millipede fauna are not easy to find and there has been a lot of geographical confusion in trying to locate old published records. The studies on the millipedes in Slovakia have a long history, with the first papers published at the end of the 19^th century (Nowicki 1867, Nowicki 1869, Nowicki 1870, Nowicki 1871, Karliński 1883a, Karliński 1883b, Latzel 1882, Latzel 1884, Chyzer 1886, Daday 1889, Daday 1896, Petricskó 1891, Petricskó 1892, Malesevics 1892, Attems 1895, Attems 1899, Verhoeff 1899a, Verhoeff 1899b). One of the most important authors working at that time in Slovakia was Ödön Tömösváry. He wanted to prepare a monograph on the myriapods of the Kingdom of Hungary, but died young and his findings were at least excerpted in the papers of Chyzer (1886), Daday (1889) and Daday (1896).

In the pre-war and interwar periods, K. W. Verhoeff, the phenomenal German zoologist, expanded the knowledge of the millipedes of our country. In addition, important information about the millipede fauna of Slovakia is included in historical monographs published by Attems (1926), Verhoeff (1928), Verhoeff (1932), Verhoeff (1937), Ortvay (1902) and Schubart (1934). A significant contribution to the knowledge of Slovak millipedes during the first half of the 20^th century was done by other authors, including Jawlowski (Jawlowski 1930, Jawlowski 1938), Jermy (1942) and Loksa (Loksa 1954, Loksa 1957, Loksa 1962, Loksa 1968). In the second half of the 20^th century, the most extensive research on millipedes in Slovakia was performed by Ján Gulička (1925–2009) (Gulička 1952, Gulička 1954, Gulička 1955, Gulička 1956a, Gulička 1956b, Gulička 1957, Gulička 1960b, Gulička 1965, Gulička 1985, Gulička 1986). Gulička's unpublished compendia (Gulička 1951, Gulička 1960a), which we have today and can work with, were not taken into account in the last published checklist (Mock 2001a). This last checklist included all the available information at that time and involved information for about 70 species reported for the whole territory of Slovakia. A new chapter of millipede research began at the end of the 20^th century, conducted by several researchers, for example, Tajovský (1997), Mock (Mock 1998, Mock 1999a, Mock 1999b) and Stašiov (Stašiov 1997, Stašiov 1998). Several studies on Slovak millipede fauna were published after Gulička’s death (Gulička 2016, Gulička et al. 2014, Gulička and Košel 2016), based on material and manuscripts from his estate. Besides this, Gulička's unpublished compendia (Gulička 1951, Gulička 1960a), which we have today available and data listed there can be actually re-evaluated and taken into consideration, represent other sources for more complex overview of the millipede fauna of Slovakia.

Since 2001, the number of species has changed significantly, thus warranting a new update. The aim of this report is to update the list of millipede species and their distribution in Slovakia, to summarise published, as well as unpublished data and records, respecting recent taxonomic changes and to provide a useful tool in the form of ecological and geographical classification of species for other studies.

Materials and methods

The classification for Diplopoda used for this checklist follows the systematic arrangement by Shear (2011), while the nomenclature follows Kime and Enghoff (2011), Kime and Enghoff (2013), Kime and Enghoff (2017). We verified the system and nomenclature using MilliBase (Sierwald and Spelda 2021); however, we are reticent towards some proposals, for example, dividing the genera Brachydesmus and Polydesmus into other genera.

We do not include the subspecific categories as subspecies, nor all the relevant literature since that information has been included by the author of the last published checklist (Mock 2001a). References are listed at the species where the first records for Slovakia were documented for the first time after the year 2000. In addition, habitat evaluation for each species is included, applied specifically to the area of Slovakia, following adapted categorisation proposed by Tuf and Tufová (2008) and marked by a letter in caps:

• Relic species (R) - stenotopic species inhabiting exclusively undisturbed habitats with low impact of human activities, for example, the nature-closest forests, remnants of steppes, caves, stony debris, mountain habitats.

• Adaptable species (A) - species inhabiting nature-close habitats (various types of forests or meadows), able to penetrate artificial and man-made habitats (parks, abandoned gardens, graveyards, greenhouses etc.).

• Eurytopic species (E) - species with the widest ecological valence, frequently found in forested and non-forested biotopes and in man-made plots (fields, brownfields etc.).

• Synanthropic species (S) - species inhabiting a wide spectrum of man-made habitats, missing in natural biotopes, species significantly synanthropic, although occasionally found in natural, mainly altered, sites.

• Exotic species (X) – this category includes mostly tropical species, non-native or introduced species, found exclusively in hothouses.

Ecological classification of each species (Kime and Enghoff 2011, Kime and Enghoff 2017) is provided in Notes, marked by lowercase letters:

• (c) - corticolous

• (e) - epigeic

• (ed) - edaphic

• (h) - hygrophilous

• (m) - mountainous

• (n) - nidicolous

• (tb) - troglobitic

• (tp) - troglophilous

• (x) - xerophilous

We would like to note that we did not adopt this habitat evaluation scheme mechanically, but we evaluate each species, based on the knowledge about its ecology in Slovakia. Question mark (?) next to the classification refers to problematic data on the occurrence of the species.

In addition, a general geographical category that outlines the distribution of each species is included in the list (Kime and Enghoff 2011, Kime and Enghoff 2017).

Data resources

Records from Slovakia compiled from the published literature and additional collected material are summarised. The actually sampled material is in the collection at the workplace of the authors' team. Part of the data was obtained by preliminary revision of the collection by J. Gulička, including his unpublished compendia. A large portion of Gulička's complete written legacy is lost. However, his views on the taxonomy, biogeography and ecology of individual species can be found in Gulička's additionally handwritten notes in the monographs of other authors (e.g. Verhoeff 1928, Schubart 1934 etc.).

