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Hidden diversity within the Nemastoma bidentatum Roewer, 1914 complex (Opiliones: Nemastomatidae) Part I: Morphological evidence

October 2021
European Journal of Taxonomy 777(1):1-67

DOI:10.5852/ejt.2021.777.1561

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CC BY 4.0

Authors:

Tone Novak

University of Maribor

Peter Kozel

Faculty of Natural Sciences and Mathematics, University of Maribor

Miriam Gudrun Schaider

University of Graz

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Nemastoma bidentatum Roewer, 1914 is a complex of closely related litter-dwelling harvestmen, characterized by a penis glans with two pairs of lateral lancet-like spines, and a femur IV with a saw-like series of pointed tubercles. Here we a) revise the hitherto known taxa within the N. bidentatum complex, and b) analyze in detail the relations among the taxa in Slovenia. The study revealed that the N. bidentatum complex consists of four species: N. bidentatum Roewer, 1914 s. str., N. relictum Gruber & Martens, 1968 stat. nov., N. pluridentatum (Hadži, 1973) stat. nov. and N. kozari Novak, Kozel, Podlesnik & Raspotnig sp. nov. Moreover, N. bidentatum s. str. consists of six subspecies: N. bidentatum bidentatum Roewer, 1914, N. bidentatum sparsum Gruber & Martens, 1968, N. bidentatum gruberi Novak, Slana Novak, Kozel & Raspotnig ssp. nov., N. bidentatum martensi Novak, Slana Novak & Raspotnig ssp. nov., N. bidentatum schmidti Novak, Raspotnig & Slana Novak ssp. nov. and N. bidentatum sneznikensis Novak, Komposch, Slana Novak & Raspotnig ssp. nov. In Slovenia, the six subspecies of N. bidentatum form a parapatric complex around N. bidentatum schmidti. Hybrids occur in the contact zones between adjacent subspecies, but they are missing between distant subspecies. The taxonomic distinction of lineages / subspecies is congruent with their distribution patterns. With six of the nine taxa present, Slovenia is considered the center of the N. bidentatum complex speciation. At the time being, this complex is the most diversified harvestman group on a subspecific and young-species level and provides important details on speciation processes in Opiliones.

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European Journal of Taxonomy 777: 1–67 ISSN 2118-9773

https://doi.org/10.5852/ejt.2021.777.1561 www.europeanjournaloftaxonomy.eu

2021 · Novak T. et al.

This work is licensed under a Creative Commons Attribution License (CC BY 4.0).

Monograph

urn:lsid:zoobank.org:pub:C86302D6-49F7-41A6-9BAC-94FDFED0D27B

Hidden diversity within the Nemastoma bidentatum Roewer, 1914

complex (Opiliones: Nemastomatidae)

Part I: Morphological evidence

Tone N OVA K SLA NA NOVA K KOZEL 

Miriam Gudrun SCHAIDER   

Jan    & Günther  





2

3

3



5



6

6



6



 Corresponding author: tone.novak@guest.um.si

2 Email: tone.novak@guest.arnes.si

3 Email: peter.kozel@um.si

4 Email: miriamschaider@hotmail.com

5 Email: c.komposch@oekoteam.at

6 Email: saska.lipovsek@um.si

7 Email: jan.podlesnik@um.si

8 Email: igor.pausic@um.si

9 Email: guenther.raspotnig@uni-graz.at

1 urn:lsid:zoobank.org:author:F6B51342-4296-473E-A3E8-AB87B0655F49

2 urn:lsid:zoobank.org:author:1603E85F-EE29-475A-8F98-B8B8E059C030

3 urn:lsid:zoobank.org:author:8C33627F-2D3D-4723-A059-B87462535460

4 urn:lsid:zoobank.org:author:8787CDCA-1F3E-48A2-8139-5CB73DE71733

5 urn:lsid:zoobank.org:author:CDE15756-4605-4C5C-AE6C-C576852C9092

6 urn:lsid:zoobank.org:author:1F7DB78E-CEE0-4840-95B4-8E4ADA3211DB

7 urn:lsid:zoobank.org:author:59572EBE-8871-4EAC-92F7-F87201CB6F67

8 urn:lsid:zoobank.org:author:35939273-4DBF-4CC4-AB06-FBDF1D17051B

9 urn:lsid:zoobank.org:author:75E9550E-01EF-439F-A3AB-6F463F35D03B



European Journal of Taxonomy 777: 1–67 (2021)

Abstract. Nemastoma bidentatum



like series of pointed tubercles. Here we a) revise the hitherto known taxa within the N. bidentatum



the N. bidentatum complex consists of four species: N. bidentatumN. relictum

 N. pluridentatumN. kozari 

N. bidentatum s. str. consists of six subspecies: N. bidentatum

bidentatumN. bidentatum sparsumN. bidentatum gruberi

N. bidentatum martensi 

N. bidentatum schmidti N. bidentatum

sneznikensis

N. bidentatum form a parapatric complex around N. bidentatum schmidti. Hybrids occur in the contact





N. bidentatum



important details on speciation processes in Opiliones.

Keywords.

                 

Raspotnig G. 2021. Hidden diversity within the Nemastoma bidentatum    

European Journal of Taxonomy 777: 1–67.

https://doi.org/10.5852/ejt.2021.777.1561

Introduction



amenable (Schönhofer 2013; Giribet & Sharma 2015; Fernández et al.









et al. 2006;

unpublished own data). One unresolved puzzle within Dyspnoi is the Nemastoma bidentatum



Nemastoma







et al.

et al. 2006; Schönhofer & Holle 2007; Komposch



N. bidentatum 

N. bidentatum from an unknown locality in





N. b. bidentatum



N OVA K T. et al., Nemastoma bidentatum (Opiliones: Nemastomatidae) revison

Fig. 1. Representatives of the Nemastoma bidentatumA. The habitus of a male

Nemastoma bidentatum gruberi 



B. The

body of a male Nemastoma relictum 





et al. 2016); Mt. Großes

C. The body of a male Nemastoma bidentatum gruberi 



European Journal of Taxonomy 777: 1–67 (2021)

subspecies: N. b. sparsumN. b. relictum



N. b. bidentatum and N. b. sparsum probably occurred. They (ibid.)

also reported N. b. bidentatum and N. b. sparsum

taxa belonging to the N. b. complex: Nemastoma (Lugubrostoma) triste pluridentatum

Nemastoma (Stridulostoma) seliskari Stridulostoma 

1973 with Nemastoma N. b. bidentatum.

Novak (2005a) substantiated N. ?bidentatum pluridentatum (the question mark denotes the unresolved



N. b. bidentatum and N. b. relictum do not interbreed

N. b. relictum should be raised to the species level if no hybrids were

N. b. bidentatum

and N. b. sparsumN. b. bidentatum

N. b. relictum and N. b. sparsum

2009). Hybrids between N. b. bidentatum and N. b. sparsum have been mentioned in Slovenia and Croatia

(Gruber & Martens 1968; Martens 1978).



et al. 2014). There is still much disagreement



et al. 2004; Haig et al.





et al. 2014).

Taxa described as subspecies must exhibit their own evolutionary trajectory and not be clinal variants

based on a single characteristic (Zink 2004; Oatley et al.



et al. 2006).



et al. 2006;

De Queiroz 2007; Miralles et al. 2010; Braby et al. 2012).



Mitostoma

olgaeNemastoma



1973a; Novak 2005a; Schönhofer 2013). These species are N. bidentatum N. bimaculatum

N. dentigerum  N. lugubre  N. schuelleri Gruber &

N. transsylvanicum N. triste 

subspecies ranked within Nemastoma bidentatum  N. b. bidentatum 

N. ?b. pluridentatum N. b. relictum  N. b. sparsum Gruber &









et al. 2014). Despite the reluctance

of many Dyspnoi to release scent gland substances (Raspotnig et al.

N OVA K T. et al., Nemastoma bidentatum (Opiliones: Nemastomatidae) revison



taxonomy (Raspotnig et al. 2017). In Paranemastoma quadripunctatumet al.

(2010) discovered an exclusively quinonic mixture of seven substances. The secretions of Nemastoma

tristeet al.

2018).

