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Research Article
Natural history collections help resurrecting Glomeris herzogowinensis Verhoeff, 1898 and further clarify the nomenclature of two Onychoglomeris subspecies of Attems (Diplopoda, Glomerida, Glomeridae)
expand article infoDragan Antić‡, Thomas Wesener§, Nesrine Akkari|
‡ University of Belgrade, Belgrade, Serbia
§ Leibniz Institute for the Study of Biodiversity Change, Zoogisches Forschungsmuseum Alexander Koenig, Bonn, Germany
| Natural History Museum Vienna, Vienna, Austria

Corresponding author: Dragan Antić ( dragan.antic@bio.bg.ac.rs )

Academic editor: Luiz Felipe Iniesta
© 2024 Dragan Antić, Thomas Wesener, Nesrine Akkari.
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.
Citation: Antić D, Wesener T, Akkari N (2024) Natural history collections help resurrecting Glomeris herzogowinensis Verhoeff, 1898 and further clarify the nomenclature of two Onychoglomeris subspecies of Attems (Diplopoda, Glomerida, Glomeridae). Zoosystematics and Evolution 100(2): 493-513. https://doi.org/10.3897/zse.100.122288
ZooBank: urn:lsid:zoobank.org:pub:CF9FDD8F-1432-4A0D-BA41-5D11B642110A
Open Access

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 Onychoglomeris australis stat. nov., along with the first barcoding data of one additional species of Onychoglomeris 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 today, 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 European species Glomeris hexasticha Brandt, 1833 alone (Kime and Enghoff 2011). Due to insufficient taxonomic information on the structure of the telopods and their uniformity 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 stable, 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 another 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 Wesener 2015; Wesener 2015a, 2015b, 2018; Wilbrandt et al. 2015; Wesener and Conrad 2016; Reip and Wesener 2018; Antić et al. 2021).

One of the taxa that have been forgotten and completely excluded from the European fauna is Glomeris herzogowinensis Verhoeff, 1898. Verhoeff (1898) described this taxon under the name “Glomeris europaea, herzogowinensis” on the basis of specimens from near Trebinje, Herzegovina, collected by Victor Apfelbeck, the then curator of the National Museum of Bosnia and Herzegovina in Sarajevo. Although the description of this taxon is relatively short, Verhoeff (1898) already points out in the first sentence: “...der marginata in der Färbung äusserst ähnlich...”, indicating a great similarity in colouration between G. herzogowinensis and one of the most common western-central-northern European species, Glomeris marginata (Villers, 1789). Albeit geographically completely separate, both species are characterised by mostly black, shiny tergites with yellowish or white posterior 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 posterior margins, evidently without examining the telopods of males from Greece. Later, it will turn out that, in these parts of Albania and Greece, there is or are one or even two species similar in appearance to G. herzogowinensis, but belonging to a different genus, Onychoglomeris Verhoeff, 1906. The fact that Verhoeff did not actually examine the telopods of the Greek male specimens is supported by the fact that, when establishing the then subgenus, Onychoglomeris, he included in it what we know today as Onychoglomeris tyrolensis (Latzel, 1884) and Simplomeris montivaga (Faës, 1902) (Verhoeff 1906). The telopods of these species differ markedly from those of the genus Glomeris and G. herzogowinensis. In his contribution to the knowledge of the genus Glomeris, Verhoeff (1911: 118, 119) included Glomeris herzogowinensis in the marginata species-group, stating some of the characteristics 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. Probably confused by Verhoeff’s (1901: 248, 249) earlier (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 genus Onychoglomeris from Albania and Greece he was studying, Attems (1935) just treated the species G. herzogowinensis as Onychoglomeris herzogowinensis. Attems (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 under G. herzogowinensis). Despite this error, Attems was, however, right in the fact that his new specimens belonged 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 (herzogowinensis 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 distributed further north on the other.