Checklists

Updated list of the millipede fauna of Slovakia

Polyxenida

51B5879F-5FC7-5A53-8809-BE8BA4E6EFD4

Polyxenidae

1474AEEA-6B12-5591-ACB8-91EDF280965C

Polyxenus lagurus

(Linnaeus, 1758)

828426A7-1F69-533A-8D81-0CB37BEC7213

Distribution

West Palearctic and Nearctic

Notes

A, c, n

Glomerida

BFC7F491-9FAA-5D78-ADF4-E7B530A4DD4E

Notes

The order Glomerida, according to published data, is represented by three genera in Slovakia – Trachysphaera, Glomeris and Geoglomeris (Lang 1954, Kime and Enghoff 2011, Kime and Enghoff 2013, Kocourek et al. 2017). Genus Glomeris is rather problematic, as it represents a complex of species, characterised by inconsistency and unreliable determination information in the old literature. G.tetrasticha, G.hexasticha and G.pustulata represent well-confirmed species, occurring in the territory of Slovakia. Only one old record of G.klugii from Slovakia exists. In the area of Slovakia, some populations of the species of the genus Glomeris are characterised by a remarkable variety of colouration, which could indicate the ongoing speciation processes.

Glomeridae

EBD87C29-C98C-5A2D-8A64-999815A34F72

Glomeris hexasticha

Brandt, 1833

A2511E1B-A91D-5850-ABB7-F2C07503C54C

Distribution

South, Central and East European

Notes

A, e, x

Glomeris klugii

Brandt, 1833

61A6C9BA-E61A-533A-8F6C-4DA9CA387815

Distribution

West and South European

Notes

A, e, ?

The species (known under synonyms G.conspersa and G.undulata in older literature) is characterised by a south-western occurrence in Europe (Kime and Enghoff 2011). It has been confirmed in most of the neighbouring countries (Poland, Czech Republic, Austria and Hungary). For Slovakia, only one record of this species has been recently discovered, in the millipede collection of Czech arachnologist František Miller (1902–1983), housed in the National Museum, Prague (Dolejš and Kocourek 2020). One female specimen of G.klugii is included in the collection, labelled with April 1930, Turčianske Teplice District, Žilina Region. Based on this information, a series of sampling from this area has been conducted, but unsuccessfully. Therefore, a mistake in localisation made by collector is highly possible.

Reference: Dolejš and Kocourek (2020)

Glomeris mnischechi

Nowicki, 1870

694103EA-A3AF-533E-A057-1E6D5868BA6A

Distribution

West Carpathian

Notes

R, e

Glomerismnischechi is considered the only endemic species of the order Glomerida north of the Alps. According to Kime and Enghoff (2011), the species inhabits mountainous biotopes of Slovakia (Belianske Tatras Mts., Pieniny Mts.) and Poland. However, literature contains many controversial data about diagnostic characteristics, as well as ecological demands (Mock 2001a, Kravcová and Mock 2014). Nowicki originally offered two descriptions of G.mnischechi (Nowicki 1870, Nowicki 1871), without any illustration. In addition, the author used different transcription for the species names in both cases, Glomerismnischechi (Nowicki 1870) and Glomerismniszechii (Nowicki 1871). After this confusion, the species was described several times in the old literature, under various synonyms (Latzel 1884, Gulička 1951, Gulička 1960a, Dziadosz 1966). Other authors considered the species to be subspecies of G.hexasticha, due to the striking similarity of both species (e.g. Jawlowski 1938). Although the taxonomic status of the species remains unresolved, recent molecular analyses of several representatives of G.hexasticha from the type locality of G.mnischechi suggest the presence of several separate species; therefore, the existence of G.mnischechi and its occurrence in Slovakia cannot be ruled out.

Glomeris pustulata

(Fabricius, 1781)

8EAAF47F-F7BE-5D57-AD24-C8CA0B7A8FC5

Distribution

South and Central European

Notes

A, e, c

Glomeris tetrasticha

Brandt, 1833

D64F6E1F-0B9A-5880-9006-DBADB01E7E7C

Distribution

Central and East European

Notes

A, e

Some authors mentioned the occurrence of Glomerisconnexa C. L. Koch, 1847 in Slovakia. This species is characteristic for its south-western European distribution (Kime and Enghoff 2011). In the literature from the last century, there is a lot of data on the findings and its occurrence. However, it should be noted that, in most cases, under this name, there were referred other species, especially G.tetrasticha (Kime and Enghoff 2011, Kocourek et al. 2017). From Slovakia, no recent reliable records of its occurrence are available; therefore, the older data on this species name are preliminarily all included under G.tetrasticha. However, due to the findings from the Czech Republic, very close to the border with Slovakia, the occurrence of this species cannot be ruled out.

Geoglomeris subterranea

Verhoeff, 1908

485B724B-7148-51A6-9F8A-F595371C7499

Distribution

West and Central European

Notes

R, ed, tp, h

This species was described by Verhoeff (1908) from the neighbourhood of Dresden, Germany, where two females were found on limestone near a brook. In his monograph on the millipede fauna of Czechoslovakia, Lang (1954) presented the finding of G.subterranea in the only Slovak locality from the vicinity of the Bratislava City (Malé Karpaty Mts.). However, he did not comment on the finding and attached only a picture taken from other literature (Schubart 1934). Since the cited monograph contains many ambiguities and unreliable data, this information is considered doubtful (Gulička 1986, Mock 2001a). In the area of Slovakia, it was found only recently (Haľková et al., unpublished), repeatedly, in karst springs and wetlands on karst bedrock. Detailed morphological study (including SEM), supported by molecular analysis, confirmed the identity of G.subterranea, without any apparent morphological adaptations to aquatic and semi-aquatic habitat. The possibility that this species is not strictly limited to the terrestrial environment has already been suggested by Noll (1939), although his findings were completely forgotten in recent literature. Noll mentioned the presence of Geoglomerisjurassica (a younger synonym of G.subterranea) in the water of three wells in Northern Bavaria, Germany. The author explained its occurrence as random, presuming the animals entered the well through crevices in the wall.

