A new species of Cryptops Leach, 1814, C. speleorex sp. nov., is described from Movile Cave, Dobrogea, Romania. The cave is remarkable for its unique ecosystem entirely dependent on methane- and sulfur-oxidising bacteria. Until now, the cave was thought to be inhabited by the epigean species C. anomalans, which is widespread in Europe. Despite its resemblance to C. anomalans, the new species is well-defined morphologically and molecularly based on two mitochondrial (cytochrome c oxidase subunit I COI and 16S rDNA) and one nuclear (28S rDNA) markers. Cryptops speleorex sp. nov. shows a number of troglomorphic traits such as a generally large body and elongated appendages and spiracles, higher number of coxal pores and saw teeth on the tibia of the ultimate leg. With this record, the number of endemic species known from the Movile Cave reaches 35, which ranks it as one of the most species-rich caves in the world.

Biospeleology, Cryptops speleorex sp. nov., Dobrogea, molecular phylogenetics, new species, troglomorphism

Located in the southeastern part of Romania not far from the Black Sea Coast, Movile Cave is the first known subterranean chemosynthesis-based ecosystem (Sarbu et al. 2019). Being completely isolated from the outside environment for 5.5 million years, the cave is remarkable for its unique ecosystem entirely dependent on methane- and sulfur-oxidising bacteria, which release nutrients through chemosynthesis for fungi and other cave animals along the food chain. This subterranean ecosystem is also notable for being rich in hydrogen sulfide, methane (1–2%), ammonia and CO₂ (1.5–3.5%) whereas it is poor in O₂ (7–16%). Relative humidity in the cave is 100% and there is no detectable air movement. The cave was first discovered in 1986 and since then, only a handful of people have visited it (Sarbu et al. 2019).

Despite its harsh living conditions, Movile Cave ecosystem is known to harbor a diverse and unique fauna. The cave hosts 51 invertebrate species, of which 34 species are endemic (Sarbu et al. 2019). Among these species, some present a number of unique adaptations to a troglobitic life in caves, such as the troglobiont water scorpion Nepa anophthalma Decu, Gruia, Keffer & Sarbu, 1994 (Hexapoda, Hemiptera, Nepidae); the nesticid and liocranid spiders Kryptonesticus georgescuae Nae, Serban & Weiss, 2018 (Araneae: Nesticidae) and Agraecina cristiani (Georgescu, 1989) (Araneae, Liocranidae); the cave leech Haemopis caeca Manoleli, Klemm & Sarbu, 1998 (Annelida, Hirudinea, Haemopidae) and the isopod Armadillidium tabacarui Gruia, Iavorschi & Sarbu, 1994 (Crustacea, Isopoda, Armadillidiidae) (Sarbu et al. 2019).

Five species of myriapods are hitherto discovered from the innermost parts of Movile viz. Archiboreoiulus serbansarbui Giurginca, Vănoaica, Šustr, & Tajovský, 2020 (Diplopoda), Symphylella Silvestri, 1902 sp. (Symphyla), Geophilus alpinus Meinert, 1870 and Clinopodes carinthiacus (Latzel, 1880) (Geophilomorpha) and a troglobitic population of Cryptops anomalans Newport, 1844 (Negrea 1993; Sarbu et al. 2019). It is worth mentioning that the latter taxon has been only studied morphologically (Negrea 1993, 2004).

Recently, we had the occasion to study freshly collected specimens of an undetermined species of the genus Cryptops Leach, 1814 from Movile Cave. Using both, morphological and molecular evidence, the cave specimens were compared with those of C. anomalans living on the surface, outside the cave. A phylogenetic analysis of 29 Cryptops specimens from different parts of Europe, including two from inside Movile Cave, based on two mitochondrial (cytochrome c oxidase subunit I COI and 16S rDNA) and one nuclear (28S rDNA) markers was performed. Morphological and molecular analyses confirmed that the cave specimens from Movile correspond to a new species, Cryptops speleorex sp. nov., that we describe herein. Additionally, we provide an annotated list and a key to the troglobitic Cryptops species in the world.