Six decades later, Mauriès et al. (1997), based on relatively abundant material of Onychoglomeris from Albania, but without studying the specimens from the type locality of G. herzogowinensis nor the Greek specimens of Onychoglomeris, questioned the existence of three subspecies, 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 almost nine decades and we consider both of Attems’ subspecies 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 containing 500 µl methylene chloride (DCM) for 5 minutes to extract their defensive secretions for future semiochemical studies. Later, the specimens were transferred to 70% ethanol.

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 für Naturkunde Berlin, Germany

ZSM Zoologische Staatssammlung München, Germany

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). Photographs 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 binocular 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 created using Google Earth Pro (version 7.3.6.9750) and Adobe 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 herzogowinensis, as well as Onychoglomeris australis stat. nov., a DNA barcoding analysis (Hebert et al. 2003) was conducted. 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 additionally Onychoglomeris ferraniensis Verhoeff, 1909 were analysed (see Table 1). In addition, sequences of similarily coloured (= black) Glomeris species were downloaded 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, 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 species, of which eight sequences from five species were newly sequenced.

Table 1.
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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 Locality Voucher # GenBank #
Glomeridella minima (Latzel, 1884) Austria ZSM MYR 00371 JN271878
Tonkinomeris huzhengkuni Liu & Golovatch, 2020 China SCAU TY01 MT522013
Glomeris pustulata Latreille, 1804 Germany ZSM MYR 00024 HM888093
Glomeris pustulata Latreille, 1804 Germany ZSM MYR 00376 JN271880
Glomeris hexasticha Brandt, 1833 Germany ZFMK MYR1460 MG931023
Glomeris hexasticha Brandt, 1833 Germany ZFMK MYR3898 MG931024
Glomeris tetrasticha Brandt, 1833 Germany ZSM MYR 00036 HM888105
Glomeris tetrasticha Brandt, 1833 Germany ZSM MYR 00035 HM888104
Glomeris marginata Villers, 1789 France ZFMK MYR6084 MG931022
Glomeris marginata Villers, 1789 Luxembourg ZFMK MYR1363 MG931021
Glomeris maerens Attems, 1927 Spain ZFMK MYR6097 MG892108
Glomeris maerens Attems, 1927 Spain ZFMK MYR6092 MG892110
Glomeris klugii Brandt, 1833 Italy ZFMK MYR637 KX714076
Glomeris klugii Brandt, 1833 Italy ZFMK MYR4734 KX714072
Glomeris apuana Verhoeff, 1911 Italy ZFMK MYR753 KT188944
Glomeris apuana Verhoeff, 1911 Italy ZFMK MYR752 KT188943
Onychoglomeris tyrolensis Latzel, 1884 Italy ZFMK MYR1276 KP205571
Glomeris pulchra Koch, 1847 Croatia, Dalmatia, Cetina River ZFMK MYR8217 PP475126
Glomeris pulchra Koch, 1847 Croatia, Dalmatia, Cetina River ZFMK MYR8217b PP475127
Glomeris balcanica Verhoeff, 1906 Greece, Dion-Olympos ZFMK MYR11331 PP475128
Onychoglomeris ferraniensis Verhoeff, 1909 Italy, Piemonte, Cuneo, Ceva ZFMK MYR623 PP475129
Onychoglomeris ferraniensis Verhoeff, 1909 Italy, Piemonte, Cuneo, Ormea ZFMK MYR2287 PP475130
Onychoglomeris australis Attems, 1935 stat. nov. Greece, Kalambaka ZFMK MYR11332 PP475131
Glomeris herzogowinensis Verhoeff, 1898 Bosnia & Herzegovina, Trebinje, Taleža ZFMK MYR8970 PP475132
Glomeris herzogowinensis Verhoeff, 1898 Bosnia & Herzegovina, Trebinje, Taleža ZFMK MYR8969 PP475133