Trachysphaera acutula

(Latzel, 1884)

E2F129A8-8841-5B7A-8884-F673C64DA7E8

Distribution

Carpathian

Notes

R, ed, tp, h

Trachysphaera costata

(Waga, 1857)

B05A6FD8-BBE1-5C39-87F4-27EE0DDCB1E5

Distribution

East and Central European

Notes

R, ed, m

Trachysphaera gibbula

(Latzel, 1884)

C8D3F0D0-4F6A-570C-A64D-2544106EAECD

Distribution

Alpine and West Carpathian

Notes

R, ed

Polyzoniida

0D81710B-775D-5A5F-B443-9D6023099906

Polyzoniidae

621915C8-EAFC-5396-9714-FE20D7462F70

Polyzonium eburneum

Verhoeff, 1907

F24C4010-6F24-56CD-9527-A5CFB0D89E06

Distribution

East Alpine and West Carpathian

Notes

R, e

Polyzonium germanicum

Brandt, 1837

0FF2874E-FB0B-579D-AE75-912804D3A386

Distribution

European

Notes

A, e, h

Polyzonium transsilvanicum

Verhoeff, 1898

6E9952B6-3AB8-57ED-927F-53563C7C8C6C

Distribution

East Carpathian

Notes

R, h

Julida

85AF59DC-241D-5259-BB9E-A2B5FD794658

Blaniulidae

EFAA8300-886A-5590-B1A3-EAECD325EF46

Archiboreoiulus pallidus

(Brade-Birks, 1920)

3481208A-BB4B-5F5A-8FB5-5089E9D8B446

Distribution

West, North, Central and East European

Notes

R, ed, tp

Reference: Mock et al. (2015)

Blaniulus guttulatus

(Fabricius, 1798)

B89DF54C-253C-5767-887F-BF4C2CF3B1A6

Distribution

European

Notes

S, ed

Cibiniulus slovacus

Antić, Mock & Enghoff, 2015

6BD50117-1D65-52DF-B16F-788C6FE3D833

Distribution

West Carpathian

Notes

R, ed, tp

Choneiulus palmatus

(Němec, 1895)

58F43795-2A1E-5F3A-917B-9BBF992DCF83

Distribution

European

Notes

S, ed

Reference: Mock (2001b)

Nopoiulus kochii

(Gervais, 1847)

82D904AB-2C93-5895-9895-696840D9AD3B

Distribution

Euro-Caucasian

Notes

S, ed

Proteroiulus fuscus

(Am Stein, 1857)

90CD54F7-E609-5EEA-A47D-CA7C9BFB183F

Distribution

European

Notes

E, c, n

Nemasomatidae

17D6B204-1121-5909-8A04-30BB23FCB962

Nemasoma varicorne

C. L. Koch, 1847

E2AE1783-D0ED-5389-99E3-EA3FDCEA4048

Distribution

European

Notes

R, c, n

Julidae

A0FD6BA1-3FBD-5217-908B-7229ADD0FA3B

Brachyiulus bagnalli

(Brolemann, 1924)

8A120E9F-8E37-5E28-A5AE-266719F586D5

Distribution

South and Central European

Notes

E, e, n, x

Brachyiulus lusitanus

Verhoeff, 1898

DA264670-66AC-5322-B965-09E7A9B62558

Materials

1. Type status:Other material. Occurrence: individualCount: 1; sex: male; lifeStage: adult; Taxon: scientificName: Brachyiuluslusitanus Verhoeff, 1898; class: Diplopoda; family: Julidae; genus: Brachyiulus; specificEpithet: lusitanus; taxonRank: species; Location: country: Slovakia; locality: Košice Basin, Košice – a city park (Mestský park) at the railway station; verbatimElevation: 210 m a.s.l.; locationRemarks: leaf litter (Acer platanoides, Aesculus hippocastaneum); verbatimCoordinates: 48°43'20.4"N 21°15'52.5"E; Identification: identifiedBy: A. Mock; Event: eventDate: 18-12-2018; fieldNotes: co-existing with Nopoiuluskochii, Cylindroiulusparisiorum, Ophyiuluspilosus; Record Level: basisOfRecord: PreservedSpecimen

2. Type status:Other material. Occurrence: individualCount: 2; sex: female; lifeStage: adult; Taxon: scientificName: Brachyiuluslusitanus Verhoeff, 1898; class: Diplopoda; family: Julidae; genus: Brachyiulus; specificEpithet: lusitanus; taxonRank: species; Location: county: Slovakia; locality: Košice Basin, Košice – a city park (Mestský park) at the railway station; verbatimElevation: 210 m a.s.l.; locationRemarks: leaf litter (Acer platanoides, Aesculus hippocastaneum); verbatimCoordinates: 48°43'20.4"N 21°15'52.5"E; Identification: identifiedBy: A. Mock; Event: eventDate: 18-12-2018; fieldNotes: co-existing with Nopoiuluskochii, Cylindroiulusparisiorum, Ophyiuluspilosus; Record Level: basisOfRecord: PreservedSpecimen

Distribution

European

Notes

A, e

According to Kime and Enghoff (2017), the species is distributed in Western and Central Europe and Balkans, but also Algeria, Egypt and Iran. It was introduced into Australia and North America. The species can be found in forests, in addition to open land, meadows, cornfields and vineyards. It has been recorded under stone debris, as well as city parks and heated greenhouses.

