Zoosyst. Evol. 100 (2) 2024, 493-513 | DOI 10.3897/zse.100.122288 > PENSUFT. yee BERLIN Natural history collections help resurrecting Glomeris herzogowinensis Verhoeff, 1898 and further clarify the nomenclature of two Onychoglomeris subspecies of Attems (Diplopoda, Glomerida, Glomeridae) Dragan Anti¢', Thomas Wesener?, Nesrine Akkari? 1 University of Belgrade — Faculty of Biology, Institute of Zoology, Center for Biospeleology, Studentski Trg 16, 11000 Belgrade, Serbia 2 Leibniz Institute for the Study of Biodiversity Change, Zoogisches Forschungsmuseum Alexander Koenig, Adenauerallee 127, 53113 Bonn, Germany 3) 37 Zoological Department, Natural History Museum Vienna, Burgring 7, 1010 Vienna, Austria https://zoobank. org/CF 9F DD8F-1432-4A0D-BA41-5D11B642110A Corresponding author: Dragan Antié (dragan.antic@bio.bg.ac.rs) Academic editor: Luiz Felipe Iniesta # Received 4 March 2024 Accepted | April 2024 Published 9 May 2024 Abstract Based on the study of freshly-collected material and old museum specimens, we have solved a decades-old riddle surrounding the name Onychoglomeris herzogowinensis (Verhoeff, 1898). The southern Dinaric coastal species Glomeris herzogowinensis Verhoeff, 1898 is revived, while Onychoglomeris herzogowinensis australis Attems, 1935 and O. h. media Attems, 1935, are treated here as full species after returning the specific name to Glomeris Latreille, 1902, O. australis Attems, 1935, stat. nov. and O. media Attems, 1935, stat. nov. Besides the designation of lectotypes, we provide comprehensive illustrations, diagnoses, detailed remarks and a distribution map for all three species. In addition, DNA barcoding provided COI sequences for Glomeris herzogowinensis and On- ychoglomeris australis stat. nov., along with the first barcoding data of one additional species of Onyvchoglomeris Verhoeff, 1906, O. ferraniensis Verhoeff, 1909 and two Glomeris species, the Balkan G. balcanica Verhoeff, 1906 and the trans-Adriatic G. pulchra Koch, 1847. The significance of historical specimens from natural history museums is briefly discussed. Key Words Balkan Peninsula, COI, Europe, Glomerinae, lectotype, syntypes, taxonomy Introduction The Western Palaearctic genus Glomeris Latreille, 1802 comprises about 75 species with a smaller number of taxa in the Canary Islands, North Africa and Anatolia and the majority of species on the European continent (Enghoff et al. 2015). Apart from the fact that this genus includes some of the most attractive and colourful millipedes in Europe, it is certainly a nightmare for taxonomists. Around 60 species of the genus live on the continent to- day, with an unfathomably large number of subspecies, varieties, forms or morphs having been described in the past, counted in hundreds (Golovatch et al. 2009). It is interesting to note that more than 80 forms have been described within the common central and southeast Eu- ropean species Glomeris hexasticha Brandt, 1833 alone (Kime and Enghoff 2011). Due to insufficient taxonomic information on the structure of the telopods and their uni- formity in this group, the species, subspecies or “lower categories” are mostly described on the basis of colour patterns. While in some species this pattern may be sta- ble, in many others, there is variability, even within the same population, where the colouration of one species may be similar or identical to the colour pattern of an- Copyright Antic, D. et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 494 Anti¢, D. et al.: Glomeris herzogowinensis and Onychoglomeris from the Balkans other species. Fortunately, the chaotic situation within the genus and the order Glomerida, in general, has improved somewhat in recent decades, largely due to an integrative approach to the problem (Hoess et al. 1997; Hoess and Scholl 1999a, 1999b, 2001; Hoess 2000; Oeyen and We- sener 2015; Wesener 2015a, 2015b, 2018; Wilbrandt et al. 2015; Wesener and Conrad 2016; Reip and Wesener 2018; Anti¢ et al. 2021). One of the taxa that have been forgotten and complete- ly excluded from the European fauna is Glomeris her- zogowinensis Verhoeff, 1898. Verhoeff (1898) described this taxon under the name “Glomeris europaea, herzo- gowinensis” on the basis of specimens from near Trebin- je, Herzegovina, collected by Victor Apfelbeck, the then curator of the National Museum of Bosnia and Herzegov- ina in Sarajevo. Although the description of this taxon is relatively short, Verhoeff (1898) already points out in the first sentence: “...der marginata in der Farbung 4usserst ahnlich...”, indicating a great similarity in colouration between G. herzogowinensis and one of the most com- mon western-central-northern European species, G/lom- eris marginata (Villers, 1789). Albeit geographically completely separate, both species are characterised by mostly black, shiny tergites with yellowish or white pos- terior margins. Verhoeff (1898) noted several differences between the two taxa, including details of the telopods, although he never illustrated them. This deficiency led to G. herzogowinensis falling into oblivion. Two years later, Verhoeff (1901: 248, 249) cited G. herzogowinensis from several localities in Albania and Greece, apparently only on the basis of a large, dark body with lighter posteri- or margins, evidently without examining the telopods of males from Greece. Later, it will turn out that, in these parts of Albania and Greece, there 1s or are one or even two species similar in appearance to G. herzogowinensis, but belonging to a different genus, Onychoglomeris Ver- hoeff, 1906. The fact that Verhoeff did not actually exam- ine the telopods of the Greek male specimens is support- ed by the fact that, when establishing the then subgenus, Onychoglomeris, he included in it what we know today as Onychoglomeris tyrolensis (Latzel, 1884) and Simplom- eris montivaga (Faés, 1902) (Verhoeff 1906). The telo- pods of these species differ markedly from those of the genus Glomeris and G. herzogowinensis. In his contri- bution to the knowledge of the genus Glomeris, Verhoeff (1911: 118, 119) included Glomeris herzogowinensis in the marginata species-group, stating some of the charac- teristics of the species. It is clear from the above that the species he described from the Trebinje area really belongs to the genus Glomeris. However, the problem emerged in the papers of Attems (1929, 1935), after which the species name Glomeris herzogowinensis was no longer mentioned. Strangely and without any explanation, Attems (1929: 289, 312) listed Verhoeff’s species under the name “Onychoglomeris herzegovinensis Verh.”