The DNA extraction, amplification and sequencing protocol was similar to earlier studies (Wesener 2015a; Sagorny and Wesener 2017), using the degenerate (Astrin and Stüben 2008) primer pair HCO-JJ/LCO-JJ (HCOJJAWACTTCVGGRTGVCCAAARAATCA/LCOJJCHACWAAYCATAAAGATAT YGG). Sequences were concatenated by hand or by utilising the software Seqman (DNASTAR Inc.). BLAST searches (Altschul et al. 1997) were performed to confirm sequence identities. The whole dataset was translated into amino acids to rule out the accidental amplification 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. Codon positions included were 1st+2nd+3rd. All ambiguous positions were removed for each sequence pair. There were a total of 657 positions in the final dataset. Evolutionary analyses were conducted in Mega11 (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) (lnL = -4292.222, Invariant = 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 Reversible 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 algorithms to a matrix of pairwise distances estimated using the Maximum Composite Likelihood (MCL) approach. Codon positions included were 1st-2nd-3rd. All positions with less than 95% site coverage were eliminated, i.e. fewer than 5% alignment gaps, missing data and ambiguous 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 Illustrator 2023 with all bootstrap values > 50% illustrated (Fig. 1).

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%.

Results

Analysis of the COI barcoding gene

All species were recovered with high statistical support (94–100%, Fig. 1), while deeper nodes and interspecific relationships were statistically not supported. Neither the families Glomeridae and Glomeridellidae, nor the genus Glomeris are recovered as monophyletic (Fig. 1). Glomeris herzogowinensis does not group with Onychoglomeris species, but is in an unsupported sister-group with the similarily coloured G. maerens from the Mediterranean coast of Spain (Fig. 1). G. herzogowinensis and G. maerens show also the lowest genetic distance to one another (11.9–12.6%), while G. herzogowinensis also shows lower genetic distances 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%.

Taxonomy

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, 2, 3, 4, 5, 6, 7

Glomeris europaea, herzogowinensis Verhoeff, 1898: 163, fig. 18.

notGlomeris herzegowinensis (sic!).– Verhoeff (1901: 248).

notGlomeris 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).

notOnychoglomeris herzegowinensis (sic!).– Thaler (1999: 198, 199, figs 16, 17).

Glomeris marginata. – Ceuca (1990: 10).

Onychoglomeris herzogowinensis in part.– Kime and Enghoff (2011: 34, 118).

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-coloured anterolateral margins of the thoracic shield which is in the form of a narrow band in G. marginata. Additionally, 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.

Paralectotypes. ● 1 male, slide preparation (ZMB-MYR12772) (Verhoeff slide 953): leg pair 18 and telopods; Trebinje. ● 1 male, 1 female (ZMB-MYR2261); Trebinje.

?Types. ● 1 female (NHMW MY10415); Herzegovina: 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 Apfelbeck near Trebinje). ● 2 tubes (ZSM-A 20070848), 1 whole male, 1 male dissected (missing telopods and posterior leg pairs), 1 female, 1 juvenile; “ehemals Trockenmaterial” [material previously dry], Etk Nb. 28; Herzegowina. ● 1 tube (ZSM-A20070848), (Etk Nb. 28): 1 entire female specimen, a detached collum and thoracic shield, “ehemals Trockenmaterial; Tier m Original determinat. Etikett C Typüs-verdächtig” [material previously dry, animal with original determination, probable type], Trebinje. ● 1 male, slide preparation (ZSM-A20031802): telopods, leg pairs ?16, 17 and 18; Schuma (= Šuma, karst region around Trebinje).