Cylindroiulus boleti

(C. L.Koch, 1847)

4B2E539A-C953-5C56-A2A3-B021375BA5F8

Distribution

South and Central European

Notes

R, e

Cylindroiulus arborum

Verhoeff 1928

B89DFBDB-1AE3-5FF3-9F8F-FC29FCCBABC9

Materials

1. Type status:Other material. Occurrence: individualCount: 1; sex: male; lifeStage: adult; Taxon: scientificName: Cylindroiulusarborum Verhoeff, 1928; class: Diplopoda; family: Julidae; genus: Cylindroiulus; specificEpithet: arborum; taxonRank: species; Location: country: Slovakia; locality: Burda Mts.; verbatimElevation: 190 m a.s.l.; locationRemarks: forest (Quercus spp., Carpinusbetulus, Fraxinusexcelsior), decomposed wood from an oak-tree cavity; verbatimCoordinates: 47°50'39.84"N 18°49'16.92"E; Identification: identifiedBy: A. Mock; Event: eventDate: 2017-06-16; fieldNotes: co-existing with Proteroiulusfuscus and juveniles of Haaseaflavescens; Record Level: basisOfRecord: PreservedSpecimen

2. Type status:Other material. Occurrence: individualCount: 1; sex: female; lifeStage: adult; Taxon: scientificName: Cylindroiulusarborum Verhoeff, 1928; class: Diplopoda; family: Julidae; genus: Cylindroiulus; specificEpithet: arborum; taxonRank: species; Location: country: Slovakia; locality: Burda Mts.; verbatimElevation: 190 m a.s.l.; locationRemarks: forest (Quercus spp., Carpinusbetulus, Fraxinusexcelsior), decomposed wood from an oak-tree cavity; verbatimCoordinates: 47°50'39.84"N 18°49'16.92"E; Identification: identifiedBy: A. Mock; Event: eventDate: 2017-06-16; fieldNotes: co-existing with Proteroiulusfuscus and juveniles of Haaseaflavescens; Record Level: basisOfRecord: PreservedSpecimen

3. Type status:Other material. Occurrence: individualCount: 16; lifeStage: juveniles; Taxon: scientificName: Cylindroiulusarborum Verhoeff, 1928; class: Diplopoda; family: Julidae; genus: Cylindroiulus; specificEpithet: arborum; taxonRank: species; Location: country: Slovakia; locality: Burda Mts.; verbatimElevation: 190 m a.s.l.; locationRemarks: forest (Quercus spp., Carpinusbetulus, Fraxinusexcelsior), decomposed wood from an oak-tree cavity; verbatimCoordinates: 47°50'39.84"N 18°49'16.92"E; Identification: identifiedBy: A. Mock; Event: eventDate: 2017-06-16; fieldNotes: co-existing with Proteroiulusfuscus and juveniles of Haaseaflavescens; Record Level: basisOfRecord: PreservedSpecimen

Distribution

Central and East European

Notes

R, e

Mainly lowland species with Central and East European distribution (Kime and Enghoff 2017). The species usually prefers forest habitats, found mostly in deadwood and leaf litter, although several records are from hothouses and other artificial habitats. New record after more than 70 years in Slovakia (see Dudich 1958, Mock 2001a).

Cylindroiulus britannicus

(Verhoeff, 1891)

00B4D62F-CAB4-5AC1-A501-206AD0322D4C

Materials

1. Type status:Other material. Occurrence: individualCount: 5; sex: male; lifeStage: adult; Taxon: scientificName: Cylindroiulusbritannicus (Verhoeff, 1891); class: Diplopoda; family: Julidae; genus: Cylindroiulus; specificEpithet: britannicus; taxonRank: species; Location: country: Slovakia; locality: Košice Basin, Košice, Public cemetery; verbatimElevation: 225 m a.s.l.; locationRemarks: leaf litter (Tilia cordata, Fraxinusexcelsior, Acer platanoides); verbatimCoordinates: 48°41'42.420"N 21° 15'35.130"E; Identification: identifiedBy: A. Mock; Event: eventDate: 2020-05-26; fieldNotes: co-existing with Blaniulusguttulatus, Choneiuluspalmatus, Cylindroiulus caeruleoncictus, Ophyiuluspilosus; Record Level: basisOfRecord: PreservedSpecimen

Distribution

Cosmopolitan

Notes

S, c, e

Species with European distribution, in Britain and Ireland found beneath the bark of dead deciduous tree trunks and stumps (Blower 1985). It appears to be predominantly or entirely synanthropic in most other countries, occurring primarily in cities and cultivated areas (Kime and Enghoff 2017). The same applies to the first findings of the species in Slovakia, where it was found in city parks.