The taxonomic problems within the N. b. complex deserve renewed consideration of the taxa involved.





in Nemastoma. Most of the unresolved questions refer to populations in Slovenia where a number of

individuals of the N. b. complex cannot be assigned to any of the currently evidenced subspecies:

N. b. bidentatum and N. b. sparsum.



data from scent gland secretions were compiled for all species and subspecies of the genus Nemastoma

and deserve a separate publication. The aims of this contribution are as follows: a) to revise recent

knowledge on the N. b. complex and b) to analyze relations among the taxa of the complex on the territory

of Slovenia in greater detail.

Material and methods

Specimens examined and repositories



 





















we have retained the original citations.

N. b. bidentatumN. b. gruberi ssp. nov.:

N. b. martensiN. b. schmidti

N. b. sneznikensisN. b. sparsum

N. kozariN. pluridentatumN. b. bidentatum × N. b. schmidti

N. b. gruberi ssp. nov. × N. b. martensi

N. b. gruberi ssp. nov. × N. b. martensi ssp. nov. × N. b. schmidti N. b. gruberi

ssp. nov. × N. b. schmidti N. b. martensi ssp. nov. × N. b. schmidti ssp. nov.:

N. b. schmidti ssp. nov. × N. b. sneznikensisN. b. schmidti ssp. nov. ×

N. b. sparsum

Repositories

  

  

  

European Journal of Taxonomy 777: 1–67 (2021)

Abbreviations used

Apo = apophysis

  

  

  

  

  

  

  

  

  

  

Leg numbering: I–IV.

Sampling area in Slovenia

N. b. complex in Slovenia were too scattered to allow proper

analysis of variation within the N. b.N. bidentatum



N. bidentatum have not been found. In locality





were presented only exemplarily.

Morphology

 IV were considered in taxa

N. bidentatum as the basic taxonomic

Nemastoma bidentatum 





of denticles on Fe 













et al.











subterminally or terminally on the 















N OVA K T. et al., Nemastoma bidentatum (Opiliones: Nemastomatidae) revison

software. Transmission light microscopy was combined with illumination from above. Images were







(up to ca 70°C) in lactic acid until the ovipositor became translucent enough to examine the Rec sem.



Ecology

All the taxa within the N. b.





Field notes on the habitat and microhabitat characteristics inhabited by particular taxon of N. b. complex

were recorded to provide further ecological data. Data on ecology and habitat provide further information



Results











Genus Nemastoma

Diagnosis 

 





or denticles.

Nemastoma bidentatum

Diagnosis

Nemastoma bidentatum



Morphology







the glans is characteristic of some taxa. The shape and size of the Ch basal article and especially of Ch-







unknown female of N. pluridentatum 

Remarks

N. bidentatum complex have been discovered in Slovenia: N. b. gruberi

N. b. martensi 

N. b. schmidtiN. b. sneznikensis

European Journal of Taxonomy 777: 1–67 (2021)

N. kozari



N. b. gruberi

 

particular cases still raises doubts about the reliability of the assignment (see the description of the taxon

for more details). This is because we are still missing a broad review of potentially syntopic hybrids that







with N. b. schmidti









these taxa. Nemastoma bidentatum

Balkan countries do not allow any further conclusion for the time being.

Nemastoma b. complex in total: Nemastoma bidentatum

 N. relictumN. pluridentatum

N. kozari

sp. nov.

Nemastoma bidentatum 

Nemastoma bidentatum

includes N. triste].

Nemastoma bidentatumN. triste].

— Gruber & Martens 1968: 141–142 (redescription). — Martens 1978: 106–107. — Schönhofer

2013: 36.

Diagnosis 

Species of the Nemastoma b.





Remarks

N. b. bidentatumN. b. sparsum Gruber &

N. b. gruberi

N. b. martensiN. b. schmidti

Novak ssp. nov. and N. b. sneznikensis

Nemastoma bidentatum bidentatum 



includes N. triste].

Nemastoma bidentatumN. triste];

N. bidentatum sparsum].

N OVA K T. et al., Nemastoma bidentatum (Opiliones: Nemastomatidae) revison

Nemastoma bidentatum bidentatum

N. bidentatum schmidti 

N. bidentatum schmidti ssp. nov.]. — Komposch &

N. bidentatum schmidti ssp. nov.]. — Novak

et al. 2006: 269. — Schönhofer 2013: 36.

Diagnosis

Subspecies of N. bidentatum 



 

 

 Rec sem of 1 tubular and 12

elongated balloon-like vesicles.

Material examined

VM44

SLOVENIA – VM44





Description 

Male (from Ljubelj)











 min:max









  



 Leg article lengths in Tables 1–2.

Female (from Ljubelj)

 Length 1.6–1.8.







European Journal of Taxonomy 777: 1–67 (2021)

   







(Fig. 11B).

 Leg article lengths in Tables 1–2.

Remarks

N. b. sparsum

N. b. bidentatum and N. triste





remained unresolved 



N. triste

Martens (1968) redescribed the nominal subspecies N. b. bidentatum upon a neotype (Gruber & Martens





 

N. b. schmidti ssp. nov. All so far published evidence on N. b. bidentatum



et al.



(Gruber & Martens 1968) also includes N. b. schmidti ssp. nov. See remarks under N. pluridentatum

N. b. schmidtiN. b. gruberi ssp. nov. × N. b. schmidti ssp. nov. and N. b. martensi

ssp. nov. × N. b. schmidti ssp. nov.

Trochanter Femur Patella Tibia Metatarsus Tarsus Total length

Pedipalp 0.25 (0.33) 0.76 (0.80) 0.53 (0.56) 0.46 (0.49) – 0.29 (0.31) 2.29 (2.49)

Leg I 0.23 (0.24) 0.68 (0.70) 0.39 (0.38) 0.61 (0.64) 1.06 (1.05) 0.97 (0.98) 3.94 (3.99)

Leg II 0.27 (0.31) 1.30 (1.56) 0.47 (0.45) 1.13 (1.01) 2.03 (1.68) 0.84 (1.47) 5.98 (6.48)

Leg III 0.19 (0.23) 0.87 (0.96) 0.36 (0.35) 0.61 (0.64) 1.15 (1.17) 0.88 (0.91) 4.06 (4.26)

Leg IV 0.25 (0.25) 1.32 (1.21) 0.39 (0.44) 0.78 (0.89) 1.84 (1.84) 1.18 (1.16) 5.75 (5.79)

Table 1. Length of appendage-segments in millimeters in N. bidentatum bidentatum



Femur Patella Tibia Metatarsus Tarsus Total length

Leg I 0.75 (0.75) 0.35 (0.35) 0.55 (0.60) 1.00 (0.90) 0.85 (0.85) 3.50 (3.50)

Leg II 1.20 (0.80) 0.35 (0.40) 1.05 (1.05) 1.90 (1.95) 1.55 (1.60) 6.05 (5.80)

Leg III 0.75 (0.85) 0.30 (0.35) 0.60 (0.55) 1.15 (1.05) 0.80 (0.80) 3.60 (3.60)

Leg IV 1.25 (1.30) 0.35 (0.35) 0.75 (0.75) 1.75 (1.70) 1.05 (1.05) 5.15 (5.15)

Table 2. Length of appendage-segments in millimeters (trochanters not presented) in N. bidentatum

bidentatum

N OVA K T. et al., Nemastoma bidentatum (Opiliones: Nemastomatidae) revison

Distribution









Ecology

Montane subspecies of N. bidentatum

Abies alba

of Picea abies (L.) H.Karst. and Fagus sylvatica

Nemastoma bidentatum sparsum 



Nemastoma bidentatum sparsum

N. bidentatum gruberi ssp. nov.].

Nemastoma bidentatumN. bidentatum bidentatum

and N. triste

 — Martens 1978:



286. — Novak et al.

Fig. 2. The complete distributional area of the Nemastoma bidentatum



positions of the type localities of the taxa.

European Journal of Taxonomy 777: 1–67 (2021)





Diagnosis 



oblong  

 Apo conical. Glans equilaterally triangular. Rec

sem with 14 balloon-like vesicles.