Other material examined

Bosnia and Herzegovina: ● 1 female (NHMW MY10414); Trebinje ● 2 males, 1 female (IZB); in front of Taleža Cave, Taleža Village, near Trebinje, under stones; 15 November 2019; D. Antić leg. ● 3 males, 7 females (IZB); same locality as previous; 8 April 2022; D. Antić and D. Stojanović leg. ● 1 female (IZB); in front of Pavlova Cave, Bihovo Village, near Trebinje, under a stone; 16 November 2019; D. Antić leg. ● 1 female (IZB); same as previous but inside Pavlova Cave. Croatia: ● 1 female (NHMW MY10427); Pridvorje ● 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. Schönhofer leg. ● 1 male (ZFMK MYR95); Dubrovnik-Neretva, Konavle Region, Vignje, near Sklenica Cave, under stones in dense, humid, mossy forest, 89 m elev.; 3 April 2010; R. Ozimec & A. Schönhofer leg. ● 3 males (ZFMK MYR153); Dubrovnik-Neretva, Konavle region, Vignje, Špilja at Vignje Cave; under stones; 3 April 2010; R. Ozimec & A. Schönhofer leg. ● 1 female (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. Schönhofer leg. Montenegro: ● 1 male, 4 females (NHMW MY10413); Savina ● 1 male (IZB) ethanol and slide with leg pairs 17 and 18 and telopods; Orjen Mountain, Balješina Lokva, 1400 m elev.; 4 July 1997; I. Karaman leg. ● 1 female (ZFMK MYR220); Rumija 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. Schönhofer leg. ● 1 ex.; Borovik, near Cetinje; 11 May 2011; D. Antić 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 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 revealed 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.

Figure 2. 

Glomeris herzogowinensis Verhoeff, 1898, lectotype male (NHMW MY3903), habitus. A. Dorsal view; B. Lateral view; C. Ventral view. Scale bar: 1 mm.

Figure 3. 

Glomeris herzogowinensis Verhoeff, 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.

Figure 4. 

Glomeris herzogowinensis Verhoeff, 1898, male from Savina, Montenegro (NHMW MY10413). 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).

Glomeris herzogowinensis shows a striking resemblance 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 prominent light-coloured posterolateral margins compared to G. marginata. However, this is not entirely correct, as one individual analysed by us (Fig. 6A) has identical margins to most G. marginata. Indeed, most of the studied specimens of G. herzogowinensis have more pronounced margins than the classic G. marginata, but some French populations of the latter present posterolateral margins that are more developed than in G. herzogowinensis (see Reip and Wesener (2018: 96, fig. 1D, E)). Verhoeff (1898, 1911) mentioned the presence of strongly-pronounced light-coloured anterolateral margins 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. herzogowinensis has very distinct anterolateral margins (Figs 5A, C, 6B, D, E), in contrast to G. marginata, where it is only present in the form of a narrow band. The colouration of the fresh specimens that we have analysed corresponds completely to the description of Verhoeff (1898). They are mostly black with clearly demarcated lighter, whitish or yellowish posterolateral margins of the tergites. 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).

Figure 5. 

Glomeris herzogowinensis Verhoeff, 1898, male from Taleža, Bosnia and Herzegovina (IZB). A. Habitus, lateral view; B. Habitus, dorsal view; C. Head, collum and thoracic shield, anterior view; D. Anal shield, posterior view. Scale bars: 1 mm.

Figure 6. 

Glomeris herzogowinensis Verhoeff, 1898, male (A–C) and female (D–F) from Taleža, 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.

Figure 7. 

Glomeris herzogowinensis Verhoeff, 1898, living specimens. A, B. Female from Pavlova Cave, Bosnia and Herzegovina (IZB); C, D. Specimen from Taleža, Bosnia and Herzegovina (IZB). Photos by Dragan Antić.

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 telopods of G. herzogowinensis, as well as of the 17th pair of legs and the entire 18th pair of legs (Figs 3, 4B–D), which are of typical Glomeris appearance.

We would also like to mention that all examined specimens 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. herzogowinensis from G. marginata by the presence of two complete striae and an incomplete one (2+1 vs. 1+2, 1+1 or 1+0 sensu Schubart (1934: 33, fig. 28)). Indeed, all but two of the specimens examined show two complete striae and an incomplete one. In two specimens, the second stria is almost complete, with only a small interruption dorsally. Some specimens are characterised by the presence of additional, 4th incomplete striae in front of the first complete one.