. The crux of the problem with this taxon happened six years later. zse.pensoft.net Probably confused by Verhoeff’s (1901: 248, 249) ear- lier (obviously incorrect) record of G. herzogowinensis from Albania and Greece and the confusing similarity in the habitus between the latter species of which he received some syntypes and the specimens of the ge- nus Onychoglomeris from Albania and Greece he was studying, Attems (1935) just treated the species G. her- zogowinensis as Onychoglomeris herzogowinensis. At- tems (1935) was not sure of his act, especially because the structure of the telopods of G. herzogowinensis was unknown to him. He stated that only Verhoeff could clarify this by examining the telopods, although Attems himself could have done so (see below in Remarks un- der G. herzogowinensis). Despite this error, Attems was, however, right in the fact that his new specimens be- longed to the genus Onychoglomeris. He described two taxa: Onychoglomeris herzegovinensis media Attems, 1935 from Albania and O. h. australis Attems, 1935 from Greece (Attems 1935). He treated the taxon from Croatia, Bosnia and Heregovina and Montenegro as the nominotypical subspecies O. h. herzegovinensis (herzo- gowinensis is the correct spelling in all cases). Attems (1935) provided illustrations of the telopods of the two subspecies, which undoubtedly speak in favour of the genus Onychoglomeris, but at the same time, he pointed out some differences in habitus between his subspecies on the one hand and the nominotypical subspecies dis- tributed further north on the other. Six decades later, Mauries et al. (1997), based on rel- atively abundant material of Onychoglomeris from Alba- nia, but without studying the specimens from the type lo- cality of G. herzogowinensis nor the Greek specimens of Onychoglomeris, questioned the existence of three sub- species, considered all under the name Onychoglomeris herzogowinensis. The name appeared as such in Thaler (1999) and Kime and Enghoff (2011). Based on newly-collected material from near the type localities and on the study of the syntypes and historical specimens of Verhoeff’s G. herzogowinensis and Attems’ subspecies O. h. australis and O. h. media, we revive Verhoeff’s species Glomeris herzogowinensis after al- most nine decades and we consider both of Attems’ sub- species as species, viz. Onychoglomeris australis Attems, 1935 stat. nov. and Onychoglomeris media Attems, 1935 Stat. nov. Materials and methods Live specimens were collected by hand and preserved in 70% ethanol for mophological and 96% ethanol for DNA analyses. Several live individuals of Glomeris herzogowinensis were first placed in glass vials contain- ing 500 ul methylene chloride (DCM) for 5 minutes to extract their defensive secretions for future semiochem- ical studies. Later, the specimens were transferred to 70% ethanol. Zoosyst. Evol. 100 (2) 2024, 493-513 Depository IZB Institute of Zoology, University of Belgrade — Faculty of Biology, Belgrade, Serbia NHMW Naturhistorisches Museum Wien, Vienna, Austria ZFMK Zoological Research Museum A. Koenig, Leibniz Institute for Biodiversity Change, Bonn, Germany ZMB Museum fiir Naturkunde Berlin, Germany ZSM Zoologische Staatssammlung Munchen, Ger- many Morphology, photography and map Specimens were examined with a Nikon SMZ 25 (NHMW), Nikon SMZ 745T, Nikon SMZ 1270 (IZB) or Olympus SZX12 (ZFMK) binocular stereomicroscopes. Old microscopic preparations were examined with a Nikon SMZ 25 (NHMW) binocular stereomicroscope or with a Carl Zeiss Axioscope 40 microscope (IZB). Photo- graphs of habitus, leg pairs 17 and 18 and telopods were taken using a Nikon DS-Ri-2 camera mounted on a Nikon SMZ25 binocular stereomicroscope using NIS-Elements Microscope Imaging Software with an Extended Depth of Focus (EDF) patch (NHMW, Figs 2-6, 8—11A—D, G) or with a Nikon DS-Fi2 camera with a Nikon DS-L3 camera controller attached to a Nikon SMZ 1270 binoc- ular stereomicroscope (IZB, Fig. 11E, F). The photos of the living animals were taken with a Canon PowerShot SX530 HS (Fig. 7A, B), Olympus Stylus Tough TG-6 (Fig. 7C, D), Nikon D750 (Fig. 12A, B) and Panasonic DMC-G81 (Fig. 12E, F) digital cameras as well as with a cellphone (Fig. 12C, D). The distribution map was creat- ed using Google Earth Pro (version 7.3.6.9750) and Ado- be Photoshop CS6. The final images were processed and assembled in Adobe Photoshop CS6. DNA extraction, amplification and sequencing In order to find close relatives to Glomeris herzogowin- ensis, as well as Onychoglomeris australis stat. nov., a DNA barcoding analysis (Hebert et al. 2003) was con- ducted. COI sequences of both taxa, as well as those of potential related Glomeris species, such as G. balcanica Verhoeff, 1906 and G. pulchra Koch, 1847 and addition- ally Onychoglomeris ferraniensis Verhoeff, 1909 were analysed (see Table 1). In addition, sequences of simi- larily coloured (= black) Glomeris species were down- loaded from GenBank: Glomeris marginata (Villers, 1789) from Central Europe, G. apuana Verhoeff, 1911 from the Apuan Alps and G. maerens Attems, 1927 from Spain. Additionally, sequences of widespread species occurring in the Balkans and surrounding areas were added from GenBank: G. pustulata Latreille, 1804, 495 G. hexasticha Brandt, 1833, G. tetrasticha Brandt, 1833 and G. klugii Brandt, 1833. As outgroup taxa, sequences of Glomeridella minima (Latzel, 1884) and Tonkinomeris huzhengkuni Liu & Golovatch, 2020 from the family Glomeridellidae Cook, 1896 were added. Our dataset included 25 COI sequences from 15 spe- cies, of which eight sequences from five species were newly sequenced. The DNA extraction, amplification and sequencing protocol was similar to earlier studies (Wesener 2015a; Sagorny and Wesener 2017), using the degenerate (As- trin and Stiben 2008) primer pair HCO-JJ/LCO-JJ (HCOJJ AWACTTCVGGRTGVCCAAARAATCA/ LCOJJ CHACWAAYCATAAAGATATYGG). — Se- quences were concatenated by hand or by utilising the software Seqman (DNASTAR Inc.). BLAST search- es (Altschul et al. 1997) were performed to confirm sequence identities. The whole dataset was translat- ed into amino acids to rule out the accidental amplifi- cation of pseudogenes. The eight new sequences have been uploaded to GenBank under the accession codes PP475126—PP475133 (Table 1). All sequences were aligned in Bioedit (Hall 1999). The number of base differences per site (p-distances) between sequences was calculated (See Suppl. material 1). The analysis involved 25 nucleotide sequences. Co- don positions included were 1°+2"+3". All ambiguous positions were removed for each sequence pair. There were a total of 657 positions in the final dataset. Evolu- tionary analyses were conducted in Megall (Tamura et al. 2021). The best fitting substitution model for a Maximum Likelihood analysis was calculated with ModelTest (Tamura and Nei 1993) as implemented in MEGA11. The best fitting model was the general time reversible (GTR)-Model (Tavaré 1986) with gamma distribution and invariant sites (GTR+G+I) (InL = -4292.222, Invari- ant = 0.609, Gamma = 0.624, Freq A: 25.7, T: 38.89, C: 14.17, G: 21.24). The evolutionary history was inferred by using the Maximum Likelihood method and the General Time Re- versible model (GTR+G+I) (Nei and Kumar 2000). The tree with the highest log likelihood (-4292.19) is shown in Fig. 1. Initial trees for heuristic search were automatically obtained by applying Neighbour-Joining and BioNJ algo- rithms to a matrix of pairwise distances estimated using the Maximum Composite Likelihood (MCL) approach. Codon positions included were 1*-2"'-3". All positions with less than 95% site coverage were eliminated, L.e. fewer than 5% alignment gaps, missing data and ambig- uous bases were allowed at any position (partial deletion option). There were a total of 657 positions in the final dataset. The bootstrap consensus tree was calculated from 1000 replicates (Felsenstein 1985) in MEGA11 (Tamura et al. 2021). The obtained tree was edited in Adobe II- lustrator 2023 with all bootstrap values > 50% illustrated (Fig. 1). zse.pensoft.net 496 Anti¢, D. et al.: Glomeris herzogowinensis and Onychoglomeris from the Balkans Table 1. Newly-analysed specimens, vouchers and GenBank numbers. More detailed localities are only given for newly-sequenced specimens. Abbreviations: SCAU = South China Agricultural University, Guangzhou, China; ZSM = Bavarian State Collection, Munich, Germany; ZFMK = Zoological Research Museum Koenig, Leibniz Institute for the Analysis of Biodiversity Change (LIB), Bonn, Germany. Species Glomeridella minima (Latzel, 1884) Tonkinomeris huzhengkuni Liu & Golovatch, 2020 Glomeris pustulata Latreille, 1804 Glomeris pustulata Latreille, 1804 Glomeris hexasticha Brandt, 1833 Glomeris hexasticha Brandt, 1833 Glomeris tetrasticha Brandt, 1833 Glomeris tetrasticha Brandt, 1833 Glomeris marginata Villers, 1789 Glomeris marginata Villers, 1789 Glomeris maerens Attems, 1927 Glomeris maerens Attems, 1927 Glomeris klugii Brandt, 1833 Glomeris klugii Brandt, 1833 Glomeris apuana Verhoeff, 1911 Glomeris apuana Verhoeff, 1911 Onychoglomeris tyrolensis Latzel, 1884 Glomeris pulchra Koch, 1847 Glomeris pulchra Koch, 1847 Glomeris balcanica Verhoeff, 1906 Onychoglomeris ferraniensis Verhoeff, 1909 Onychoglomeris ferraniensis Verhoeff, 1909 Onychoglomeris australis Attems, 1935 stat. nov. Glomeris herzogowinensis Verhoeff, 1898 Glomeris herzogowinensis Verhoeff, 1898 Results Analysis of the COI barcoding gene All species were recovered with high statistical sup- port (94-100%, Fig. 1), while deeper nodes and inter- specific relationships were statistically not supported. Neither the families Glomeridae and Glomeridellidae, nor the genus G/omeris are recovered as monophylet- ic (Fig. 1). Glomeris herzogowinensis does not group with Onychoglomeris species, but is in an unsupport- ed sister-group with the similarily coloured G. mae- rens from the Mediterranean coast of Spain (Fig. 1). G. herzogowinensis and G. maerens show also the low- est genetic distance to one another (11.9-12.6%), while G. herzogowinensis also shows lower genetic distanc- es to the similarily coloured (black) G. apuana (12.2- 12.9%) and the Balkan G. balcanica (12.6—-13.4%), while it shows genetic distances of 13.4-16.4% to all other analysed species. The genus Onychoglomeris is recovered as monophyletic with moderate statistical support (74), with O. australis stat. nov. and the Italian O. tyrolensis in a weakly-supported sister-group (54, Fig. 1). O. australis stat. nov. shows the lowest genetic distance to O. tyrolensis (10.5%) and O. ferraniensis (11.1-11.4%), while it differs from species of the other genera by 12.8—-15.7%. zse.pensoft.net Taxonomy Locality Voucher # GenBank # Austria ZSM MYR 00371 JN271878 China SCAU TY0O1 MT522013 Germany ZSM MYR 00024 HM888093 Germany ZSM MYR 00376 JN271880 Germany ZFMK MYR1460 MG931023 Germany ZFMK MYR3898 MG931024 Germany ZSM MYR 00036 HM888105 Germany ZSM MYR 00035 HM888104 France ZFMK MYR6084 MG931022 Luxembourg ZFMK MYR1363 MG931021 Spain ZFMK MYR6097 MG892108 Spain ZFMK MYR6092 MG892110 Italy ZFMK MYR637 KX714076 Italy ZFMK MYR4734 KX714072 Italy ZFMK MYR753 KT188944 Italy ZFMK MYR752 KT188943 Italy ZFMK MYR1276 KP205571 Croatia, Dalmatia, Cetina River ZFMK MYR8217 PP475126 Croatia, Dalmatia, Cetina River ZFMK MYR8217b PP475127 Greece, Dion-Olympos ZFMK MYR11331 PPA/5128 Italy, Piemonte, Cuneo, Ceva ZFMK MYR623 PP475129 Italy, Piemonte, Cuneo, Ormea ZFMK MYR2287 PP475130 Greece, Kalambaka ZFMK MYR11332 PP475131 Bosnia & Herzegovina, Trebinje, Taleza ZFMK MYR8970 PP475132 Bosnia & Herzegovina, Trebinje, Taleza ZFMK MYR8969 PP475133 Class Diplopoda de Blainville in Gervais, 1844 Order Glomerida Brandt, 1833 Family Glomeridae Leach, 1816 Subfamily Glomerinae Leach, 1816 Genus Glomeris Latreille, 1802 Glomeris herzogowinensis Verhoeff, 1898 Figs 1-7 Glomeris europaea, herzogowinensis Verhoeff, 1898: 163, fig. 18. not Glomeris herzegowinensis (sic!).— Verhoeff (1901: 248). not Glomeris herzogowinensis — Verhoeff (1901: 249). Gl. herzegowinensis (sic!)— Verhoeff (1906: 211). herzegowinensis (sic!).— Verhoeff (1911: 119). [in the genus Glomeris]. Onychoglomeris hercegovinensis (sic!) in part—Attems (1929: 289, 312). Onychoglomeris hercegovinensis hercegovinensis (sic!).— Attems (1935: 149). Onychoglomeris hercegovinensis hercegovinensis (sic!).— Attems (1959: 323). Onychoglomeris herzegowinensis (sic!).— Strasser (1971: 12). not Onychoglomeris herzegowinensis (sic!).— Thaler (1999: 198, 199, figs 16, 17). Glomeris marginata.— Ceuca (1990: 10). Onychoglomeris herzogowinensis in part.— Kime and Enghoff (2011: 34, 118). Zoosyst. Evol. 100 (2) 2024, 493-513 100 497 MG931022.1 Glomeris marginata MG931021.1 Glomeris marginata MT522013.1 Tonkinomeris huzhengkuni 94 MG892110.1 Glomeris maerens MG892108.1 Glomeris maerens TW359 Glomeris balcanica 100 KT188943.1 Glomeris apuana KT188944.1 Glomeris apuana 100 fF ZFMK-TIS-2544369 Glomeris pulchra ZFMK-TIS-2544370 Glomeris pulchra 100 | HM888093.1 Glomeris pustulata JN271880.1 Glomeris pustulata | | 100 HM888105.1 Glomeris tetrasticha HM888104.1 Glomeris tetrasticha KX/14072.1 Glomeris klugil 99 MG931024.1 Glomeris hexasticha we KX/140/6.1 Glomeris klugii | | | 100 MG931023.1 Glomeris hexasticha — es 0,10 JN271878.1 Glomeridella minima Figure 1. Maximum Likelihood tree (ML) of millipedes of the order Glomerida, based on 657 bp of the COI gene. Yellow box = Onychoglomeris Verhoeff, 1906; blue box = Glomeris herzogowinensis Verhoeff, 1898. Numbers on nodes are bootstrap values from the ML analysis and are shown when > 50%. Diagnosis. Large species (up to 20 mm) with mostly black, shiny tergites with contrasting yellowish or white posterior margins. Similar to G. marginata in general appearance, but differs by strongly-pronounced light-co- loured anterolateral margins of the thoracic shield which is in the form of a narrow band in G. marginata. Addi- tionally, G. herzogowinensis has two complete or almost complete striae on the thoracic shield (tergite 2), while G. marginata has one complete stria. Material studied. Lectotype. 