Fig. 3.Nemastoma

bidentatum

N OVA K T. et al., Nemastoma bidentatum (Opiliones: Nemastomatidae) revison

Material examined

CROATIA – XM05



SLOVENIA – WM06

– WM42





WM45



WM47

WM52

WM53

WM56





WM57



WM66







WM67







WM73WM74 •





WM75











WM77





WM78

WM86











WM87





WM88

WM89

European Journal of Taxonomy 777: 1–67 (2021)

WM94

WM95

WM96 

WM97







WM98





WM99 •



– XM05

XM06









XM07



XM08









XM15















       





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



XM16







XM25



Description 

Male 



1.44.

N OVA K T. et al., Nemastoma bidentatum (Opiliones: Nemastomatidae) revison

 Ch basal article with nearly straight dorsal margin in front of dorsal indentation and small

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



 

lengths in Table 3.

.   

. Leg article lengths in Tables 3–4.

Female 





 





 





. Leg article lengths in Tables 3–4.

Remarks

N. b. sparsum



N. b. gruberi ssp. nov. distributed south of the Sava River

N. b. sparsum south of the Sava River (Novak et al.

N. b. gruberi ssp. nov. Martens (1978: 112) reported a sparsum-like

population living in the mountain pine association (Hyperico grisebachii-Pinetum mugo Horvat 1938)



may refer to N. b. gruberiN. b. martensi ssp. nov. or N. b. sneznikensis ssp. nov. Individuals



N. b. gruberi ssp. nov. and N. b. gruberi ssp. nov. ×

N. b. schmidtiN. b. martensi ssp. nov. and N. b. sneznikensis ssp. nov.

Distribution













European Journal of Taxonomy 777: 1–67 (2021)

Ecology

Nemastoma b. sparsum is a lowland and hilly land subspecies inhabiting various types of acidophilous

and neutrophilous forests of Fagus sylvaticaCarpinus betulus L. and Quercus petraea





Nemastoma relictum 



Nemastoma bidentatum relictum

Nemastoma bidentatum relictum 

485. — Komposch 2009: 476; 2011: 65. — Schönhofer 2013: 36.

Diagnosis

Species within the N. b. complex with button-shaped 



saccular vesicles.

Material examined





Description 

Male 



 Ch 



Trochanter Femur Patella Tibia Metatarsus Tarsus Total length

Pedipalp 0.25 (0.28) 0.75 (0.74) 0.52 (0.60) 0.45 (0.47) – 0.29 (0.28) 2.26 (2.37)

Leg I 0.23 (0.24) 0.68 (0.70) 0.39 (0.38) 0.61 (0.64) 1.06 (1.05) 0.97 (0.98) 3.94 (3.99)

Leg II 0.27 (0.31) 1.30 (1.56) 0.47 (0.45) 1.13 (1.01) 2.03 (1.68) 0.84 (1.47) 5.98 (6.48)

Leg III 0.19 (0.23) 0.87 (0.96) 0.36 (0.35) 0.61 (0.64) 1.15 (1.17) 0.88 (0.91) 4.06 (4.26)

Leg IV 0.25 (0.25) 1.32 (1.21) 0.39 (0.44) 0.78 (0.89) 1.84 (1.84) 1.18 (1.16) 5.75 (5.79)

Table 3. Length of appendage-segments in millimeters in N. bidentatum sparsum



Femur Patella Tibia Metatarsus Tarsus Total length

Leg I 0.75 (0.75) 0.30 (0.30) 0.55 (0.55) 0.95 (0.95) 0.80 (0.90) 3.35 (3.45)

Leg II 1.30 (1.25) 0.35 (0.40) 1.00 (1.00) 1.95 (1.95) 1.45 (1.45) 6.05 (6.05)

Leg III 0.75 (0.75) 0.30 (0.30) 0.50 (0.50) 1.05 (1.10) 0.60 (0.75) 3.20 (3.40)

Leg IV 1.20 (1.20) 0.35 (0.30) 0.75 (0.75) 1.70 (1.70) 1.05 (1.00) 5.05 (4.95)

Table 4. Length of appendage-segments in millimeters (trochanters not presented) in N. bidentatum

sparsum

1968).

N OVA K T. et al., Nemastoma bidentatum (Opiliones: Nemastomatidae) revison





 





   

 Leg article lengths in Tables 5–6.

Female 

 Length 1.70.

 







  





 Leg article lengths in Tables 5–6.

Remarks



N. b. complex.

Trochanter Femur Patella Tibia Metatarsus Tarsus Total length

Pedipalp 0.25 (0.25) 0.65 (0.63) 0.51 (0.47) 0.41 (0.37) – 0.32 (0.25) 2.14 (1.97)

Leg I 0.21 (0.23) 0.78 (0.74) 0.34 (0.40) 0.63 (0.52) 0.90 (0.90) 0.89 (0.76) 3.75 (3.55)

Leg II 0.20 (0.23) 1.23 (1.23) 0.42 (0.39) 0.98 (0.98) 1.82 (1.62) 1.21 (1.35) 5.86 (5.80)

Leg III 0.20 (0.20) 0.78 (0.56) 0.31 (0.31) 0.56 (0.54) 1.07 (1.00) 0.67 (0.80) 3.59 (3.41)

Leg IV 0.24 (0.20) 1.13 (1.15) 0.39 (0.38) 0.83 (0.77) 1.56 (1.45) 1.12 (1.04) 5.27 (4.99)

Table 5. Length of appendage-segments in millimeters in N. relictum



Femur Patella Tibia Metatarsus Tarsus Total length

Leg I 0.65 (0.75) 0.30 (0.30) 0.50 (0.50) 0.85 (0.90) 0.80 (0.85) 3.10 (3.30)

Leg II 1.10 (1.15) 0.40 (0.40) 0.95 (0.90) 1.70 (1.65) 1.30 (1.30) 5.45 (5.40)

Leg III 0.70 (0.75) 0.30 (0.30) 0.50 (0.55) 1.00 (1.00) 0.75 (0.80) 3.25 (3.40)

Leg IV 1.10 (1.15) 0.35 (0.30) 0.75 (0.75) 1.50 (1.50) 1.00 (1.00) 4.70 (4.70)

Table 6. Length of appendage-segments in millimeters (trochanters not presented) in N. relictum Gruber &

 

Gruber & Martens 1968).

European Journal of Taxonomy 777: 1–67 (2021)

Distribution









Ecology

Nemastoma relictum stat. nov. is a montane to alpine species within the N. b. complex inhabiting forests







probably eurychronous.

Nemastoma pluridentatum  stat. nov. Novak



Nemastoma (Lugubrostoma) triste pluridentatum

the description also includes N. bidentatum schmidti N. bidentatum gruberi ssp. nov. ×

N. bidentatum schmidti ssp. nov. and N. bidentatum martensi ssp. nov. × N. bidentatum schmidti

ssp. nov.].

Nemastoma bidentatum bidentatum – Martens 1978: 107.

Nemastoma ?bidentatum pluridentatum – 

Nemastoma bidentatum pluridentatum – Schönhofer 2013: 36.

Diagnosis 

Nemastoma b.





Material examined

Holotype

BOSNIA AND HERZEGOVINA – YK10



was erroneously cited as YJ19.)

Partial redescription

Note: male body and penis missing; female unknown.

Male (holotype)











lengths in Table 7.

N OVA K T. et al., Nemastoma bidentatum (Opiliones: Nemastomatidae) revison



lengths in Table 7.