All Cryptops specimens from Movile Cave were hand-collected by the biospeleologists Serban Sarbu and A. Hillebrand and preserved in 70% or 96% ethanol. Microphotographs were obtained with a Nikon DS-Ri-2 camera mounted on a Nikon SMZ25 stereomicroscope using NIS-Elements Microscope Imaging Software with an Extended Depth of Focus (EDF) patch. Images were edited in Photoshop CS6 and assembled in InDesign CS6. Material is shared between the ISER – Emil Racoviță Institute of Speleology, Bucharest, Romania; IZB – University of Belgrade – Institute of Zoology, Faculty of Biology, Belgrade, Serbia; NHMW – Naturhistorisches Museum Wien, Austria; NMNHS – National Museum of Natural History, Sofia, Bulgaria and the ZMUT – University of Turku – Zoological Museum, Finland. In addition to the type material of the new species we have morphologically studied material of C. anomalans from Serbia and Romania.

Morphological terminology follows Bonato et al. (2010).

Abbreviations: T – tergite, S – sternite.

Order Scolopendromorpha Pocock, 1895

Family Cryptopidae Kohlrausch, 1881

Genus Cryptops Leach, 1814

Cryptops (Cryptops) anomalans Newport, 1844

Material examined

Romania: SE Romania: Lalomiţa County, Călugărească Forest, 18.II.2016, leg. and det. S. Baba, 1 subad. ex. (ISER); Lalomiţa County, Călugărească Forest, oak forest, 28.II.2019, leg. and det. S. Baba, 2 ex. (ISER) (lab code 54a); Lalomiţa County, Călugărească Forest, rotten wood, 13.III.2016, leg. and det. S. Baba, 1 ex. (ISER); Bucharest, Herăstrău Park, under stones, 10.X.2019, leg. and det. S. Baba, 1 ex. (ISER) (lab code 57a); Mangalia, Obanul Mare, Cave Drilling, -3 m, 10.VIII.1999, det. St. and A. Negrea, 1 ex. (ISER); Mangalia, Obanul Mare, Cave Drilling, -8 m, 27.V.2000, det. St. and A. Negrea, 1 ex. (ISER); Mangalia, Obanul Mare, Cave Drilling, -8 m, 28.VI.2000, det. St. and A. Negrea, 1 ex. (ISER); Mangalia, Obanul Mare, Cave Drilling, -12 m, 27.V.2000, det. St. and A. Negrea, 1 ex. (ISER). Serbia: Valley of the Izbice River, v. Izbice, near Novi Pazar, SW Serbia (43°07.333'N, 20°34.354'E; elevation about 700 m a.s.l.): 5♂, 5♀, collected in 2012 (May-October), leg. D. Stojanović (lab code 1) (IZB); Prolom Banja Spa, near Kuršumlija, southern Serbia (43°02.449'N, 21°23.448'E; elevation about 620 m a.s.l.): 3♀, collected 30.04.2016., leg. D. Stojanović (lab code 2) (IZB); village Kacabać, near Bojnik, Leskovac, southern Serbia (43°03.415'N, 21°46.368'E; elevation about 200 m a.s.l.): 2♂, 1♀, collected 01.05.2016., leg. D. Stojanović (lab code 3) (IZB); Pećina Rasnica 1 Cave, village Rasnica, near Pirot, SE Serbia: 1♂, 1♀, collected 18.07.2018., leg. D. Antić (lab code 4) (IZB); Novopazarska Banja Spa, near Novi Pazar, SW Serbia (43°09.269'N, 20°33.132'E; elevation about 650 m a.s.l.): 3♂, 4♀, collected 30.05.2012., leg. D. Stojanović (IZB); Spomen Park, Leskovac, southern Serbia (42°59.051'N, 21°56.349'E; elevation about 200 m a.s.l.): 1♀, collected 28.07.2012., leg. D. Stojanović (IZB); pine forest near the Đurđevi Stupovi Monastery, Novi Pazar, SW Serbia (43°09.183'N, 20°30.049'E): 1♀, collected 15.05.2015., leg. D. Stojanović (lab code 7) (IZB); village Dobanovci, near Surčin, Belgrade, Serbia (44°49.197'N, 20°13.334'E): 1♀, collected 03.11.2013., leg. D. Stojanović (lab code 8) (IZB); Bojčinska šuma forest, village Progar-Jakovo, near Surčin, Belgrade, Serbia (44°43.528'N, 20°09.245'E): 3♀, 1♂, collected 09.06.2013., leg. D. Stojanović, K. Bjelanović (lab code 9) (IZB); Vrla River, Mt. Vlasina, near “Rosa” water factory, v. Topli Do, near Surdulica, SE Serbia (42°38.213'N, 22°17.565'E; elevation about 1070 m a.s.l.): 1♂, collected 07.07.2011, leg. D. Stojanović (IZB); Višnjička Banja, Belgrade, Serbia (44°49.073'N, 20°32.337'E; elevation about 350 m a.s.l.): 1♂, collected 09.06.2006, leg. Ž. Pavković (IZB); Spomen Park, Leskovac, southern Serbia (42°59.051'N, 21°56.349'E; elevation about 200 m a.s.l.): 3♀, 1♂, collected 14.04.2012., leg. D. Stojanović (lab code 12) (IZB). Bulgaria: Pirin Mts, between Sandanski and Lilyanovo, 12.8.1988, litter, mainly Platanus, P. Beron leg. 1 ex. (NMNHS) (Figs 2B, 3B, 4B,D, 5B).