Cylindroiulus burzenlandicus

Verhoeff, 1907

87350C1D-256A-5255-80AB-B78FB5330452

Distribution

Carpathian

Notes

R, e, m

Cylindroiulus caeruleocinctus

(Wood, 1864)

59F9CDA8-596B-5293-B1F4-B134097D301F

Distribution

Cosmopolitan

Notes

S, e

Reference: Mock (2006)

Cylindroiulus latestriatus

(Curtis, 1845)

015DCC44-B303-577D-8F68-8D8FD23D8EBD

Distribution

Atlantic, North-western and Central European

Notes

S, e

Reference: Mock (2001b)

Cylindroiulus parisiorum

(Brölemann & Verhoeff, 1896)

2169FE95-512D-5B5C-9687-EE993EA34BAF

Materials

1. Type status:Other material. Occurrence: individualCount: 21; sex: male; lifeStage: adult; Taxon: scientificName: Cylindroiulusparisiorum (Brölemann & Verhoeff, 1896); class: Diplopoda; family: Julidae; genus: Cylindroiulus; specificEpithet: parisiorum; taxonRank: species; Location: country: Slovakia; locality: Košice Basin, Košice – a city park (Mestský park) at the railway station; verbatimElevation: 210 m a.s.l.; locationRemarks: leaf litter (Acer platanoides, Aesculus hippocastaneum); verbatimCoordinates: 48°43'20.4"N 21°15'52.5"E; Identification: identifiedBy: A. Mock; Event: eventDate: 2018-12-18; fieldNotes: co-existing with Nopoiuluskochii, Brachiulus lusitanus, Ophyiuluspilosus; Record Level: basisOfRecord: PreservedSpecimen

2. Type status:Other material. Occurrence: individualCount: 28; sex: female; lifeStage: adult; Taxon: scientificName: Cylindroiulusparisiorum (Brölemann & Verhoeff, 1896); class: Diplopoda; family: Julidae; genus: Cylindroiulus; specificEpithet: parisiorum; taxonRank: species; Location: country: Slovakia; locality: Košice Basin, Košice – a city park (Mestský park) at the railway station; verbatimElevation: 210 m a.s.l.; locationRemarks: leaf litter (Acer platanoides, Aesculus hippocastaneum); verbatimCoordinates: 48°43'20.4"N 21°15'52.5"E; Identification: identifiedBy: A. Mock; Event: eventDate: 2018-12-18; fieldNotes: co-existing with Nopoiuluskochii, Brachiulus lusitanus, Ophyiuluspilosus; Record Level: basisOfRecord: PreservedSpecimen

3. Type status:Other material. Occurrence: individualCount: 28; sex: female; lifeStage: adult; Taxon: scientificName: Cylindroiulusparisiorum (Brölemann & Verhoeff, 1896); class: Diplopoda; family: Julidae; genus: Cylindroiulus; specificEpithet: parisiorum; taxonRank: species; Location: country: Slovakia; locality: Košice Basin, Košice – a city park (Mestský park) at the railway station; verbatimElevation: 210 m a.s.l.; locationRemarks: leaf litter (Acer platanoides, Aesculus hippocastaneum); verbatimCoordinates: 48°43'20.4"N 21°15'52.5"E; Identification: identifiedBy: A. Mock; Event: eventDate: 2018-12-18; fieldNotes: co-existing with Nopoiuluskochii, Brachiulus lusitanus, Ophyiuluspilosus; Record Level: basisOfRecord: PreservedSpecimen

Distribution

European

Notes

S, e, c

The species has European distribution; however, the captures apart from England are scattered and isolated. Its occurrence is associated with human activity. According to Kime and Enghoff (2017), it is relatively rare, some records are likely to be incorrect due to possible confusion with other Cylindroiulus species. Findings in synanthropic habitat represent the first records for Slovakia.

Cylindroiulus vulnerarius

(Berlese, 1888)

3F2C90F6-E45B-5522-BEE0-7729C555013F

Distribution

West and Central European

Notes

X, e, h

Reference: Mock (2001b)

Enantiulus nanus

(Latzel, 1884)

6A9BDF51-A050-5A30-8414-E0B0559A7BD7

Distribution

European

Notes

A, e, h

Enantiulus tatranus

(Verhoeff, 1907)

A4829F1E-DA65-527C-B7B1-D37F201BA8C6

Distribution

West Carpathian

Notes

R, e, m

Enantiulus transsilvanicus

(Verhoeff, 1899)

FC523E99-BEB5-5322-BB58-4FFED80B2107

Distribution

East Carpathian

Notes

R, e

Reference: Gulička et al. (2014)

Julus curvicornis

Verhoeff, 1899

C4854698-B0A2-5FEA-88FC-10B1D892E657

Distribution

Carpathian

Notes

R, e

Julus scandinavius

Latzel, 1884

838E1EA4-4893-559E-AC0B-12AF67C88986

Distribution

Central European and Scandinavian

Notes

A, e, n

Julus scanicus

Lohmander, 1925

ACE7AF66-74D7-5242-A32D-D26CEF6D988E

Distribution

Central European

Notes

A, e, h

Julus terrestris

Linnaeus, 1758

991BA89E-2910-5CC7-A987-3ADC15B2DB20

Distribution

North, Central, South and East European

Notes

R, e, h

Kryphioiulus occultus

(C. L. Koch, 1847)

F1365525-3B19-500B-BE44-03B9C771BE26

Distribution

East European

Notes

A, e, x

Leptoiulus baconyensis

(Verhoeff, 1899)

15D295F6-7BF1-5386-A52E-66E7A0C596D2

Distribution

East and Central European

Notes

R, e

Leptoiulus cibdellus

(Chamberlin, 1921)

08E79850-A0D8-5202-90FB-1DCFEE256CFF

Distribution

North and Central European, Baltic

Notes

R, e, h, n

Leptoiulus liptauensis

(Verhoeff, 1899)

37631AAA-BF6F-5888-9930-B3D690E6A46D

Distribution

West Carpathian

Notes

R, e, m

Leptoiulus noricus

Verhoeff, 1913

8C2CE56F-DB40-5F25-A980-976BB5F8386E

Distribution

Sudetico-Carpathian

Notes

R, e, m

Leptoiulus mariae

Gulička, 1952

528708CF-BDA4-5421-8648-F5613BD4F96A

Distribution

West Carpathian

Notes

R, e, tp

Leptoiulus proximus

(Němec, 1896)