Remarks

Nemastoma (Lugubrostoma) triste pluridentatum

N. pluridentatum

N. b. gruberi ssp. nov. × N. b. schmidti ssp. nov. 

N. b. martensi ssp. nov. × N. b. schmidti



 Nemastoma b. schmidti 







original microscopic preparation labeled “Nemastoma bidentatum



Nemastoma (Stridulostoma) seliskari 

under N. b. martensi ssp. nov. × N. b. schmidti





N. pluridentatum

a mistyped name: “N. (L.) triste pleuridentatum



the type locality of N. pluridentatum







 IV in preserved legs is the only available character allowing us to place this

taxon into the N. b.N. ?bidentatum pluridentatum

N. b. pluridentatum

N. pluridentatumN. kozari sp. nov.

N. pluridentatum

status to N. pluridentatum. See remarks under N. relictumN. kozari sp. nov. and hybrids

N. b. gruberi ssp. nov. × N. b. schmidti ssp. nov. and N. b. martensi ssp. nov. × N. b. schmidti ssp. nov.

Distribution





Trochanter Femur Patella Tibia Metatarsus Tarsus Total length

Pedipalp 0.27 0.68 0.49 0.38 – 0.29 2.11

Leg I 0.19 0.60 0.31 0.41 0.50 0.49 2.50

Leg II 0.22 1.05 0.28 1.25 1.05 0.80 4.65

Leg III 0.21 0.70 0.25 0.50 0.65 0.65 3.16

Leg IV 0.22 0.95 0.25   0.75 3.95

Table 7. Length of appendage-segments in millimeters in N. pluridentatum stat. nov. from



European Journal of Taxonomy 777: 1–67 (2021)

Ecology

Nemastoma pluridentatum stat. nov. inhabit the highest peak areas with the Mastgrass Gentianello

crispatae-Nardetum strictae 

Nemastoma bidentatum gruberi 

urn:lsid:zoobank.org:act:17EA28A1-5F36-4DFC-9191-9C8680496FAC



Nemastoma bidentatum sparsum – Novak et al.





Diagnosis

Relatively long-legged subspecies of Nemastoma bidentatum





one tubular and 12 elongated balloon-like vesicles.

Etymology

gruberi 

who generously introduced us to the taxonomy of harvestmen and helped in identifying problematic

specimens.

Material examined

Holotype

SLOVENIA – WL18



Paratypes

SLOVENIA – WL18





Other material

ITALY – VL05

VL15



CROATIA – UL92 N. b. sparsum

VL01

VL13

XL90



SLOVENIA – UL93









N OVA K T. et al., Nemastoma bidentatum (Opiliones: Nemastomatidae) revison



























VL03

VL04











VL07

VL13



VL14







– VL15VL16

VL17

VL18

VL23

VL25



VL28



VL29

VL36

VL45

VL47

VL54











VL36

VL66

VL67





European Journal of Taxonomy 777: 1–67 (2021)



VL69

VL74





VL75



VL76









VL77





VL78



VL79

VL83

VL84



VL85







VL86



VL87

VL89VL93

VL94 •











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VL97

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WL03

N OVA K T. et al., Nemastoma bidentatum (Opiliones: Nemastomatidae) revison

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WL06



WL07

WL08





WL13











WL14











WL15















WL16



WL17









WL18

WL19



WL23



European Journal of Taxonomy 777: 1–67 (2021)

WL24





WL25

















WL26



WL27











WL28





WL29





WL37





WL38









 WL39













WL48 •





WL49

WL59





WM01

WM10

N OVA K T. et al., Nemastoma bidentatum (Opiliones: Nemastomatidae) revison



WM20



WM30







WM40



















WM41



– WM50







Description

Male (holotype)



 











 



 

 

  

 Leg

article lengths in Table 8.

Female



European Journal of Taxonomy 777: 1–67 (2021)









 

 Ovipositor 

(Fig. 11E).

 Leg article lengths in Table 8.

Variability







N. b. bidentatum. Further

investigation in Croatian Istria is required to elucidate the whole range of morphological variation and



Remarks

Nemastoma b. gruberi ssp. nov. is the most variable subspecies within N. bidentatum

N. b. sparsum

N. b. sparsum

et al.

N. b. sparsum



N. b. gruberi



Distribution





Ecology

Nemastoma b. gruberi 

shrub and Quercetalia pubescenti–petraeae 



Trochanter Femur Patella Tibia Metatarsus Tarsus Total length

Pedipalp 0.28 (0.31) 0.77 (0.81) 0.56 (0.63) 0.47 (0.49) – 0.30 (0.31) 2.38 (2.55)

Leg I 0.26 (0.27) 0.94 (0.83) 0.39 (0.35) 0.67 (0.63) 1.11 (1.04) 1.10 (0.98) 4.47 (4.10)

Leg II 0.23 (0.23) 1.55 (1.40) 0.47 (0.50) 0.91 (1.16) 2.21 (2.24) 1.80 (1.61) 7.17 (7.14)

Leg III 0.22 (0.22) 0.82 (0.83) 0.36 (0.35) 0.63 (0.63) 1.22 (1.19) 0.97 (0.95) 4.22 (4.17)

Leg IV 0.23 (0.23) 1.43 (1.35) 0.39 (0.41) 0.79 (0.91) 1.94 (1.90) 1.45 (1.23) 6.23 (6.03)

Table 8. Length of appendage-segments in millimeters in N. bidentatum gruberi 



N OVA K T. et al., Nemastoma bidentatum (Opiliones: Nemastomatidae) revison

Nemastoma kozari 

urn:lsid:zoobank.org:act:7F207FF7-9241-4854-A4C3-B7DED507F740



Diagnosis

Species of the N. b.



Etymology

kozari 



Material examined

Holotype

BOSNIA AND HERZEGOVINA – YK10





Paratypes

BOSNIA AND HERZEGOVINA – YK10

Opiliones-TN 137/2019.

Other material

BOSNIA AND HERZEGOVINA – YK10



Description

Male (holotype)













  height:length ~

 

 

Table 9.

  



lengths in Table 9.

Female



European Journal of Taxonomy 777: 1–67 (2021)









and  



(Fig. 11F).

 Leg lengths in Table 9.

Distribution







Ecology

Nemastoma kozari 



Nemastoma bidentatum martensi 

urn:lsid:zoobank.org:act:CCDC54EF-D41A-4547-A4D4-50370212C243



Diagnosis

Subspecies of Nemastoma bidentatum



Rec sem of 12–14 slightly elongated balloon-like vesicles.

Etymology

martensi 



Material examined

Holotype

SLOVENIA – VL23 



Trochanter Femur Patella Tibia Metatarsus Tarsus Total length

Pedipalp 0.25 (0.28) 0.74 (0.69) 0.51 (0.55) 0.41 (0.43) – 0.28 (0.29) 2.19 (2.24)

Leg I 0.22 (0.23) 0.87 (0.76) 0.35 (0.30) 0.59 (0.62) 0.97 (0.91) 0.82 (0.88) 3.82 (3.70)

Leg II 0.24 (0.24) 1.32 (1.22) 0.44 (0.36) 1.08 (1.00) 1.87 (1.72) 1.46 (1.37) 6.41 (5.91)

Leg III 0.23 (0.26) 0.82 (0.78) 0.33 (0.31) 0.53 (0.51) 1.09 (1.06) 0.82 (0.72) 3.82 (3.64)

Leg IV 0.27 (0.26) 1.28 (1.23) 0.43 (0.38) 0.88 (0.77) 1.74 (1.69) 1.04 (0.79) 5.64 (5.12)

Table 9. Length of appendage-segments in millimeters in N. kozari



N OVA K T. et al., Nemastoma bidentatum (Opiliones: Nemastomatidae) revison

Paratypes

SLOVENIA – VL23

Other material

SLOVENIA – VL15



VL23











VL24













VL25

VL26

VL33

VL34

VL43



VL44



– VL53

– VL54





VL84

VL87



Description

Male (holotype)















  height:length ~



  

lengths in Table 10.

European Journal of Taxonomy 777: 1–67 (2021)

  

 Leg

article lengths in Table 10.

Female











  



 Leg article lengths in Table 10.

Distribution





Ecology

Nemastoma b. martensi ssp. nov. is native to lowland to submontane habitats with thermophile scrub and

Quercetalia pubescenti–petraeae and Tilia platyphyllos

admixture of Fagus sylvatica and Corylus avellana

probably eurychronous.