Habitat

Known from almost near sea level up to 1400 m elev. in the Orjen Mountain. Scrubs of Carpinus, Quercus, Juniperus, under stones in limestone areas. Inside caves.

Distribution

The extreme south of Croatia and Bosnia and Herzegovina, as well as the coastal part of 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. Dražina pers. comm.; present study), Vignje (present study); Bosnia and Herzegovina: Surroundings of Trebinje (Verhoeff 1898; Attems 1929, 1935), Taleža near Trebinje (present study), Bihovo near Trebinje (present study). Montenegro: Savina (Attems 1929, 1935), Orjen (present study), Rumija, near Sutorman (present study), Cetinje, Borovik (present study).

Type locality

Near Trebinje, Bosnia and Herzegovina.

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).

Diagnosis

Similar in colouration (Fig. 12A–D) and morphology 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 telopoditomere 2 (= femur) and a shorter telopoditomere 4 (= tarsus), brownish stripes at the base of posteromesal process of telopoditomere 2 absent (vs. present in O. media stat. nov.), telopoditomere 3 (= tibia) with a well-developed posterior tooth that is more or less conical (vs. tooth poorly developed, subtriangular, sometimes almost absent 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 MY10424); 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 Platanoussa; 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. ● 5 males (NHMW MY10420); Epirus, Aoos Gorge near Konitsa, 550 m elev., Carpinus; 9 September 1996; K. Thaler and B. Knoflach leg. ● 8 males, 5 females (NHMW MY10421); Epirus, Timfi Mountain near Micropapingo, 800 m elev., bush; 10 September 1996; K. Thaler and B. Knoflach leg. ● 1 female (ZFMK MYR122); Epirus, Pindus Mountain, Zagori, Monodendri - Ano Pedina junction, under stone on the road, 835 m elev., 39.868002, 20.722076; 3 April 2006; A. Schönhofer leg. ● 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. Schönhofer 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. Huber & A. Schönhofer leg. ● 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. Schönhofer leg. ● 2 males, 4 females (ZFMK MYR124); Thessaly, 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. Schönhofer leg. ● 1 male (ZFMK MYR11332); Thessaly, Kalambaka, Meteora; September 2019; P. Knautt leg. ● 2 males, 1 juvenile (ZFMK MYR11334); same data ● 2 females (NHMW MY10417); Central Greece, Karpenisi; V. Apfelbeck leg. ● 1 female “?type”, (ZSM-A20070858), Epirus.

Remarks

Attems (1935), although he examined only a few males, already pointed out differences in the telopods between his australis and media, which we found to be constant after examining more males. The median lobe 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 structure is always the same, conical and well developed (Fig. 9A, B, E, white arrows). Such a structure is mentioned for O. media stat. nov. by Attems (1935) as much smaller compared to O. australis stat. nov. Our observation 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 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. media 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 beginnings of the second ridge (Fig. 8G) or that the second ridge is interrupted only in the centre (Fig. 8I).

Figure 8. 

Onychoglomeris australis Attems, 1935, stat. nov. A. Lectotype male (NHMW MY10424), habitus, lateral view; B–G. Males from Konitsa, Greece (NHMW MY10420). 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.

Figure 9. 

Onychoglomeris australis Attems, 1935, stat. nov. A. Male 4 from Konitsa, Greece (NHMW MY10420), 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 MY10420), leg pair 18, anterior view; H. Male 1 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.

We would like to emphasise that juveniles of this species 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 region (Fig. 12C, D).

The two southernmost finds of this species in Central Greece were apparently misidentified as G. herzogowinensis 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 identification 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, Carpinus, Quercus, Juniperus, Pinus, under stones, under tree trunks, under mossy limestone debris, leaf litter in limestone areas, open areas, bushland.

Distribution

Known from Epirus, Thessaly and central Greece (Fig. 13). Epirus: Paraskevi on Athamanika (Attems 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 Mikropapingo (Thaler 1999), Monodendri - Ano Pedina (present study), Zoodochos Pigi (present study). Thessaly: Kastanea, Elafi (present study), Panagia (present study), Kalambaka (present study). Central Greece: Karpenisi (Verhoeff 1901 [missidentification]; Attems 1935), Velouchi on Tymfristos (Verhoeff 1901 [missidentification]).