1 male (NHMW 3903); BOSNIA AND HERZEGOVINA, Trebinje; V. Apfelbeck leg.; K. Verhoeff don. 1897. Lectotype here designated. zse.pensoft.net 498 Anti¢, D. et al.: Glomeris herzogowinensis and Onychoglomeris from the Balkans Paralectotypes. @ | male, slide preparation (ZMB- MYR12772) (Verhoeff slide 953): leg pair 18 and telo- pods; Trebinje. e 1 male, 1 female (ZMB-MYR2261); Trebinje. ? Types. e | female (NHMW MY10415); Herzegovi- na: K. Verhoeff don. 1899. (Although this female arrived later in the NHMW collection than the lectotype, it may well represent another type specimen collected by Ap- felbeck near Trebinje). e 2 tubes (ZSM-A 20070848), 1 whole male, | male dissected (missing telopods and pos- terior leg pairs), 1 female, 1 juvenile; “ehemals Trock- enmaterial” [material previously dry], Etk Nb. 28; Her- zegowina. @ | tube (ZSM-A20070848), (Etk Nb. 28): 1 entire female specimen, a detached collum and thoracic shield, “ehemals Trockenmaterial; Tier m Original deter- minat. Etikett C Typtis-verdachtig” [material previously dry, animal with original determination, probable type], Trebinje. @ 1 male, slide preparation (ZSM-A2003 1802): telopods, leg pairs 716, 17 and 18; Schuma (= Suma, karst region around Trebinje). Other material examined. BosNiIA AND HERZEGOVI- NA: @ | female (NHMW MY10414); Trebinje e 2 males, 1 female (IZB); in front of Taleza Cave, Taleza Village, near Trebinje, under stones; 15 November 2019; D. Anti¢ leg. e 3 males, 7 females (IZB); same locality as previous; 8 April 2022; D. Anti¢ and D. Stojanovic leg. @ 1 female (IZB); in front of Pavlova Cave, Bihovo Village, near Trebinje, under a stone; 16 November 2019; D. Antic leg. e | female (IZB); same as previous but inside Pavlova Cave. Croatia: @ 1 female (NHMW MY10427); Prid- vorje @ 2 females (ZFMK MYR89); Dubrovnik-Neretva, Konavle Region, Gruda, Konavoski dvori, under stones close to river, 50 m elev.; 3 April 2010; R. Ozimec & A. Schonhofer leg. e 1 male (ZFMK MYR95); Du- brovnik-Neretva, Konavle Region, Vignje, near Sklenica Cave, under stones in dense, humid, mossy forest, 89 m elev.; 3 April 2010; R. Ozimec & A. Schonhofer leg. e 3 males (ZFMK MYRI153); Dubrovnik-Neretva, Konav- le region, Vignje, Spilja at Vignje Cave; under stones: 3 April 2010; R. Ozimec & A. Schonhofer leg. e 1 fe- male (ZFMK MYR173); Dubrovnik-Neretva, Konavle Region, Vignje, surroundings of entrance of Tunnel of Konavle Polje, under stones, 50 m elev.; 3 April 2010; R. Ozimec & A. Schonhofer leg. MONTENEGRO: @ 1 male, 4 females (NHMW MY10413); Savina e 1 male (IZB) ethanol and slide with leg pairs 17 and 18 and telopods; Orjen Mountain, BaljeSina Lokva, 1400 m elev.; 4 July 1997; I. Karaman leg. @ 1 female (ZFMK MYR220); Ru- mija Mountain, near Sutorman, sieving in oak forest near rocks and under stones along open path, 42°9'22.8"N, 19°6'32.1"E, 805 m elev.; 9 May 2006; A. Schonhofer leg. e 1 ex.; Borovik, near Cetinje; 11 May 2011; D. An- tic observed. Remarks. After examining type and old museum specimens, as well as freshly-collected animals, we confidently conclude that Verhoeff’s herzogowinensis has typical Glomeris telopods. Attems (1935) examined zse.pensoft.net Figure 2. Glomeris herzogowinensis Verhoeff, 1898, lectotype male (NHMW MY3903), habitus. A. Dorsal view; B. Lateral view; C. Ventral view. Scale bar: 1 mm. Verhoeff’s material sent to the NHMW and listed that they were both females, so there was no possibility of examining the telopods. Interestingly, we found and examined these two specimens, among which one re- vealed to actually be a male (now lectotype, see Figs 2, 3). In addition, Attems (1929) indicated the locality Savina in Montenegro as one of the collecting cites of Onychoglomeris hercegovinensis (sic!). We found one male (Fig. 4) among five specimens from this locality, again with typical Glomeris telopods (Fig. 4D). Thus, Attems missed the opportunity to see the telopods in two males, including Verhoeff’s syntype and to conclude that it was, indeed, a species of the genus Glomeris and not of Onychoglomeris. Glomeris herzogowinensis shows a striking resem- blance with G. marginata, both in habitus (Figs 5-7) and in the structure of the telopods (Figs 3C, 4D), which are almost identical in both species. Verhoeff (1898) pointed out that G. herzogowinensis has more promi- nent light-coloured posterolateral margins compared to G. marginata. However, this is not entirely correct, as one individual analysed by us (Fig. 6A) has identi- cal margins to most G. marginata. Indeed, most of the studied specimens of G. herzogowinensis have more pronounced margins than the classic G. marginata, but Zoosyst. Evol. 100 (2) 2024, 493-513 499 Figure 3. Glomeris herzogowinensis Verhoetf, 1898, lectotype male (NHMW MY3903). A. Leg pair 17, anterior view; B. Leg pair 18, anterior view; C. Telopods, anterior view. Scale bars: 0.2 mm. some French populations of the latter present postero- lateral margins that are more developed than in G. her- zogowinensis (see Reip and Wesener (2018: 96, fig. 1D, E)). Verhoeff (1898, 1911) mentioned the presence of strongly-pronounced light-coloured anterolateral mar- gins of the thoracic shield as one of the most important features distinguishing these two species. Indeed, the thoracic shield of all examined individuals of G. herzo- gowinensis has very distinct anterolateral margins (Figs 5A, C, 6B, D, E), in contrast to G. marginata, where it 1s only present in the form of a narrow band. The coloura- tion of the fresh specimens that we have analysed corre- sponds completely to the description of Verhoeff (1898). They are mostly black with clearly demarcated lighter, whitish or yellowish posterolateral margins of the terg- ites. The collum also has a lighter posterior margin, as does the anal shield. As already mentioned, the thoracic shield also has a pronounced anterolateral margin. Some specimens are characterised by the presence of a pair of pale marbled patches on the tergites, including the thoracic shield, as well as an unpaired patch on the anal shield (Figs 5A, B, D, 6B). The presence of demarcated posterolateral light-coloured margins is clearly visible in old museum specimens too (Figs 2A, B, 4A). Verhoeff (1898) listed some differences in the structure of the telopods, but they were apparently so insignificant that he never drew these structures. Nevertheless, in this paper, we present for the first time illustrations of the telo- pods of G. herzogowinensis, as well as of the 17" pair of legs and the entire 