5D4CD2CC-5E74-584E-B453-8BD294C5FA8D

Distribution

North, Central and East European

Notes

A, e

Leptoiulus tatricus

Gulička, 1956

5CFD59DC-D3C9-51A1-A7EC-982F005C6A4B

Distribution

West Carpathian

Notes

R, e, m

Leptoiulus trilobatus

(Verhoeff, 1894)

E70CBA85-A434-5CCF-8054-9F30CB9DE647

Distribution

Central and East European

Notes

A, e

Leptoiulus tussilaginis

(Verhoeff, 1907)

10EE350B-3B20-5539-A3F8-F06E5320DA26

Distribution

West Carpathian

Notes

R, e, m

Megaphyllum projectum

Verhoeff, 1894

2F4ADCC2-0CAE-5197-96DE-987A6C1EE653

Distribution

Central and East European

Notes

A, e

Megaphyllum silvaticum

(Verhoeff, 1898)

18575707-9CD0-58B8-AC98-F6C577D56B10

Distribution

East and Central European

Notes

R, e, m

Megaphyllum unilineatum

(C. L. Koch, 1838)

82675623-7D8A-587F-9DA2-ECC464C45BD1

Distribution

South and Central European, Balkan

Notes

E, e, x

Ommatoiulus sabulosus

(Linnaeus, 1758)

F3581CAB-3DDD-5006-9F26-42C2AB926E38

Distribution

European

Notes

E, e, x

Ophyiulus pilosus

(Newport, 1842)

002768BF-03DC-5C20-AF68-E1D1D6813B89

Distribution

European

Notes

E, e

Unciger foetidus

(C. L. Koch, 1838)

968A4371-833F-5CF1-8DEB-6403DC30EB83

Distribution

European

Notes

E, e, h

Unciger transsilvanicus

(Verhoeff, 1899)

590A2C07-04D4-5B08-82B9-6E7AB7E2C297

Distribution

Central European

Notes

A, e

Xestoiulus carpathicus

(Verhoeff, 1907)

2AC4FCC1-001A-5F77-88D6-0BF22F3560D6

Distribution

North Carpathian

Notes

R, e, m

Xestoiulus laeticollis

(Porat, 1889)

9DECA411-8B99-5EBF-BFCC-9236A24D3D0B

Distribution

East and Central European

Notes

R, e, h, n

Reference: Tajovský et al. (2001)

Chordeumatida

0E3D0A19-D96F-53D6-B9B2-04BDD47C8879

Chordeumatidae

F11EB5B5-46CA-5729-AA43-7CDCC80AFBAA

Melogona broelemanni

(Verhoeff, 1897)

8779CE31-7CCF-5ED2-844B-F748C93AF4F1

Distribution

Central European, Balkan

Notes

S, e

Reference: Mock and Tajovský (2002)

Melogona transsylvanica

(Verhoeff, 1897)

160BA32D-8C74-58FD-A7F6-B98739996D13

Distribution

East Carpathian

Notes

A, e

Reference: Mock and Tajovský (2002)

Melogona voigtii

(Verhoeff, 1899)

A36ACD63-2589-50F5-BCEE-180E889D5584

Distribution

North and Central European

Notes

S, e

Reference: Mock and Tajovský (2002)

Brachychaeteumatidae

331205F4-1713-5FBA-8F29-22CB684628EA

Brachychaeteuma bradeae

(Brolemann & Brade-Birks, 1917)

CD723F79-E5EA-5909-A6A2-ABF03F0F0BAC

Distribution

West European

Notes

R, tp

Reference: Kováč et al. (2005)

Trachygonidae

BCF46DAF-6878-5C36-8C31-E0AFFA337A96

Heteroacrochordum evae

Loksa, 1960

FF9DAFFD-2F98-5440-B74F-938C13BBFF45

Distribution

West Carpathian

Notes

R, ed

Reference: Mock et al. (2019)

Hungarosomatidae

E6E1CFB1-CB6A-50D7-BF31-8C5C8BFE1358

Hungarosoma bokori

Verhoeff, 1928

10E8DB47-2C13-54BF-97D3-62954C7C3AB6

Distribution

West Carpathian

Notes

R, e, h

The first description of the species was published by Verhoeff (1928), based on a single female specimen from the Abaliget Cave in Hungary. Detailed analysis of diagnostic characteristics, based on the fresh material from the type locality, as well as all available museum material, was presented only recently by Mock et al. (2016). However, the authors overlooked the apparent similarity of the diagnostic features with that of Ochogonacervina (Verhoeff, 1899), recently pointed out by Antić et al. (2018). Gonopods of both species appear to be identical, nevertheless, a synonymy was not formally established. Minute differences between Verhoeff’s description of O.cervina and our knowledge of H.bokori must be reviewed. In addition, in order to justify the name of the family Hungarosomatidae and its position in the Chordeumatida system, the decision as to which genus the species belongs: Ochogona, Octeicosisoma, Triakontizona or Ceratosoma, has to be resolved (Haľková and Mock 2018, Korsós and Lazányi 2020).