Nemastoma bidentatum schmidti 

urn:lsid:zoobank.org:act:389E4C27-24B9-44F9-A724-11475CB9E12B



Nemastoma triste



Nemastoma bidentatum       

Zagreb / Agram; the other localities refer to N. bidentatum sparsum



Trochanter Femur Patella Tibia Metatarsus Tarsus Total length

Pedipalp 0.27 (0.26) 0.73 (0.76) 0.48 (0.55) 0.44 (0.47) – 0.27 (0.29) 2.19 (2.33)

Leg I 0.23 (0.23) 0.93 (0.80) 0.35 (0.35) 0.60 (0.56) 1.03 (0.94) 0.78 (0.89) 3.92 (3.88)

Leg II 0.22 (0.23) 1.43 (1.35) 0.48 (0.45) 1.15 (1.25) 1.93 (1.80) 1.69 (1.65) 6.88 (6.63)

Leg III 0.24 (0.21) 0.83 (0.80) 0.33 (0.33) 0.60 (0.59) 1.09 (1.05) 0.83 (0.83) 3.93 (3.81)

Leg IV 0.23 (0.21) 1.33 (1.30) 0.48 (0.38) 0.83 (0.83) 1.59 (1.63) 0.84 (0.83) 5.30 (5.18)

Table 10. Length of appendage-segments in millimeters in N. bidentatum martensi 



N OVA K T. et al., Nemastoma bidentatum (Opiliones: Nemastomatidae) revison

Nemastoma bidentatum bidentatum





Nemastoma quadripunctatum humerale 

Nemastoma (Lugubrostoma) triste pluridentatum



?Nemastoma bidentatum bidentatum × N. bidentatum sparsum – Novak & Gruber 2000: 285.

Diagnosis

Subspecies of Nemastoma bidentatum with elongated oval-trapezoid 





  Rec sem

of one tubular and 12 slightly elongated vesicles.

Etymology

schmidti 





Material examined

Holotype

SLOVENIA – VM31 



Paratypes

SLOVENIA – VM31 

Other material

VM74





CROATIA – WL67

ITALY – UM52

UM62







UM65

UM80



N. triste

N. triste

UL99





UM72

European Journal of Taxonomy 777: 1–67 (2021)

UM73

UL82



UM90







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VL16VL19



VL26 •

N OVA K T. et al., Nemastoma bidentatum (Opiliones: Nemastomatidae) revison

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



VL36







VL37



VL38



VL39

VL46

VL47







VL48









VL49



VL54



VL56



VL57

VL58



VL59







VL68













European Journal of Taxonomy 777: 1–67 (2021)





VL69



VL79

VL85

VL89

VL93

VL99

VM00







VM05

VM09

– VM10



VM12

VM13

VM14

VM20



VM21 •





VM22

VM23

VM30

VM31

                

VM32

VM33



VM40

VM41 •

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VM42

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

VM43

VM50 •

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



VM51

N OVA K T. et al., Nemastoma bidentatum (Opiliones: Nemastomatidae) revison

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

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







European Journal of Taxonomy 777: 1–67 (2021)

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

VM60



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

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

VM70







VM71





VM72

VM73

– VM74VM80



VM81





VM82



VM83VM84

VM85





VM90

N OVA K T. et al., Nemastoma bidentatum (Opiliones: Nemastomatidae) revison





VM91







VM93



– VM94

VM95



VM96



WL07

WL19

WL39

WL48

WM00



























WM02



– WM03

– WM04



WM06













WM13





WM14 •

WM15





European Journal of Taxonomy 777: 1–67 (2021)

– WM16





























WM23

WM32

WM33

WM34



WM35

WM36

WM43

WM45







– WM46











WM54

WM55

WM64



WM65



WM74



WM75







WM76

WM95



N OVA K T. et al., Nemastoma bidentatum (Opiliones: Nemastomatidae) revison

Description

Male (holotype)











 

 





  



article lengths in Table 11.

Female















 Leg article lengths in Table 11.

Remarks

N. bidentatumN. b. schmidti

N. bidentatum or N. b. bidentatum in

N. b. schmidti ssp. nov.

 N. b. bidentatum



1978; Novak et al.



as N. bidentatum  

N. b. schmidti ssp. nov. a possible hybrid N. b. bidentatum × N. b. sparsum. The glans tip of a specimen



 N. b. schmidti ssp. nov. See remarks

under N. pluridentatum N. b. gruberi ssp. nov. × N. b. schmidti ssp. nov. and N. b. martensi

ssp. nov. × N. b. schmidti ssp. nov.

Distribution





European Journal of Taxonomy 777: 1–67 (2021)

Ecology

Nemastoma b. schmidti ssp. nov. is a mesophilous colline to montane subspecies occupying most regions



from mixed hardwood riverine forests in the lowland to subalpine Nardion strictae Br.-Bl. 1926 with

isolated patches of Pinus mugo

eurychronous.

Nemastoma bidentatum sneznikensis 

urn:lsid:zoobank.org:act:7690FA07-5D60-4545-8660-282ADED3F576



Etymology

sneznikensis 

Diagnosis

N. b.



vesicles.

Material examined

Holotype

SLOVENIA – VL54

Novak and Tone Novak leg.; deep-humus in Dinaric Pinus mugo scrub in contact with the Dinaric Fagus

sylvatica–Abies alba

Paratypes

SLOVENIA – VL54

Other material

SLOVENIA – VL54



Description

Male (holotype)











Trochanter Femur Patella Tibia Metatarsus Tarsus Total length

Pedipalp 0.28 (0.30) 0.77 (0.83) 0.45 (0.60) 0.46 (0.48) – 0.26 (0.33) 2.22 (2.54)

Leg I 0.21 (0.25) 0.86 (0.85) 0.33 (0.35) 0.63 (0.60) 0.93 (1.01) 0.95 (0.99) 3.91 (4.05)

Leg II 0.23 (0.29) 1.35 (1.35) 0.41 (0.49) 1.12 (1.08) 1.98 (1.95) 1.58 (1.63) 6.67 (6.79)

Leg III 0.22 (0.18) 0.86 (0.85) 0.35 (0.35) 0.59 (0.60) 1.08 (1.04) 0.87 (0.78) 3.97 (3.80)

Leg IV 0.22 (0.26) 1.32 (1.33) 0.41 (0.40) 0.82 (0.81) 1.60 (1.81) 1.11 (1.08) 5.48 (5.69)

Table 11. Length of appendage-segments in millimeters in N. bidentatum schmidti 



N OVA K T. et al., Nemastoma bidentatum (Opiliones: Nemastomatidae) revison

  min:max





 



lengths in Table 12.

Female















 Leg article lengths in Table 12.

Remarks

N. b. sneznikensis ssp. nov. is very similar to N. relictum



found in the last decade. See remarks on taxa of NemastomaN. b. gruberi ssp. nov.

Distribution





Ecology

Nemastoma b. sneznikensis ssp. nov. is a psychrophilous subspecies. To date it has been recorded only

 Dinaric Pinus mugo scrub in contact with the Dinaric Fagus

sylvatica–Abies alba treeline on calcareous soils. 

N. relictum (Komposch 1999;



et al. 2016; Halse 2018). Since this habitat is restricted to an

area of less than 1 km



probably eurychronous.

Trochanter Femur Patella Tibia Metatarsus Tarsus Total length

Pedipalp 0.20 (0.15) 0.59 (0.58) 0.43 (0.48) 0.33 (0.38) – 0.24 (0.25) 1.79 (1.84)

Leg I 0.19 (0.18) 0.73 (0.62) 0.31 (0.28) 0.52 (0.51) 0.79 (0.96) 0.74 (0.63) 3.28 (3.18)

Leg II 0.19 (0.23) 1.16 (0.95) 0.38 (0.38) 1.03 (0.93) 1.46 (1.61) 1.27 (1.24) 5.49 (5.34)

Leg III 0.18 (0.17) 0.70 (0.78) 0.29 (0.27) 0.53 (0.52) 0.86 (1.04) 0.63 (0.57) 3.19 (3.35)

Leg IV 0.17 (0.18) 1.06 (0.98) 0.33 (0.28) 0.74 (0.73) 1.42 (1.44) 1.01 (0.92) 4.73 (4.53)

Table 12. Length of appendage-segments in millimeters in Nemastoma bidentatum sneznikensis ssp. nov.



European Journal of Taxonomy 777: 1–67 (2021)

Hybrids among the subspecies of Nemastoma bidentatum

Remarks

N. b. bidentatum lives in syntopy with N. relictum (Gruber &

Martens 1968; Martens 1978; in both papers sub N. b. relictumN. b. sparsum (Komposch &



considered as potential species (Gruber & Martens 1968; Martens 1978; Komposch & Gruber 2004;

N. b. schmidti  interbreeds with

all other adjacent subspecies: N. b. bidentatumN. b. martensiN. b. sneznikensis

N. b. gruberi ssp. nov. and N. b. sparsum from the Southwest to the Northeast of the country.

N. b. gruberi ssp. nov. interbreed with N. b. martensiN. b. sparsum

N. pluridentatum

N. b. complex allow it to be

considered a separate species. This is because interbreeding of N. pluridentatum 

N. kozari

Nemastoma in Bosnia and Herzegovina are nearly



 than pure subspecies and are well





of either one or two pure subspecies and a series of hybrids with varying Ch-Apo shapes.