18" pair of legs (Figs 3, 4B—D), which are of typical Glomeris appearance. We would also like to mention that all examined spec- imens show two transverse ridges on the collum (Figs 5C, 6B, E). Verhoeff (1911) found that, in addition to the two characteristic ridges, a third ridge starts on both sides of the collum. In the fresh material, the beginning of the third ridge was only observed in one specimen and only on the left side. Concerning the thoracic shield (tergite 2), Verhoeff (1898) distinguished G. herzogow- inensis from G. marginata by the presence of two com- plete striae and an incomplete one (2+1 vs. 1+2, 1+1 or 1+0 sensu Schubart (1934: 33, fig. 28)). Indeed, all but zse.pensoft.net 500 Anti¢, D. et al.: Glomeris herzogowinensis and Onychoglomeris from the Balkans Figure 4. Glomeris herzogowinensis Verhoeff, 1898, male from Savina, Montenegro (NHMW MY 10413). A. Habitus, dorsal view; B. Leg pair 17, anterior view; C. Leg pair 18, anterior view; D. Telopods, anterior view. Scale bars: 1 mm (A); 0.5 mm (B-D). two of the specimens examined show two complete stri- Habitat. Known from almost near sea level up to ae and an incomplete one. In two specimens, the second 1400 m elev. in the Orjen Mountain. Scrubs of Carpi- stria is almost complete, with only a small interruption nus, Quercus, Juniperus, under stones in limestone ar- dorsally. Some specimens are characterised by the pres- eas. Inside caves. ence of additional, 4" incomplete striae in front of the Distribution. The extreme south of Croatia and first complete one. Bosnia and Herzegovina, as well as the coastal part of zse.pensoft.net Zoosyst. Evol. 100 (2) 2024, 493-513 Figure 5. Glomeris herz 501 B. Habitus, dorsal view; C. Head, collum and thoracic shield, anterior view; D. Anal shield, posterior view. Scale bars: 1 mm. Montenegro (Fig. 13). Endemic south Dinaric coastal species. Croatia: Pridvorje (Attems 1929), Konavoski Dvori (Ceuca 1990, as G. marginata; present study), Gruda near Konavle (T. Drazina pers. comm.; present study), Vignje (present study); Bosnia and Herzegov- ina: Surroundings of Trebinje (Verhoeff 1898; Attems 1929, 1935), Taleza near Trebinje (present study), Bi- hovo near Trebinje (present study). Montenegro: Sav- ina (Attems 1929, 1935), Orjen (present study), Ru- mija, near Sutorman (present study), Cetinje, Borovik (present study). Type locality. Near Trebinje, Bosnia and Herzegovina. zse.pensoft.net 502 Antic, D. et al.: Glomeris herzogowinensis and Onychoglomeris from the Balkans : - ~ ~ aay . - git a - ~~ 7 ‘ is Fath . en *, at ¥ = aN Figure 6. Glomeris herzogowinensis Verhoeff, 1898, male (A—C) and female (D-F) from Taleza, Bosnia and Herzegovina (IZB). A. Habitus, lateral view; B. Collum and thoracic shield, anterior view, C. Anal shield, posterior view; D. Habitus, lateral view; E. Head, collum and thoracic shield, anterior view; F. Anal shield, posterior view. Scale bars: 1 mm. Genus Onychoglomeris Verhoeff, 1906 Onychoglomeris australis Attems, 1935, stat. nov. Figs 8, 9, 12A—D Onychoglomeris hercegovinensis australis (sic!).— Attems (1935: 150, figs 6-8). Glomeris herzogowinensis in part.— Verhoeff (1901: 249). Onychoglomeris herzegowinensis australis (sic!).— Strasser (1976: 580). Onychoglomeris herzegowinensis (sic!).— Thaler (1999: 198, 199, figs 16, 17). Onychoglomeris herzogowinensis in part.— Kime and Enghoff (2011: 34, 118). zse.pensoft.net Diagnosis. Similar in colouration (Fig. 12A—D) and mor- phology to the geographically very close O. media stat. nov., but differs in the appearance of the anal shield, leg pair 18 and telopods. Anal shield straight in lateral view (vs. distinctly concave in O. media stat. nov.). Leg pair 18 with short podomere 2, which is 1.5 times longer than wide, with straight mesal margin (vs. podomere 2 longer, twice as long as wide with distinctly convex mesal margin in O. media stat. nov.). Telopods apparently less robust, with a less developed posteriomesal process of telopo- ditomere 2 (= femur) and a shorter telopoditomere 4 (= tarsus), brownish stripes at the base of posteromesal pro- cess of telopoditomere 2 absent (vs. present in O. media Zoosyst. Evol. 100 (2) 2024, 493-513 503 (IZB); C, D. Specimen from TaleZa, Bosnia and Herzegovina (IZB). Photos by Dragan Antic¢. stat. nov.), telopoditomere 3 (= tibia) with a well-devel- oped posterior tooth that is more or less conical (vs. tooth poorly developed, subtriangular, sometimes almost ab- sent in O. media stat. nov.), the syncoxite is usually high, rounded (vs. syncoxite mostly lower, bilobed in O. media stat. nov.). For more details see remarks below. Material studied. Lectotype. 1 male (NHMW MY 10424): Greece, Epirus, Athamanika (= Tzoumerka) Mountain, Paraskevi, Abies, 1400 m elev.; 16 June 1933; M. Beier leg. Lectotype here designated. Paralectotypes. 13 males, 11 females (NHMW MY3900); same data as for lectotype. Including one slide (NHMW MY3900) with two pairs of leg pair 18, two pairs of leg pair 17 and additional leg ?17. Other material examined. All in Greece: @ 1 male (NHMW MY10418); Epirus, Buka Chalasmata near Plat- anoussa; 14 May 1932: M. Beier leg. @ 2 males, 2 females (NHMW MY10419); Epirus, Katarraktis; 1932/1933; M. Beier leg. @ 1 female (NHMW MY10416); Prosgoli; V. Apfelbeck leg. e 5 males (NHMW MY10420); Epirus, Aoos Gorge near Konitsa, 550 m elev., Carpinus; 9 Sep- tember 1996; K. Thaler and B. Knoflach leg. e 8 males, 5 females (NHMW MY10421); Epirus, Timfi Mountain near Micropapingo, 800 m elev., bush; 10 September 1996; K. Thaler and B. Knoflach leg. e 1 female (ZFMK MYR122); Epirus, Pindus Mountain, Zagori, Monoden- dri - Ano Pedina junction, under stone on the road, 835 m elev., 39.868002, 20.722076; 3 April 2006; A. Schonhofer leg. e 5 males, 5 females, 2 juveniles (ZFMK MYR4517); Epirus, Vikos Gorge, near Monodendri, Quercus forest with lichens, 1000 m elev., 39.881527, 20.755473; 4 April 2006; A. Schénhofer leg. @ 7 males, 2 females, (ZFMK MYR4518); Epirus, SW Ioannina, Zoodochos Pigi, open bushland with partly evergreen Quercus close to stream under stones, old tree trunks and sieving from leaf litter, 460 m elev., 39.56492, 20.72300; 13 August 2009; S. Hu- ber & A. Schonhofer leg. e 1 female (ZFMK MYR162); Thessaly, road to Kastanea, Elafi, Carpinus, Quercus, N-exposition, sieving from depressions in trees, 454 m elev., 39.723250, 21.475917; 1 April 2006; A. Schon- hofer leg. e 2 males, 4 females (ZFMK MYR124); Thes- saly, road E92a between Panagia and Metsovo; sieving in a damp, shady stream valley, moss and between stones, pine forest and alpine meadows, 1084 m elev., 39.80344, 21.306998; 2 April 2006; A. Schonhofer leg. @ 1 male (ZFMK MYR11332); Thessaly, Kalambaka, Meteora; September 2019; P. Knautt leg. e 2 males, 1 juvenile (ZFMK MYR11334); same data e 2 females (NHMW