Reference: Papáč et al. (2014)

Craspedosomatidae

0A4EF9DD-BE59-578D-AE90-9AF360667BBF

Beskidia jankowskii

(Jawlowski, 1938)

5465705D-4E49-593D-8E26-4B084410442C

Distribution

East Carpathian

Notes

R, e, m

Reference: Gulička et al. (2014)

Craspedosoma raulinsii

Leach, 1814

18001744-0F61-565E-9304-C5DC381DEE11

Distribution

West, Central and South European

Notes

A, e

Craspedosoma transsilvanicum

Verhoeff, 1897

C034B994-8521-5B2A-BF62-86B943A61F9C

Distribution

South and Eastern European

Notes

R, e

Chelogona carpathicum

(Latzel, 1882)

9791AA65-998E-5E80-8ACA-379C1C6853A6

Distribution

West Carpathian

Notes

R, e, m

Ochogona caroli

(Rothenbuhler, 1900)

F85EE3C3-6D8D-598A-800C-A0E921089325

Distribution

Central European

Notes

R, e

Reference: Gulička et al. (2014)

Haasidae

8465A86A-22AF-58EA-A29D-62CCEF45F63C

Haasea flavescens

(Latzel, 1884)

C84E98DF-B346-5CFA-997A-1DBA43A3F8F5

Distribution

South and Central European

Notes

A, e

Hylebainosoma gulickai

(Tajovský, Mock & Papáč, 2014)

A999896F-BD64-58F6-A12B-B3C7541A1B9F

Distribution

West Carpathian

Notes

R, tb

Hylebainosoma tatranum

Verhoeff, 1899

140D933F-8B82-55A7-98C8-2C3B87E733AD

Distribution

North Carpathian

Notes

R, e, m

Entomobielziidae

E7D5C923-C05B-5F7F-9308-8679655690C8

Entomobielzia kimakowizii

(Verhoeff, 1897)

A238419E-E84D-5864-94E8-30FCFEB8B7DA

Distribution

East Carpathian

Notes

R, e

Reference: Gulička et al. (2014)

Attemsiidae

65C7370E-1D2E-5072-9B36-54FF3C108D9C

Allorhiscosoma sphinx

(Verhoeff, 1907)

2A92FCE4-9022-5340-833A-B55860611281

Distribution

West Carpathian

Notes

R, tp

Mecogonopodium carpathicum

Mock & Tajovský, 2008

8E374C6F-3304-51BB-B71F-3023B348134D

Distribution

West Carpathian

Notes

R, tp

Mastigophorophyllidae

F3341C42-CE6E-5A79-97D3-5A0080CB7F45

Haploporatia eremita

(Verhoeff, 1909)

9CADC961-AE1B-5FD5-9F57-4B13B5EA3CCF

Distribution

Central European

Notes

R, e

Mastigona bosniensis

(Verhoeff, 1897)

10F158A8-343B-5272-98A4-A5C356723B03

Distribution

Central and East European

Notes

E, e

Mastigophorophyllon cirriferum

Verhoeff, 1899

304B8977-AF62-58F9-BDF5-7CCD99B24798

Distribution

West Carpathian

Notes

R, e, m

Verhoeffiidae

725C200A-8CB2-5B3F-899B-73FEFDE65557

Haplogona oculodistincta

(Verhoeff, 1893)

3B6E25C9-4384-59EA-8A19-7E147787968D

Distribution

Alpine

Notes

S, e

Reference: Haľková and Mock (2018)

Polydesmida

04B3F154-EEB0-589A-AAB6-EA53CFF83B6F

Polydesmidae

CE25DD76-98C6-5791-BFB1-E37E5C39CA55

Brachydesmus dadayi

Verhoeff, 1895

533BBE4D-9FCB-563B-B251-E8258224F4C2

Distribution

South, Central and East European

Notes

A, e, h

Brachydesmus superus

Latzel, 1884

694DE3E8-C9E8-52CC-A576-03D8521719E2

Distribution

European

Notes

S, e, h

Polydesmus burzenlandicus

Verhoeff, 1925

3F6D92C3-5485-51AF-A747-5E482E314050

Distribution

South-eastern Carpathian

Notes

R, e, c

Polydesmusburzenlandicus is a small hygrophilous representative of the order Polydesmida, widespread mainly in the south-eastern Carpathians (Kime and Enghoff 2011) The species inhabits forest habitats of the mountainous landscape. It was recently documented from the area of the Latorica PLA (Mock et al. 2021). These findings represent the first record for Slovakia.

Reference: Mock et al. (2021)

Polydesmus complanatus

(Linnaeus, 1761)

2F776C36-2364-557E-9339-FA5E7F1A6C08

Distribution

North, Central, South and East European

Notes

E, e, h

Polydesmus denticulatus

C. L. Koch, 1847

DFBCDACE-498E-578F-9B2B-6666B24578C4

Distribution

European

Notes

E, e

Polydesmus inconstans

Latzel, 1884

41C36D84-A0C7-5A99-A3F6-40C3FF568B9B

Distribution

European

Notes

S, e, h

Reference: Mock (2004)

Polydesmus komareki

Ložek & Gulička, 1962

7A252CA8-152E-5F89-9304-7D099E79DD1B

Distribution

East Carpathian

Notes

R, e, m

Polydesmus montanus

Daday, 1889

AED868E6-77C3-5328-A02B-3A599AD52E30

Distribution

East Carpathian

Notes

R, e, c, m

Polydesmus polonicus

Latzel, 1884

7602842C-F4AB-568C-A312-4A9572FEE2B7

Distribution

East Carpathian

Notes

R, e, m

Polydesmus subscabratus

Latzel, 1884

8387BDA8-3A08-5193-B8A5-708A271BC0AB

Distribution

South Carpathian

Notes

R, e, tp, ?

South Carpathian endemic species with affinity to wet forested habitats. From Slovakia, only one questionable historical record is documented from the vicinity of the Veľaty Village (Daday 1889), significantly isolated from the rest of the area of its known distribution in the Southern Carpathians (Kime and Enghoff 2011, Kime and Enghoff 2013). Its occurrence in Slovakia, however, can be supported by the recent findings of P.transylvanicus Daday, 1889 in nearby locations within the Eastern Slovak Plain (Haľková and Mock 2018), the species with similar biogeography and ecology as P.subscabratus.