An overview of interbreeding relations among the taxa of the N. b. complex is presented in Table 13.

N. b. bidentatum and N. b. sparsum



N. b. gruberi ssp. nov. × N. b. schmidti ssp. nov. × N. b. sparsum

N. b. sneznikensis

ssp. nov. are too scarce to be properly evaluated in full. The zone of hybridization between N. b. schmidti

ssp. nov. and N. b. sparsum



Hybrids N. b. gruberi ssp. nov. × N. b. schmidti

N. b. martensi ssp. nov. × N. b. schmidti ssp. nov. and N. b. gruberi ssp. nov. × N. b. martensi ssp. nov. are

N. b. martensi

ssp. nov. and N. b. sparsum











Nemastoma bidentatum bidentatum  × N. bidentatum schmidti ssp. nov.



Diagnosis





Material examined

SLOVENIA – VM44

WM06

N OVA K T. et al., Nemastoma bidentatum (Opiliones: Nemastomatidae) revison













Nemastoma bidentatum bidentatum N. bidentatum sparsum 

Remarks

Every allegation of hybrids N. b. bidentatum × N. b. sparsum in Slovenia and Croatia (Gruber & Martens

1968; Martens 1978; Novak & Gruber 2000; Novak et al. 2002; Novak 2004b) refer either to N. b. schmidti

ssp. nov. × N. b. sparsum or N. b. gruberi ssp. nov. × N. b. schmidti ssp. nov.; see these references for

further remarks.

Nemastoma bidentatum gruberi ssp. nov. × N. bidentatum martensi ssp. nov.



Diagnosis







Material examined

CROATIA – WL00WL20 •





SLOVENIA – VL26

VL33

VL54

bidentatum

gruberi

kozari

martensi

pluriden-

tatum

relictum

schmidti

sneznikensis

gruberi

kozari

martensi +

pluridentatum

relictum

schmidti + + +

sneznikensis +

sparsum + +

Table 13.N. bidentatum complex.

European Journal of Taxonomy 777: 1–67 (2021)





VL95

VL96



WL37



Nemastoma bidentatum gruberi ssp. nov. × N. bidentatum martensi ssp. nov.

× N. bidentatum schmidti ssp. nov.



Diagnosis

Typical male hybrids are characterized by Ch-Apo with nearly symetrically convex ventro-anterior margin





Material examined

SLOVENIA – VL09

VL26



Nemastoma bidentatum gruberi ssp. nov. × N. bidentatum schmidti ssp. nov.



Nemastoma bidentatum bidentatum



Nemastoma bidentatum bidentatum × Nemastoma bidentatum sparsum – Novak & Gruber 2000: 285



Diagnosis



 without proximal dorso-medial hump.

Material examined

CROATIA – WL97et al.

XL06













SLOVENIA – VL13   

VL26



VL27

N OVA K T. et al., Nemastoma bidentatum (Opiliones: Nemastomatidae) revison

VL34



VL58

VL67

VL78

VL79

VL84



VL88



– VL93VL99



WL04



WL05

WL06



WL08



WL09WL19

VL28

WL29

WM01



WM40

WM42



Nemastoma bidentatum gruberi ssp. nov. × N. bidentatum sparsum 



Diagnosis



 



the tip.

Material examined

CROATIA – XL03

Nemastoma bidentatum martensi ssp. nov. × N. bidentatum schmidti ssp. nov.



Nemastoma (Stridulostoma) seliskari 

Nemastoma bidentatum bidentatumNemastoma (Stridulostoma) seliskari



Nemastoma bidentatum bidentatum × N. bidentatum sparsum – 



European Journal of Taxonomy 777: 1–67 (2021)

Nemastoma bidentatum pluridentatumNemastoma (Stridulostoma)

seliskari Nemastoma seliskari 

Diagnosis





N. b. schmidti 

minute hump.

Material examined

SLOVENIA – VL15

VL35

VL53

WL05

– WL17



Remarks

  Nemastoma bidentatum 



type specimen of Nemastoma (Stridulostoma) seliskari

Stridulostoma based on the existence of an alleged “stridulatory

 



from N. (Lugubrostoma) triste pluridentatum

Stridulostoma with Nemastoma

36) synonymized N. seliskari with N. b. pluridentatum







Nemastoma collected later on Mt. Mirna







N. (Lugubrostoma)

triste pluridentatum. See remark under N. pluridentatum stat. nov.

Nemastoma bidentatum schmidti ssp. nov. × N. bidentatum sneznikensis ssp. nov.

Diagnosis

sneznikensis

N. b. schmidti 

schmidti

Material examined

SLOVENIA – VL54



N OVA K T. et al., Nemastoma bidentatum (Opiliones: Nemastomatidae) revison

Remarks



Nemastoma bidentatum schmidti ssp. nov. × N. bidentatum sparsum 



Nemastoma triste

Nemastoma bidentatum bidentatum × N. bidentatum sparsum



et al. 2002:



Nemastoma bidentatum

Diagnosis



either being relatively tiny with more or less elongated superior portion or more massive with superior

 

both subspecies.

Material examined

SLOVENIA – VM91 









VM92

WM01

WM05



WM06 •

WM10



WM11



WM12



WM13





WM16

WM20

WM21





WM22 



WM23





European Journal of Taxonomy 777: 1–67 (2021)























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



WM24



WM25





WM30

WM31



WM33



WM40

WM41





WM42







WM43 •



WM44

WM45

– WM52

   



WM53



WM62













N OVA K T. et al., Nemastoma bidentatum (Opiliones: Nemastomatidae) revison

WM63

WM64

WM65



WM73











WM76



WM83

WM84 •



WM85

WM94

WM95

WM95







Remarks

N. b. schmidti ssp. nov. has been reported under N. b. bidentatumN. b. gruberi ssp. nov. under

N. b. sparsum. Since hybrids N. b. bidentatum × N. b. sparsum

all these published data likely refer either to N. b. schmidti ssp. nov. × N. b. sparsum or N. b. gruberi

ssp. nov. × N. b. schmidti ssp. nov. Nemastoma b. bidentatum referred from Germany (Schönhofer &

N. b. schmidti ssp. nov. × N. b. sparsum. Since both N. b. sparsum and

N. b. gruberi 

belong to N. b. gruberi ssp. nov. × N. b. schmidti ssp. nov.

Identication key to taxa within the Nemastoma bidentatum Roewer, 1914 complex

N. bidentatumN. bidentatum

s. str. including typical male hybrids. Female hybrids show intermediate traits and are not considered



not referred in the key. Detailed morphology is presented in Figs 6–13. Consult characters presented in



  ....................................................................................................................................................... 2

  ..................................................................................................................................................... 16

2. Ch and     ........................................................ 3

– Ch and     ................................................

...........................................................................................N. pluridentatum 

   .................................... 4

 

shaped ............................................................................................................................................... 5

European Journal of Taxonomy 777: 1–67 (2021)

4. Ch-Apo     

(Fig. 12C) .................................................................. N. relictum 

– Ch-Apo  

  ........................................... N. b. sneznikensis ssp. nov.

5. Ch-Apo  

protrusions ........................................................................................................................................ 6

   

isosceles triangular (Fig. 12G) ...................................................................... N. b. martensi ssp. nov.

 

 ......................................................................................................................................... 7

  ........................ 13

  .................................................... 8

  .............................................................................. 10

8. Ch-Apo either broadly quarter moon-like or elongated oval-trapezoid ........................................... 9

  .......................................

.................................................................N. b. bidentatum N. b. schmidti ssp. nov.

 

 

glans terminally roundly truncated (Fig. 12A) .................................N. b. bidentatum 

 



  glans terminally evenly truncated (Fig. 12H)

........................................................................................................................N. b. schmidti ssp. nov.

 .....................................................................11

  ........................................................................ 12

 .....................................