MY10417); Central Greece, Karpenisi; V. Apfelbeck leg. e | female “?type”, (ZSM-A20070858), Epirus. Remarks. Attems (1935), although he examined only a few males, already pointed out differences in the telo- pods between his australis and media, which we found to be constant after examining more males. The median lobe zse.pensoft.net 504 Anti¢, D. et al.: Glomeris herzogowinensis and Onychoglomeris from the Balkans Figure 8. Onychoglomeris australis Attems, 1935, stat. nov. A. Lectotype male (NHMW MY10424), habitus, lateral view; B-G. Males from Konitsa, Greece (NHMW MY 10420). B. Male 1, habitus, lateral view; C. Male 1, habitus, dorsal view; D. Male 1, anterior part of body, lateral view; E. Male 3, anal shield, lateral view; F. Male 1, collum, anterior view; G. Male 3, collum, anterior view; H. Female from Katarraktis, Greece (NHMW MY10419), collum, anterior view; I. Male from Katarraktis, Greece (NHMW MY10419), collum, anterior view. Scale bars: 1 mm. of the syncoxite is high and rounded distally (Fig. 9A, B, D, H) in all but one of the males examined. In one, it is lower and flattened distally (Fig. 9C), which looks more like an anomaly. Attems (1935: 150, fig. 7) also noted a strongly developed conical tooth on the telopoditomere 3 (= tibia). The same was clearly illustrated by Thaler (1999: 199, figs 16, 17). In the males examined by us, this zse.pensoft.net structure is always the same, conical and well developed (Fig. 9A, B, E, white arrows). Such a structure 1s men- tioned for O. media stat. nov. by Attems (1935) as much smaller compared to O. australis stat. nov. Our observa- tion was the same (see remarks under O. media stat. nov.). As one of the differences, Attems (1935: 150, fig. 6) mentioned the absence of the medial syncoxital lobe of Zoosyst. Evol. 100 (2) 2024, 493-513 505 Figure 9. Onychoglomeris australis Attems, 1935, stat. nov. A. Male 4 from Konitsa, Greece (NHMW MY 10420), telopods, anterior view; B. Male 4 from Konitsa, Greece (NHMW MY10420), telopods, posterior view; C. Paralectotype male 1 (NHMW MY3900), telopod syncoxite, posterior view; D. Paralectotype male 2 (NHMW MY3900), telopod syncoxite, posterior view; E. Male 4 from Konitsa, Greece (NHMW MY10420), part of right telopod, posterior view; F. Male 4 from Konitsa, Greece (NHMW MY10420), leg pair 17, anterior view; G. Male 4 from Konitsa, Greece (NHMW MY 10420), leg pair 18, anterior view; H. Male | from Konitsa, Greece (NHMW MY10420), leg pair 18 and telopods in situ, anterior view. White arrow indicates posterior tooth of telopoditomere 3. Scale bars: 0.5 mm. zse.pensoft.net 506 Antic, D. et al.: Glomeris herzogowinensis and Onychoglomeris from the Balkans leg pair 18 in O. australis stat. nov. However, after having checked all males available to us, we conclude that this feature is variable, as some males present this lobe (Fig. 9G). On the other hand, we found that podomere 2 is short and has a straight mesal margin (Fig. 9G, H), as also drawn by Attems (1935: 150, fig. 6), quite different from O. me- dia stat. nov. (see Remarks under O. media stat. nov.). In all examined males, the anal shield is predominantly straight in lateral view (Fig. 8A, B), in some only slightly concave (Fig. 8E), but never as distinct as in O. media stat. nov. (see below, Fig. 10A). Attems (1935) reported two transverse ridges on the collum. After examining all males and females, we found that this feature is variable and that, in addition to specimens with one (Fig. 8F) or two (Fig. 8H) ridges, there are also those with lateral be- ginnings of the second ridge (Fig. 8G) or that the second ridge is interrupted only in the centre (Fig. 81). We would like to emphasise that juveniles of this spe- cies are lighter in colour and are characterised by colour patterns that are not seen or not that obvious in adults and should not be confused with other glomerids from the re- gion (Fig. 12C, D). The two southernmost finds of this species in Central Greece were apparently misidentified as G. herzogowin- ensis by Verhoeff (1901: 249). Although Verhoeff stated that he had three males from Karpenisi, it is very likely that he did not check the telopods, but made his identi- fication on the basis of the very similar habitus with G. herzogowinensis. Unfortunately, we were unable to track down this Verhoeff material. We only found two females in the NHMW collection. As we were unable to look at the males, these two southernmost localities are marked with a question mark on the map. Habitat. From 170 m to 1400 m elev. Abies, Carpi- nus, Quercus, Juniperus, Pinus, under stones, under tree trunks, under mossy limestone debris, leaf litter in lime- stone areas, open areas, bushland. Distribution. Known from Epirus, Thessaly and central Greece (Fig. 13). Epirus: Paraskevi on Athamanika (At- tems 1935), Buka Chalasmata near Platanoussa (Attems 1935), Katarraktis (Attems 1935), Prosgoli (Verhoeff 1901 [missidentification]; Attems 1935), Graveniti (Strasser 1976), Elati (Strasser 1976), Ligiades (Strasser 1976), Metsovon (Strasser 1976), Filiate (Strasser 1976), Aoos near Konitsa (Thaler 1999), Timfi near Mikropap- ingo (Thaler 1999), Monodendri - Ano Pedina (present study), Zoodochos Pigi (present study). Thessaly: Kas- tanea, Elafi (present study), Panagia (present study), Ka- lambaka (present study). Central Greece: Karpenisi (Ver- hoeff 1901 [missidentification]; Attems 1935), Velouchi on Tymfristos (Verhoeff 1901 [missidentification]). Type locality. Paraskevi, Epirus, Greece. Attems (1935: 143) stated: “Paraskevi ist ein Gipfel des Cumer- ka-Gebirges” which translates that Paraskevi is a summit on the Tzoumerka (= Athamanika) Mountain. We could not find out where exactly Paraskevi is located. zse.pensoft.net Onychoglomeris media Attems, 1935, stat. nov. Figs 10, 11, 12E, F Onychoglomeris hercegovinensis media (sic!).— Attems (1935: 149, figs 4, 5). Onychoglomeris hercegovinensis (sic!) in part— Attems (1929: 289, 312), Onychoglomeris herzogowinensis— Mauries et al. (1997: 258-260, fig. 2). Onychoglomeris herzegowinensis (sic!).— Curéi¢ et al. (1999: 11P). Onychoglomeris herzogowinensis in part—— Kime and Enghoff (2011: 34, 118). Glomeris herzogowinensis.— Verhoeff (1901: 248). Glomeris herzogowinensis in part.