Polydesmus tatranus

Latzel, 1884

6135C067-D8A4-5AEF-B37B-BDACE0170229

Distribution

Carpathian

Notes

R, e, h, m

Polydesmus transylvanicus

Daday, 1889

F19BF5DF-6F23-5EBD-BC58-0DDB68335FC0

Distribution

Carpathian

Notes

R, e, h

Reference: Haľková and Mock (2018)

Paradoxosomatidae

CA7A8050-3949-55B0-8481-75FBA30951DF

Oxidus gracilis

(C. L. Koch, 1847)

00AF0722-FD46-50DD-A656-DB0CEE6CB79E

Distribution

Cosmopolitan

Notes

X, e

Strongylosoma stigmatosum

(Eichwald, 1830)

58E29EC2-4A4D-5FF3-8A8E-6095FD2EB639

Distribution

East and Central European, Baltic

Notes

A, e, tp, h

Pyrgodesmidae

39B0E373-F2B2-5294-A509-D0FC38803166

Poratia digitata

(Porat, 1889)

8C690F2D-6B47-56FC-B104-FF46490EB680

Distribution

North, West and Central European

Notes

X, e, c, h

Reference: Mock (2001b)

Oniscodesmidae

5A793D16-C55A-5C43-9D9B-5CC8B6D5FECA

Amphitomeus attemsi

(Schubart, 1934)

FC24AFCE-C982-58AC-A70A-8C2F9A09C482

Distribution

East and Central European

Notes

X, e, c, h

Reference: Mock (2001b)

Discussion

The increase in faunistic research over the past two decades has resulted in significant expansion of knowledge of the Slovak millipede fauna. Since the last published review of millipede species from the territory of Slovakia (Mock 2001a), 23 species have been added, resulting at present in 93 species. Within the whole Carpathian Region, Romania still represents the country with the richest fauna, with 170 recorded millipede species (Giurginca 2021). Nevertheless, compared to the total species number of millipedes from neighbouring countries, 94 from Poland (Stojałowska and Staręga 1974, Wytwer 1997), 77 from the Czech Republic (Tajovský and Tuf 2016, Kocourek et al. 2017), 107 species from Hungary (Korsós and Lazányi 2020), 190 from Austria (extrapolation, Geiser 2018) and 75 from Germany (Hauser and Voigtländer 2019), 93 species recorded for Slovakia constitute a strikingly high number, considering the rather small area of the country. In addition, new findings are still expected for the whole territory of Slovakia since numerous locations remain less investigated, as well as records of non-native species are likely to emerge.

Most of the unclear and questionable data that were preliminarily excluded from the list are from the older literature. One of the most recent studies, creating confusion due to imprecision in accurate and reliable determination, was published by Topp et al. (2006). The authors investigated the biodiversity of woodlice and millipede faunas of the primeval forest of Central Slovakia. The authors' team, however, lacked a taxonomist and, as a result, species not occurring in our country or species with strictly synanthropic occurrence were listed, such as such as Mycogonagermanica, Allaiulusnitidus or Blaniulusguttulatus and Cylindroiuluslatestriatus (as Aneuloboiulusfrisius). From the faunistic point of view, the article is very inaccurate and should be approached with great caution.

The millipede fauna of Slovakia can be characterised as a combination of European, Alpine-Atlantic and Carpathian elements, with the occasional influence of Caucasian and Balkan fauna (Kime and Enghoff 2011, Kime and Enghoff 2017). From the biogeographical aspect, a relatively high number of endemic species are represented in the country. This is presumably linked to the Carpathians. Compared to the Alps, the geographical repartition of the Carpathian fauna was less affected by glaciations, allowing the isolation of the Carpathian Regions surrounded by the Paratethys (Holdhaus 1954). The presence of a significant proportion of endemic fauna characterises the millipede fauna of Slovakia as exceptional in Central Europe. Gulička and Košel (2016) have mentioned the presence of 28 Carpathian endemic species of millipedes in Slovakia. A large part of these species is defined as Western-Carpathian. However, the border between the Eastern and Western Carpathians crosses the eastern part of the country, resulting in the representation of the eastern Carpathian fauna. Nevertheless, it should be noted that this border is not strict and is not consistent for all the species. For some species, the term Northern-Carpathian endemic is more appropriate. Some species even can be characterised by microendemism (e.g. Leptoiulustatricus, Mecogonopodiumcarpathicum and Hylebainosomagulickai). The presence of the Western Carpathians has an undeniable influence on the composition of Slovak millipedes. Almost 20% of the species are represented by rare mountainous fauna and more than 10% are characteristic for their preference of cave habitat.

According to ecological classification, 50% of Slovak millipede fauna are represented by stenotopic species inhabiting exclusively undisturbed habitats with a low impact of human activities. A total of 20% can be classified as adaptable and only 7% can be classified as eurytopic. While the original classification, proposed by Tuf and Tufová (2008), is applicable to most of the species, the categorisation does not sufficiently segregate non-native, introduced species occurring exclusively in a specific environment (e.g. tropical species in greenhouses). Therefore, we added and used two categories to this classification, in order to distinguish synanthropic and exotic species. These categories include the remaining 13% of millipede species of Slovakia.

Despite the increased interest in faunistic research, the millipede fauna of Slovakia has not yet been completely investigated. Several findings are still to be determined at the species level. Such findings include subadult and female individuals of Typhloiulus sp. from the Domica Cave (Papáč et al. 2014) and a representative of the family Trichopolydesmidae (Akkari and Enghoff 2011). Both taxa are connected to the cave environment.