Cryptops (Cryptops) speleorexsp. nov.

http://zoobank.org/8A28E7DF-168B-485C-8A0C-CD7EB218E650

Figs 1A, B, 2A, 3A, 4A, C, 5A, 6A–C

Previous records

Cryptops anomalans: Negrea, 1993: p. 87 and all subsequent records (Negrea 1994, 1997, 2004; Negrea and Minelli 1994; Sarbu et al. 2019).

Figure 1. 

Cryptops speleorex sp. nov. A holotype, habitus, dorsal view B paratype (ZMUT), posteriormost segments and ultimate legs, dorsal view.

Material examined

Holotype : Romania: Constanța County, Mangalia, Movile Cave (Peștera Movile), Lake Hall, June, 2014, leg. S. Sarbu, 1 ex. (NMNHS, Myriapoda Collection Id: 10 812); Paratypes: same locality and collector leg. S. Serban, 1 ex. (NHMW10177); same locality, 22.XI.2017, leg. A. Hillebrand, 1 ad. ex., identified as C. anomalans by Stefan Baba (ISER); 1 ad. ex., same locality, date and collector, identified as C. anomalans by Stefan Baba (http://mus.utu.fi/ZMUT.MYR-TYPE001).

Diagnosis

A species morphologically similar to Cryptops anomalans, but differing from it by the much elongated antennae and legs, generally less setose forcipules and body, coxopleures with more than 300 coxal pores (vs. less than 100 in anomalans), ultimate leg with 13–17 saw teeth on tibia (usually 7–10, occasionally 12 in anomalans), and larger and elongated spiracles (see Table 2). Genetically, Cryptops speleorex sp. nov. differs from the C. anomalans specimens from Romania and Serbia by 9.2–12.2% in COI and 6.6–8.7% in 16S rDNA.

Table 2.

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Differences in morphological characters between Cryptops anomalans and C. speleorex sp. nov.

Morphological character	Cryptops anomalans	Cryptops speleorex sp. nov.
Body size (mm)	25–50	>46–52
Antennae length	Until posterior end of T3	Until mid of T5
Antennal article 7 L/W (mm)	0.5 × 0.25	1.0 × 0.5
Antennae: spines on basal articles	Present, numerous	Lacking or just a few
Ultimate leg length	7.65 mm	13.25 mm
Ultimate leg pretarsus (mm)	0.25	1
Ultimate leg saw teeth on tibia and tarsus 1	Tibia: 7–12 (usually 7–10); Tarsus: 3–5	Tibia: 13–17; Tarsus: 5–6
Legs	Short, compact, pretarsus short	Strongly elongated, pretarsus long
Spiracles	Ovoid, small to medium sized (Fig. 4D)	Strongly elongated, large (Fig. 4C)
Forcipular trochanteroprefemur	With spines medially (4–6)	Without spines, just stout setae
Coxopleural pore field	Approx. 2/3 of coxapleura; composed of less than 100 pores (86–90)	Approx. 4/5 of coxopleura; composed of more than 310 pores (317–320)

Description (holotype)

Length (anterior margin of head plate to posterior margin of telson) approx. 52 mm (46 mm in an adult paratype) (Figs 1A, B). Head plate (Fig. 2A) 3.2 mm long, 3.4 mm broad; antenna approx. 10 mm long. Body yellow-brownish (Fig. 1A); antennae and legs pale yellow; posterior edge of head and tergites with irregular light brownish band, darker in the middle (Figs 1B, 2A); forcipular tarsungulum and leg claws dark brown. Head plate overlaps approx. 1/3 of tergite 1; head plate slightly broader than long (3.2 mm × 3.4 mm), posterior corners strongly rounded, sides convex outwards, anterior apex slightly indented at the base of antennae, bisected by longitudinal median furrow; paramedian sutures diverging anteriorly on head plate; head punctate, sparsely covered with fine setae.