Type locality

Paraskevi, Epirus, Greece. Attems (1935: 143) stated: “Paraskevi ist ein Gipfel des Cumerka-Gebirges” which translates that Paraskevi is a summit on the Tzoumerka (= Athamanika) Mountain. We could not find out where exactly Paraskevi is located.

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.– Mauriès et al. (1997: 258–260, fig. 2).

Onychoglomeris herzegowinensis (sic!).– Ćurčić 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.– Sekulić and Živić (2017: 193). [Missidentification, but see Remarks below].

Diagnosis

Similar in colouration (Fig. 12E, F) and morphology 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 lateral 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 margin 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. australis stat. nov.), telopoditomere 3 (= tibia) with a poorly developed, sometimes almost absent, posterior tooth that is subtriangular (vs. tooth well developed, conical in O. australis stat. nov.), the syncoxite is mostly low, somewhat bilobed (vs. syncoxite usually higher and rounded in O. australis stat. nov.). For some more details, see Remarks below.

Material studied

Lectotype. 1 male (NHMW MY3901) in ethanol; Albania, Dukati [= Dukat]; 5 August 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. ● 2 females (NHMW MY10425); same data as for lectotype; ● 1 female (NHMW MY3902); Albania, Kanina [= Kaninë]; November 1908; A. Winneguth leg.

Other material examined

Albania: ● 1 male, 1 female (NHMW MY10412); Valona [= Vlorë]; Dr. K. Patsch leg.; ● 5 males, 2 females (IZB); Gjirocastro [= Gjirokastër]; 10 May, 1973; M. Karaman leg. ● 1 male, 2 females (ZFMK MYR13662); Gjirokastër District, Vjosa Valley, Përmet, Strëmbec, hiking trail Ri Soptit Waterfall; forest of low Carpinus, Quercus, Platanus and Crataegus, in leaf litter, 40.1488, 20.4543; 6 October 2023; H. Reip leg. Serbia: ● 1 male (IZB), slide with a male telopods and leg pairs 17 and 18; Visoki Dečani, Kosovo and Metohija; 1973; M. Karaman leg.

Remarks

As written above, one of the differences between O. media stat. nov. and O. australis stat. nov. is a much 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 Mauriès 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 observed in all males available to us (Fig. 11A, C, E, black arrows). Such stripes are absent from all males of O. australis 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 Mauriès et al. (1997: 259, fig. 2B, F)). All males at our disposal have a distinctly concave anal shield (Fig. 10A). In contrast to Mauriès 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).

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.

Figure 11. 

Onychoglomeris media Attems, 1935, stat. nov. A. Lectotype male (NHMW MY3901), telopods, anterior view; B. Lectotype male (NHMW MY3901), leg pair 18, anterior view; C. Male from Vlorë, Albania (NHMW MY10412), telopods, anterior view; D. Male from Vlorë, Albania (NHMW MY10412), telopods, posterior view; E. Male from Gjirokastër, Albania (IZB), part of the left telopod, anterior view; F. Male from Gjirokastër, Albania (IZB), part of the left telopod, posterior view; G. Male from Vlorë, 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.

It is of interest to mention a very isolated find in southern Serbia, near Visoki Dečani. This site is almost 250 km by air from the nearest site in the core area of southern Albanian sites (Fig. 13). In the IZB collection, only the microslide 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 in labelling or the species is really so widespread must be clarified in the future. The latter is supported by the fact that Sekulić and Živić (2017) recorded the occurrence of Glomeris marginata in southern Serbia (Znosek, Leposavić), about 80 km north-east of Visoki Dečani. It is obvious that this is not G. marginata, but it remains questionable which species Sekulić and Živić (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.