............................................................................... N. b. gruberi ssp. nov. × N. b. schmidti ssp. nov.

 

 ...........................................................................................................

....................................................N. b. schmidti ssp. nov. × N. b. sparsum

  



(Fig. 12F) ................................................................................................................ N. kozari sp. nov.

    

10L) .............................N. b. gruberi ssp. nov. × N. b. martensi ssp. nov. × N. b. schmidti ssp. nov.

13. Ch-Apo rhomboidal or apically extended ....................................................................................... 14

  with either 2 stout spines or tubercle and

 .............N. b. sparsum

14. Ch-Apo apically extended .............................................................................................................. 15

  with either 4 subequidistantly positioned spines or pointed tubercles or



 ....................

......................................................................................................................... N. b. gruberi ssp. nov.

N OVA K T. et al., Nemastoma bidentatum (Opiliones: Nemastomatidae) revison

  ...........................

......................................................N. b. gruberi ssp. nov. × N. b. sparsum 

  with 5 equidistant denticles ......

...............................................................................N. b. gruberi ssp. nov. × N. b. martensi ssp. nov.

 .......................... 17

 

 ......................................N. b. sneznikensis ssp. nov.

17. Apex of  ....................................................... 18

  ....................................................................................... 19

18. Frontal margin of Ch 

 

  ........................................

...........................................................................................................N. b. bidentatum 

– Frontal margin of Ch 

  .........................................

......................................................................................................................... N. b. schmidti sp. nov.

19. Rec sem elongated saccular-like ..................................................................................................... 20

– Rec sem balloon-like ...................................................................................................................... 21

20. Ch  

alveolus (Fig. 13C) ...................................................N. relictum 

– Ch 

alveolus originating in bladder-like structure (Fig. 13E) ........................................N. kozari sp. nov.

  ...... 22

    .............

....................................................................................................................... N. b. martensi ssp. nov.

 

 ................................................................. N. b. sparsum 

   

 ...................................................................... N. b. gruberi ssp. nov.

Biogeography

N. b.N. bidentatum exist: N. b. bidentatum

N. b. gruberiN. b. martensiN. b. schmidtiN. b. sparsum and N. b. sneznikensis

ssp. nov. The distributional areas of these taxa cover most of the country (Fig. 3). Southern Austria with

hitherto known three subspecies of N. bidentatum and one more species: N. b. bidentatumN. b. schmidti

 N. b. sparsum and N. relictum

N. b. gruberiN. b. martensiN. b. schmidti ssp. nov. and N. b. sparsum additionally

contribute to the bulk of the N. b.

known in Bosnia and Herzegovina: N. b. sparsumN. pluridentatum stat. nov. and N. kozari

two subspecies: N. b. gruberi ssp. nov. and N. b. schmidti N. b. sparsum

N. b. schmidti  only N. b. sparsum occurs in all the other mentioned



subspecies areas are not simple lines.

European Journal of Taxonomy 777: 1–67 (2021)

Discussion

Although the process of dividing the initially single genus Nemastoma



Nemastoma s. str. and began the revision of the N. b. complex.

N. b.

(Gruber 1976; Gruber & Martens 1968; Martens 1978; Novak & Gruber 2000; Komposch & Gruber 2004;



these with an uncertain status: N. b. bidentatumN. b. sparsumN. b. relictum and N. b. pluridentatum







subspecies of N. bidentatum N. bidentatum

and one species in Austria and four subspecies of N. bidentatum in Croatia. Only one or two taxa sojourn

in all the other mentioned countries. 

the N. bidentatum N. bidentatum

complex and nearly unexplored Nemastoma fauna might be the center of the N. bidentatum complex

species diversity.

Fig. 4. Nemastoma bidentatum

N OVA K T. et al., Nemastoma bidentatum (Opiliones: Nemastomatidae) revison

Fig. 5. Nemastoma bidentatum 

European Journal of Taxonomy 777: 1–67 (2021)

Fig. 6. Nemastoma bidentatum basal



N OVA K T. et al., Nemastoma bidentatum (Opiliones: Nemastomatidae) revison

N. b. schmidti

N. b. bidentatumN. b. martensiN. b. gruberiN. b. sneznikensis

ssp. nov. and N. b. sparsum in the southwest to the northeast of the country. The central position of

N. b. schmidti    







in the bidentatum – schmidti ssp. nov. – sparsum N. b. schmidti ssp. nov. interbreeds with

N. b. bidentatum and N. b. sparsum (Gruber & Martens

1968; Martens 1978; Komposch & Gruber 2004; Komposch 2009). This indicates a speciation process





distant populations if co-occuring secondarily in syntopy.

 N. b.



useful information. This is because of the varying stability of particular characters across the taxa. The

Fig. 7. Nemastoma bidentatum    

N. pluridentatum

European Journal of Taxonomy 777: 1–67 (2021)



N. b. bidentatum and N. b. schmidti



N. b. bidentatum

N. b. martensi the number and shape of denticles vary from



N. b. sneznikensis

N. b. gruberi







 indicates either N. bidentaum b. or N. b. schmidti ssp. nov. As for



Fig. 8. Nemastoma bidentatum

of N. pluridentatum

N OVA K T. et al., Nemastoma bidentatum (Opiliones: Nemastomatidae) revison

N. b. bidentatum and N. b. schmidti ssp. nov.

N. relictum stat. nov. and N. kozari sp. nov. alone allow credible







Fig. 9. Nemastoma bidentatum



European Journal of Taxonomy 777: 1–67 (2021)





of females with Rec sem consisting of balloon-like vesicles alone is not certain.

Fig. 10. Nemastoma bidentatumA–B

CD–EF–GH–JL–MN–O: lateral view; the others in dorso-medial

view to show the armament).

N OVA K T. et al., Nemastoma bidentatum (Opiliones: Nemastomatidae) revison



with respect to other such adjacent populations while interbreeding with them. The standard quantitative



valid if at least 75% of a population lies outside 99% of the range of the overlapping population for a



N. bidentatum

N. b. gruberi ssp. nov.

et al. (2012) recommendation for subspecies status. This states that under











N. b. gruberi

ssp. nov. × N. b. sparsum



N. bidentatum

N. b. bidentatum is markedly distinctive from all

N. relictum stat. nov. because it has Rec



N. bidentatum

with another taxon of the N. b.N. pluridentatum



the N. b.N. kozari sp. nov. inhabiting the same mountain.

Fig. 11. Nemastoma bidentatum

FA. The frame encloses the ovipositor portion with Rec sem. Note:

Female N. pluridentatum

European Journal of Taxonomy 777: 1–67 (2021)

Fig. 12. Nemastoma bidentatum



N. pluridentatum

N OVA K T. et al., Nemastoma bidentatum (Opiliones: Nemastomatidae) revison

Fig. 13. Nemastoma bidentatum



female N. pluridentatum

European Journal of Taxonomy 777: 1–67 (2021)

Nemastoma relictumN. pluridentatum stat. nov. and N. b. sneznikensis ssp. nov. are relic taxa

inhabiting montane to alpine habitats.





these are currently lacking for N. b. sneznikensis ssp. nov. due to its sparseness. Further taxa within the

N. b.

A clear-cut contact zone between the subspecies of N. bidentatum



between the two populations (cf. Monahan et al.









or Nardion strictae and Hyperico grisebachii-Pinetum mugo







area of N. b. schmidti ssp. nov. and N. b. sparsum is over 100 km wide in eastern Slovenia. Taking into

account various environmental conditions related to the subspecies requirements in the main distribution



Field notes on microhabitats inhabited by individual taxa of N. b. complex allowed us to conclude that all

these taxa – with the exception of N. pluridentatum with unknown habitat preferences – are specialized





et al.







are very scarce or lacking at sites with compact litter due to abundant precipitation or the formation of

macroaggregates by fungal activity (Coleman et al.

N. b. complex are indicators of loose organic





N. b.

Dyspnoi in particular. In the following contributions we shall present chemotaxonomy based on the



N. bidentatum



alleviate current faunistic and ecological investigations.

Acknowledgements











N OVA K T. et al., Nemastoma bidentatum (Opiliones: Nemastomatidae) revison



















and suggestions for improvement of the manuscript.

Competing interests

The authors declare no competing interests.