Figure 2. 

Cryptops spp., head and anteriormost segments A Cryptops speleorex sp. nov., holotype, dorsal view B Cryptops anomalans, Pirin Mts (Bulgaria), dorsolateral view (slightly apical).

Antenna relatively long, extending to the middle of tergite 5 when folded backward (Figs 1A, 2A); composed of 17 articles; article length formula: 17<1<2=16<3=4=13=14<5=6=11=12<7–10; basal two articles relatively stout, in general articles increase in length to a maximum at articles 7–10, then gradually shortening; article 17 is more than half length of article 16 (approx. 60%); articles 5–10 much longer than wide, length up to 3 times the width. All surfaces of antennal articles with scattered long setae, densest on articles 1–3; short, fine setae abundant on all articles except for articles 1 and 2, as well as basal part of 3.

Clypeus with 2 setae; prelabral setae in one row of 21–22; 4 short setae between clypeus and prelabral row, irregularly or more evenly scattered. Labral mid piece with a short, but well-developed tooth; side pieces rounded (Fig. 3A).

Figure 3. 

Cryptops spp., forcipular coxosternum, ventral view A Cryptops speleorex sp. nov., holotype B Cryptops anomalans, Pirin Mts (Bulgaria).

Forcipular segment anterior margin of coxosternite convex on each side, with a weak median diastema, fringed by 2 marginal setae on each side. Surface of coxosternite (Fig. 3A) covered with scarce short setae, 10–15 in total; trochanteroprefemur stout, median margin slightly expanded proximally, with 4 setae; femur and tibia very short; tarsungulum long, curved, almost equal in length to trochanteroprefemur’s height.

Maxilla 2 with a well-developed pretarsus; dorsal brush white, dense, situated on the distalmost part of article 3 of telopodite. Proximal side of first maxillary telopodite covered by 10–15 setae (Fig. 3A).

Tergites Tergite 1 with a complete anterior transverse suture and cruciform sutures (Figs 1A, 2A). Oblique sutures present on tergites 2–8; complete paramedian sutures on tergites 2–20; lateral crescentic sulci visible on tergites 6–20; all tergites nearly devoid of setae, occasionally individual scattered short setae. Tergite 21 longer than wide, posterior margin subtriangular, with rounded apex; shallow median depression along posterior half of tergite (Fig. 1B).

Sternites 1–2 and 19–21 without transverse and median sutures; S 3–18 with median longitudinal and curved transverse sutures, more prominent from sternite 5 onward (Fig. 4A). All sternites covered by minute setae. Endosternite: subtrapezoidal, lateral margins very slightly convex, posterior margin slightly concave in the middle; surface with several (6–10) moderately long and sparse setae.

Figure 4. 

Cryptops spp., sternites 8–9 and spiracles A, B sternites 8–9 A Cryptops speleorex sp. nov., holotype, arrow indicating the endosternite B Cryptops anomalans, Pirin Mts (Bulgaria) C, D spiracles C Cryptops speleorex sp. nov., holotype D Cryptops anomalans, Pirin Mts (Bulgaria).

Spiracles strongly elongated on T3, reducing in size towards the posterior end of the body; slit-like (Fig. 4C).

Coxopleural pore field elliptical, covering 4/5 of surface, with more than 310 coxal pores (317–320), extending nearly to posterior margin of coxopleuron (Fig. 5A). Approx. 15–20 sparse spiniform setae emerging between pores and from the dorsal and posterior margins of coxopleuron.

Figure 5. 

Cryptops spp., Coxopleural pore field A Cryptops speleorex sp. nov., holotype B Cryptops anomalans, Pirin Mts (Bulgaria).

Legs generally long; leg 10: prefemur 1.47 mm long, femur 1.59 mm, tibia 1.76 mm, tarsus 2.35 mm, pretarsus 0.7 mm. All tarsi single (Fig. 6A). Walking legs (Fig. 6A, B) smooth, generally poor in setae; spiniform setae sparsely present on the surface of prefemur, and occasionally also on the femur; all pretarsi long, with an anterior and posterior accessory spines of different size, the larger being 2/3^rd of pretarsus; accessory spines absent on leg 21; 20 leg: prefemur, femur and tibia slightly swollen; femur and tibia being slightly concave at midlength; a specific field of dense, minute setae present on the ventral, lateral and mesal sides of prefemur, femur and part of tibia.

Figure 6. 