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 Përmet, Albania. Photos by Morris Fleck (A, B), Peter Kautt (C, D) and Hans Reip (E, F).

Figure 13. 

Distribution map of Glomeris herzogowinensis Verhoeff, 1898, Onychoglomeris australis Attems, 1935, stat. nov. and Onychoglomeris media Attems, 1935, stat. nov.

Habitat

There is no information about the habitat of this species in the literature, except that Mauriès 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.

Distribution

Southern Albanian species with a single, isolated locality in southern Serbia (Fig. 13). Albania: Dukat (Attems 1935; Mauriès et al. 1997), Kaninë (Attems 1935); Vlorë (Verhoeff 1901 [missidentification]; Attems 1929 [missidentification], 1935); Dhërmi (Mauriès et al. 1997), Himarë (Mauriès et al. 1997), Llogara Pass (Mauriès et al. 1997), Gjirokastër (Mauriès et al. 1997; present study), Përmet (present study). Serbia: Visoki Dečani (Ćurčić et al. 1999), ?Leposavić (Sekulić and Živić 2017 [missidentification]).

Type locality

Dukat, Vlorë County, southern Albania.

Discussion

Our DNA barcoding analysis clearly confirms the results of the morphological analysis of the telopods: Glomeris herzogowinensis groups with other Glomeris species and not with Onychoglomeris, while O. australis stat. nov. clearly groups with Onychoglomeris. Interestingly, the sister species to G. herzogowinensis seems to be G. maerens from Spain, a similarly-coloured species living in a similar Mediterranean habitat. However, there are indications that more than one species is currently hiding 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 studies 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 genus 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 (Wesener 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 timeless resource allowing us to study organisms and their systematics, sometimes even discover and describe completely unknown taxa, awaiting on shelves of museums to be determined, described and documented. The average 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 Pleonopurus tanzanicus Enghoff & Akkari, 2022 and reaching as high as 149 years such as Ommatoiulus schubarti Akkari & Enghoff, 2012 (Akkari and Enghoff 2012; Enghoff and Akkari 2022). In other cases, taxa have inadevertedly been mixed with other hitherto described species, therefore remaining hidden for decades. One of the latest examples is perhaps that of Lophostreptus neglectus Enghoff & 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 (Enghoff and Akkari 2024). The scientific collections, especially type series and historical specimens, are most defintely an invaluable source of information for taxonomists to update information, unravel the identity of obscure historical names (e.g. Akkari et al. (2010); Akkari (2013); Antić and Akkari (2020); Antić et al. (2021)), clarify the taxonomic status of taxa and solve complicated riddles like the one presented in this work. Morphology-based taxonomy remains a subjective exercise, especially when the studied groups did not traditionally have well-defined characters for species characterisation, which is the case for the order Glomerida. Taxonomy is also very prone to human error and this has been illustrated in numerous cases, especially 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. Amending 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.

In this article, we tried to solve the case of three species that have been hidden under the same name. Glomeris herzogowinensis was confirmed as an unquestionably good taxon. We have raised the other two taxa of the genus Onychoglomeris, former subspecies, to species level. Considering 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 Vjosa (in Albanian) or Aoos (in Greek) river valley in southern 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 Schönhofer, Siegfried Huber, Hans Reip, Peter Kautt (all from Germany), R. Ozimec (Croatia) and I. Karaman (Serbia) for kindly providing specimens. 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 barcoding 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 Stojanović (Serbia), Ivo Karaman (Serbia) and Marjan Komnenov (North Macedonia) for the wonderful time during the field trips in Bosnia and Herzegovina. DA’s field research was partly financed by the Serbian Ministry of Science, Technological Development and Innovation (grant no. 451-03-65/2024-03/ 200178). DA’s visit to the NHMW in December 2023 was funded by the Synthesys + project AT-TAF.

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Supplementary material

Supplementary material 1 

Number of base differences per site (p-distances) between sequences

Dragan Antić, Thomas Wesener, Nesrine Akkari

Data type: xls

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 users 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.
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