— Verhoeff (1901: 249). ?Glomeris marginata— Sekulié and Zivié (2017: 193). [Missidentifica- tion, but see Remarks below]. Diagnosis. Similar in colouration (Fig. 12E, F) and mor- phology to the geographically very close O. australis stat. nov., but differs in the appearance of the anal shield, leg pair 18 and telopods. Anal shield distinctly concave in lat- eral view (vs. straight in O. australis stat. nov.). Leg pair 18 with podomere 2 longer, twice as long as wide with distinctly convex mesal margin (vs. podomere 2 shorter, ca. 1.5 times longer than wide, with straight mesal mar- gin in O. australis stat. nov.). Telopods apparently more robust, with a well-developed posteriomesal process of telopoditomere 2 (= femur) and longer telopoditomere 4 (= tarsus), brownish stripes at the base of posteromesal process of telopoditomere 2 present (vs. absent in O. aus- tralis stat. nov.), telopoditomere 3 (= tibia) with a poor- ly developed, sometimes almost absent, posterior tooth that is subtriangular (vs. tooth well developed, conical in O. australis stat. nov.), the syncoxite 1s mostly low, some- what bilobed (vs. syncoxite usually higher and rounded in O. australis stat. nov.). For some more details, see Re- marks below. Material studied. Lectotype. 1 male (NHMW MY3901) in ethanol; ALBANIA, Dukati [= Dukat]; 5 Au- gust 1911; A. Winneguth leg. Including two slides: one with telopods, second one with leg pairs 16—18 and right leg 13 or 14. Lectotype here designated. Paralectotypes. e 2 females (NHMW MY10425); same data as for lectotype; @ 1 female (NHMW MY3902); ALBANIA, Kanina [= Kaniné]; November 1908; A. Win- neguth leg. Other material examined. ALBANIA: @ 1 male, 1 female (NHMW MY10412); Valona [= Vloré]; Dr. K. Patsch leg.; @ 5 males, 2 females (IZB); Gjirocastro [= Gyirokastér]; 10 May, 1973; M. Karaman leg. e 1 male, 2 females (ZFMK MYR13662); Gyjirokastér District, Vjosa Valley, Pérmet, Strémbec, hiking trail Ri Soptit Waterfall; forest of low Carpinus, Quercus, Platanus and Cratae- gus, in leaf litter, 40.1488, 20.4543; 6 October 2023; H. Reip leg. SERBIA: @ | male (IZB), slide with a male telo- pods and leg pairs 17 and 18; Visoki Decani, Kosovo and Metohiya; 1973; M. Karaman leg. Zoosyst. Evol. 100 (2) 2024, 493-513 507 *-: — _S Figure 10. Onychoglomeris media Attems, 1935, stat. nov., lectotype male (A-C, NHMW MY3901) and paralectotype female (D-F, NHMW MY10425). A, D. Habitus, lateral views; B, E. Habitus, dorsal views; C, F. Collum, anterior and anterodorsal views respectively. Scale bars: 1 mm. Remarks. As written above, one of the differences be- tween O. media stat. nov. and O. australis stat. nov. is amuch smaller tooth of telopoditomere 3 (= tibia) of the telopods in O. media stat. nov. In all males we had, this tooth is poorly developed and sometimes almost absent (Fig. 11D—F, white arrows). This structure was probably overlooked by Mauries et al. (1997). It is interesting to note that, at the base of the strongly-developed posteromesal process of telopoditomere 2 (= femur), one or more brownish darker stripes were ob- served in all males available to us (Fig. 11A, C, E, black arrows). Such stripes are absent from all males of O. aus- tralis stat. nov. at hand. In comparison with O. australis stat. nov., podomere 2 of leg pair 18 is longer and has a convex mesal margin that looks somewhat like a blade (Fig. 11B, G). The medial syncoxital lobe of leg pair 18 may be present or absent as in O. australis stat. nov. (Fig. 11B, G; see also Mauries et al. (1997: 259, fig. 2B, F)). All males at our dis- posal have a distinctly concave anal shield (Fig. 10A). In contrast to Mauries et al. (1997), who found consistency in Albanian specimens with regard to the presence of only one transverse ridge on the collum, we found it variable as in O. australis stat. nov. with one or two complete ridges, sometimes a second only as lateral remains (Fig. 10C, F). It is of interest to mention a very isolated find in southern Serbia, near Visoki Deéani. This site is almost 250 km by air from the nearest site in the core area of southern Alba- nian sites (Fig. 13). In the IZB collection, only the micros- lide with the telopods and the leg pairs 17 and 18 have been found so far and both the telopods and the leg pair 18 fit into the concept of O. media stat. nov. Whether it was a mistake zse.pensoft.net i 1B Anti¢, D. et al.: Glomeris herzogowinensis and Onychoglomeris from the Balkans 4 . +>. ' * lo - ie 1m e a iy r Figure 11. Onychoglomeris media Attems, 1935, stat. nov. A. Lectotype male (NHMW MY3901), telopods, anterior view; B. Lec- totype male (NHMW MY3901), leg pair 18, anterior view; C. Male from Vloré, Albania (NHMW MY10412), telopods, anterior view; D. Male from Vloré, Albania (NHMW MY10412), telopods, posterior view; E. Male from Gjirokastér, Albania (IZB), part of the left telopod, anterior view; F. Male from Gjirokastér, Albania (IZB), part of the left telopod, posterior view; G. Male from Vloré, Albania (NHMW MY10412), leg pair 18, anterior view. White arrows indicate posterior tooth on telopoditomere 3, black arrows indicate characteristic brownish stripes of telopoditomere 2. Scale bars: 0.5 mm. in labelling or the species is really so widespread must be clarified in the future. The latter is supported by the fact that Sekuli¢ and Zivié (2017) recorded the occurrence of Glomeris marginata in southern Serbia (Znosek, Leposav- ic), about 80 km north-east of Visoki Deéani. It is obvious that this is not G. marginata, but it remains questionable which species Sekuli¢ and Zivié (2017) actually found. For the purposes of this paper, we will refer to these two Serbian records as O. media stat. nov. with a question mark. zse.pensoft.net Habitat. There is no information about the habitat of this species in the literature, except that Mauries et al. (1997) mentioned that specimens were found under stones and in leaf litter. Considering the distribution of the species, the habitat should be considered the same as for G. herzogowinensis and O. australis stat. nov. According to new data, it can be found in Carpinus, Quercus, Platanus and Crataegus forest, in leaf litter. Zoosyst. Evol. 100 (2) 2024, 493-513 509 Figure 12. Living specimens. A-D. Onychoglomeris australis Attems, 1935, stat. nov., specimens from Kalambaka, Greece. E-F. Onychoglomeris media Attems, 1935, stat. nov., specimens from Pérmet, Albania. Photos by Morris Fleck (A, B), Peter Kautt (C, D) and Hans Reip (E, F). Distribution. Southern Albanian species with a single, isolated locality in southern Serbia (Fig. 13). Albania: Du- kat (Attems 1935; Mauries et al. 1997), Kaniné (Attems 1935); Vloré (Verhoeff 1901 [missidentification]; Attems 1929 [missidentification], 1935); Dhérmi (Mauries et al. 1997), Himaré (Mauries et al. 1997), Llogara Pass (Mau- ries et al. 1997), Gjirokastér (Mauries et al. 1997; pres- ent study), Pérmet (present study). Serbia: Visoki De¢éani (Curéi¢ et al. 1999), ?7Leposavié (Sekulié and Zivicé 2017 [missidentification]). Type locality. Dukat, Vloré County, southern Albania. Discussion Our DNA barcoding analysis clearly confirms the results of the morphological analysis of the telopods: Glomer- is herzogowinensis groups with other Glomeris species and not with Onychoglomeris, while O. australis stat. nov. clearly groups with Onychoglomeris. Interesting- ly, the sister