Funding

This study was partly supported by the Slovenian Research Agency within the Biodiversity Research









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https://doi.org/10.1111/j.1463-6395.2008.00341.x



Opiliones). In: Smith A.M. (ed.) Biological Adhesives: 281–301. Springer.

https://doi.org/10.1007/978-3-319-46082-6

Zink R.M. 2004. The role of subspecies in obscuring avian biological diversity and misleading conservation

policy. Proceedings of the Royal Society London B 271: 561–564. https://doi.org/10.1098/rspb.2003.2617

Manuscript received:30 April 2021

Manuscript accepted: 24 August 2021

Published on: 28 October 2021

Topic editor: Rudy C.A.M. Jocqué

Desk editor: Radka Rosenbaumová

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ResearchGate has not been able to resolve any citations for this publication.

Weberknechte (Opiliones) des Zentral-Europäischen zoogeographischen Gebietes Sloweniens.

Article

Full-text available

Jan 1995

A faunistic review of the harvestmen (Opiliones) of the Central European zoogeographical region in Slovenia is presented and some remarks on species are added.

Morphologie, Systematik und Ökologie der Gattung Nemastoma C. L. Koch (s. str.) (Opiliones, Nemastomatidae). [Morphology, systematics and Ecology of the genus Nemastoma C.L. Koch (s. str.) (Opiliones, Nemastomatidae).]

Article

Full-text available

May 1968
Senckenberg Biol

Genus Nemastoma C.L. Koch is re-circumscribed based on male genital morphology; it comprises seven southeast, central and northwest European species. Two species are described as new: Nemastoma schuelleri n. sp. (Austrian Alps, male/female) and Nemastoma transsylvanicum n. sp. (Romania, male/female). Within Nemastoma bidentatum three subspecies are characterized mainly based male cheliceral and pedipalpal morphology: nominate N. bidentatum, N. bidentatum sparsum n. ssp. and N. bidentatum relictum n. ssp. (all mainly Austrian and Slovenian Alps). Nemastoma lugubre and Nemastoma bimaculatum are recognized as different largely allopatrically distributed species which are not closely related. Detailed synonymies, distributional and ecological data for all species are given. For the enlarged dorsal scutal plate of Nemastomatidae the term Scutum compositum is introduced.

A CONTRIBUTION TO THE KNOWLEDGE OF THE HARVESTMEN (OPIUONES) FROM THE SUBMEDITERRANEAN REGION OF SLOVENIA

Article

Full-text available

Jul 2019

A faunistic review of the harvestmen (Opiliones) of the Submediterranean region of Slovenia is represented and the current taxonomlcal and zoogeographical problems of some species are discussed.

The harvestmen fauna (Arachnida: Opiliones) from the Sub Mediterranean region of Slovenia − II.

Article

Full-text available

Jul 2019

Tone Novak

In the present paper, further findings of harvestmen in the Sub Mediterranean region of Slovenia are presented, and comments on some actual problems with the species are added. With the exception of the possible discovery of a troglobiotic species in the region, all species living there have presumably been recorded. With the exclusion of Trogulus gr. nepaeformis and T. gr. tricarinatus, where more species are expected, 34 species and one subspecies have been found in the region. About 20 opilionid species re typical of the region; the others occur in the marginal habitats bordering other zoogeographical regions.

Harvestmen of the Museo Friulano di Storia Naturale in Udine (Arachnida: Opiliones). Part 1.

Article

Full-text available

May 2019

Tone Novak

Abstract - Since 1996, intensive faunal research has taken place in Friuli-Venezia Giulia in the regions of Udine, Pordenone and Gorizia, organised by the Museo Friulano di Storia Naturale in Udine. In addition, some opilionids from other regions have been deposited in the Museum. In this contribution, 46 opilionid species from 112 localities are considered, with the exception of the Trogulus nepaeformis–group. Trogulus falcipenis KOMPOSCH, 2000, and Ischyropsalis hellwigi hellwigi (PANZER, 1794) have for the first time been cited among the Italian fauna. The problems concerning the Trogulus nepaeformis–group will be considered separately.

Cave Ecology

Book

Full-text available

Nov 2018

Cave organisms are the ‘monsters’ of the underground world and studying them invariably raises interesting questions about the ways evolution has equipped them to survive in permanent darkness and low-energy environments. Undertaking ecological studies in caves and other subterranean habitats is not only challenging because they are difficult to access, but also because the domain is so different from what we know from the surface, with no plants at the base of food chains and with a nearly constant microclimate year-round. The research presented here answers key questions such as how a constant environment can produce the enormous biodiversity seen below ground, what adaptations and peculiarities allow subterranean organisms to thrive, and how they are affected by the constraints of their environment. This book is divided into six main parts, which address: the habitats of cave animals; their complex diversity; the environmental factors that support that diversity; individual case studies of cave ecosystems; and of the conservation challenges they face; all of which culminate in proposals for future research directions. Given its breadth of coverage, it offers an essential reference guide for graduate students and established researchers alike.

Methyl-ketones in the scent glands of Opiliones: a chemical trait of cyphophthalmi retrieved in the dyspnoan Nemastoma triste

Article

Full-text available

Apr 2018
CHEMOECOLOGY

The homologous and phylogenetically old scent glands of harvestmen—also called defensive or repugnatorial glands—represent an ideal system for a model reconstruction of the evolutionary history of exocrine secretion chemistry (“phylogenetic chemosystematics”). While the secretions of Laniatores (mainly phenols, benzoquinones), Cyphophthalmi (naphthoquinones, chloro-naphthoquinones, methyl-ketones) and some Eupnoi (naphthoquinones, ethyl-ketones) are fairly well studied, one open question refers to the still largely enigmatic scent gland chemistry of representatives of the suborder Dyspnoi and the relation of dyspnoan chemistry to the remaining suborders. We here report on the secretion of a nemastomatid Dyspnoi, Nemastoma triste, which is composed of straight-chain methyl-ketones (heptan-2-one, nonan-2-one, 6-tridecen-2-one, 8-tridecen-2-one), methyl-branched methyl-ketones (5-methyl-heptan-2-one, 6-methyl-nonan-2-one), naphthoquinones (1,4-naphthoquinone, 6-methyl-1,4-naphthoquinone) and chloro-naphthoquinones (4-chloro-1,2-naphthoquinone, 4-chloro-6-methyl-1,2-naphthoquinone). Chemically, the secretions of N. triste are remarkably reminiscent of those found in Cyphophthalmi. While naphthoquinones are widely distributed across the scent gland secretions of harvestmen (all suborders except Laniatores), methyl-ketones and chloro-naphthoquinones arise as linking elements between cyphophthalmid and dyspnoan scent gland chemistry.

Remarks on Opiliones froma cavities in Slovenia (Yugoslavia

Article

Full-text available

Jan 1984

Tone Novak

Morphology and Functional Anatomy

Chapter

Feb 2007

Research in Calcretes and Other Deep Subterranean Habitats Outside Caves: Analysis and Synthesis

Chapter

Jan 2018

Stuart Halse

The outstanding difference between traditional subterranean fauna studies and those carried out recently in Australia is the Australian emphasis on the fauna that occurs deep underground, but outside caves, across large parts of the landscape. This work has shown that the Australian arid zone, particularly in the western half of the continent, is rich in subterranean fauna, with the geologies supporting most species being calcrete and alluvium in the case of stygofauna and iron-rich rocks in the case of troglofauna. It is likely that, altogether, as many as 4500 species of stygofauna and troglofauna occur in the two most species-rich regions of Western Australia — the Pilbara and Yilgarn. Striking characteristics of the stygofauna communities in these regions include little overlap in species composition of communities of the hyporheic zone and deeper groundwater, very high levels of endemism in individual calcrete bodies, and the existence of extensive radiations of candonid ostracods in the Pilbara and copepods in calcretes of the Yilgarn. Characteristics of the troglofauna communities include extremely small ranges of many species, with linear ranges of 1–2 km apparently being common, and extensive radiation of schizomids and some other invertebrate groups in iron formations of the Pilbara.

Hidden diversity within the Nemastoma bidentatum Roewer, 1914 complex (Opiliones: Nemastomatidae) Part I: Morphological evidence

Abstract

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