species to G. herzogowinensis seems to be G. maerens from Spain, a similarly-coloured species liv- ing in a similar Mediterranean habitat. However, there are indications that more than one species 1s currently hiding zse.pensoft.net 510 Anti¢, D. et al.: Glomeris herzogowinensis and Onychoglomeris from the Balkans Glomeris herzogowinensis Verhoeff, 1898 Onychoglomeris australis Attems, 1935 stat. nov. ap Onychoglomeris media Attems, 1935 stat. nov. Figure 13. Distribution map of Glomeris herzogowinensis Verhoeff, 1898, Onychoglomeris australis Attems, 1935, stat. nov. and Onychoglomeris media Attems, 1935, stat. nov. under the name G. maerens (Reip and Wesener 2018). The observed genetic distances of the COI barcoding gene between G. herzogowinensis and other Glomeris Species, as well as those between O. australis stat. nov. and other Onychoglomeris, are with 11-16% similarity to interspecific distances found in other barcoding stud- ies of species of the family Glomeridae (Wesener and Conrad 2016; Kuroda et al. 2022; Recuero and Caterino 2023), but lower than those observed in the diverse ge- nus Trachysphaera Heller, 1858 (Wilbrandt et al. 2015). The interspecific distances observed here fit well within the range observed in millipedes from other taxonomic groups and other geographic areas, such as the related (Oeyen and Wesener 2018) giant pill-millipedes (order Sphaerotheriida) from Madagascar (Wesener et al. 2014: Wesener and Sagorny 2021) and southeast Asia (Wesen- er 2019; Bhansali and Wesener 2022) or in Spirobolida from Madagascar (Wesener et al. 2011; Wesener 2020) and Thailand (Pimvichai et al. 2020, 2022). This work represents another example demonstrating the importance of natural history collections as a time- less resource allowing us to study organisms and their systematics, sometimes even discover and describe com- pletely unknown taxa, awaiting on shelves of museums to be determined, described and documented. The aver- age shelf-life of all kinds of species of living organisms was estimated to be around 20.7 years (see Fontaine et al. (2012)) with extreme cases exceeding 100 years like Pleo- nopurus tanzanicus Enghoff & Akkari, 2022 and reaching as high as 149 years such as Ommatoiulus schubarti Ak- kari & Enghoff, 2012 (Akkari and Enghoff 2012; Enghoff and Akkari 2022). In other cases, taxa have inadevertedly zse.pensoft.net been mixed with other hitherto described species, there- fore remaining hidden for decades. One of the latest ex- amples is perhaps that of Lophostreptus neglectus Eng- hoff & Akkari, 2024 discovered amongst the syntypes of its congener Lophostreptus regularis Attems, 1909 in two different collections in Sweden and Vienna and described more than a century after it was originally collected (Eng- hoff and Akkari 2024). The scientific collections, especial- ly type series and historical specimens, are most defintely an invaluable source of information for taxonomists to up- date information, unravel the identity of obscure historical names (e.g. Akkari et al. (2010); Akkari (2013); Anti¢ and Akkari (2020); Anti¢ et al. (2021)), clarify the taxonomic status of taxa and solve complicated riddles like the one presented in this work. Morphology-based taxonomy re- mains a subjective exercise, especially when the studied groups did not traditionally have well-defined characters for species characterisation, which 1s the case for the order Glomerida. Taxonomy is also very prone to human error and this has been illustrated in numerous cases, especial- ly in times when a tremendeous amount of taxa had to be described by a generation of taxonomists who did not enjoy the same advantages of communication means and technological facilities, not the least microscopy. Amend- ing these mistakes and updating the nomenclature of taxa, adding pieces of knowledge on their genetic information remains an ongoing process that make us acknowledge the colossal work accomplished by myriapod experts like Attems and Verhoeff, but also humble us once we also think towards the future and what could be achieved in perhaps less time given the same resources and further technological progress. Zoosyst. Evol. 100 (2) 2024, 493-513 In this article, we tried to solve the case of three spe- cies that have been hidden under the same name. G/lom- eris herzogowinensis was confirmed as an unquestionably good taxon. We have raised the other two taxa of the genus Onychoglomeris, former subspecies, to species level. Con- sidering the fact that we have no genetic data for O. media stat. nov. and that both O. media stat. nov. and O. australis stat. nov. very likely occur sympatrically at least in the Vjo- sa (in Albanian) or Aoos (in Greek) river valley in south- ern Albania and north-western Greece, respectively, some might disagree with such an act. In this context, and due to some morphological differences that obviously exist, we believe that the Albanian and Greek populations should be treated as separate species for the time being. Acknowledgements We are gratefull to Stefan Friedrich (ZSM) and Jason Dunlop (ZMB) for making the material under their care available to us for study. Oliver Macek (NHMW) kindly provided technical support handling the specimens and useful comments on the molecular part. Many thanks to the collectors Axel Schonhofer, Siegfried Huber, Hans Reip, Peter Kautt (all from Germany), R. Ozimec (Cro- atia) and I. Karaman (Serbia) for kindly providing spec- imens. In addition, Hans Reip, Peter Kautt and Morris Fleck (ZFMK) contributed beautiful photos (Fig. 12) of living Onychoglomeris australis Attems, 1935 stat. nov. and O. media Attems, 1935 stat. nov. Claudia Etzbauer and Jana Thormann extracted and sequenced the barcod- ing data at the ZFMK, for which we are very grateful. Last but not least, we would like to thank the reviewers Sergei Golovatch (Russia), Henrik Enghoff (Denmark) and Nikolaus Szucsich (Austria) for their comments and corrections, which have improved this manuscript. Luiz Felipe Iniesta (Brazil) helped with the editing of this manuscript. DA would like to thank his friends Dalibor Stojanovic (Serbia), Ivo Karaman (Serbia) and Marjan Komnenov (North Macedonia) for the wonderful time during the field trips in Bosnia and Herzegovina. 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Biodiversity Data Journal 3: e5176. https://doi.org/10.3897/ BDJ.3.e5176 Supplementary material | Number of base differences per site (p-distances) between sequences Authors: Dragan Antic, Thomas Wesener, Nesrine Akkari Data type: xls Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons. org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow us- ers to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited. Link: https://do1.org/10.3897/zse.100.122288 suppll zse.pensoft.net