Cryptops speleorex sp. nov., legs. A holotype, walking leg B paratype (ZMUT), walking leg, close-up of apical claw C holotype, ultimate legs, lateral view D, E paratype (NHMW), distal articles of ultimate teeth showing saw teeth.

Ultimate leg (Fig. 6C): prefemur 3.61 mm long, femur 3.05 mm, tibia 1.94 mm, tarsus 1: 1.28 mm, tarsus 2: 2.22, pretarsus 0.56 mm.; numerous robust spiniform setae on the ventral, mesal and less so on lateral and dorsal sides of prefemur; spiniform setae present also on the ventral and mesal sides of femur; tibia, tarsus 1 and tarsus 2 covered by tiny dense setae on all sides; 13–14 saw teeth on tibia (17 in an adult paratype) and 5–6 on tarsus 1.

Etymology

The species epithet is a noun in apposition, meaning "king of the cave", referring to the species top position in the food chain of the Movile ecosystem.

Distribution

The species is hitherto known only from the aphotic zone of the Cave Movile in the southern part of Romanian Dobrogea.

Ecological remarks

Cryptops speleorex sp. nov. is the largest invertebrate species in Movile Cave. It has been observed feeding on terrestrial isopods (Trachelipus troglobius Tabacaru & Boghean, 1989, Armadillidium tabacarui Gruia, Iavorschi & Sarbu, 1994), smaller beetles, Diplura or spiders (Sarbu et al. 2019).

Parsimony analysis resulted in a single most-parsimonious (MP) tree of length 1586 steps (Fig. 7). Two C. speleorex sp. nov. specimens collected from Movile Cave (samples K3 and K4) group within C. anomalans as a separate clade supported by jackknife resampling value (hereafter JF) of 99. The phylogeny shows the Movile Cave clade being evolutionary most closely related to the clade (JF = 75) including C. anomalans samples from southern Serbia and Belgrade area (JF = 100) and Romania and SW Serbia (JF = 84). This Serbian/Romanian clade forms a sister group with the clade (JF = 95) containing a single C. anomalans specimen (lab code 4) from southeast Serbia (collected from a cave) and identical sequences of C. anomalans from London, UK and different parts of Germany (JF = 100). All specimens above form a clade with strong support (JF = 92). Outside this clade are Cryptops sp. from Austria and an unsupported clade containing Cryptops spp. from Croatia and Slovenia together with C. hortensis (Donovan, 1810). Basal to these are resolved C. parisi Brolemann, 1920 and C. croaticus Verhoeff, 1931 (JF = 82) followed by Cryptops sp. from Germany.

Figure 7. 

The single most parsimonious tree of length 1586 steps with jackknife resampling values > 50% shown on the nodes. Branch lengths represent the number of optimized character-state changes.

Regarding the placement of C. speleorex sp. nov. and the relationships among the C. anomalans specimens, the likelihood analysis (Fig. 8) resulted in a mostly congruent tree topology with the parsimony tree, the only difference being that in the parsimony analysis C. speleorex sp. nov. is resolved basal to the Serbian/Romanian clade whereas in the likelihood tree it is resolved within it. The C. speleorex sp. nov. specimens form a clade supported by bootstrap value (hereafter BS) of 100. Cryptops speleorex sp. nov. groups together with the C. anomalans specimens from Serbia (excluding a single Serbian C. anomalans specimen, lab code 4) and Romania. All the specimens above form a sister clade to a group including C. anomalans specimens from Serbia (lab code 4), Germany and the UK. As in the parsimony analysis, the additional Cryptops species (other than C. anomalans) were resolved as basal to C. anomalans. Their internal grouping varies from that in the parsimony tree, which is not surprising due to the lack of nodal support in the basal-most nodes.

Figure 8. 

Likelihood tree with bootstrap values > 50% shown for each node.

When analyzed separately (only likelihood, tree not shown), the mitochondrial COI and 16S resolved C. speleorex sp. nov. as a distinct clade (BS = 100) within C. anomalans specimens, the tree topology regarding C. speleorex sp. nov./C. anomalans being identical to that of the parsimony tree. Not surprisingly, the level of variation in the nuclear 28S was low and the likelihood analysis based on it could not resolve the relationships among the C. anomalans/C.speleorex sp. nov. specimens (tree not shown).

Scolopendromorphs are strictly terrestrial and most species are found in forest leaf litter, decomposed wood, under bark of dead trees, in the soil, under stones or in caves in the temperate and tropical areas of the world. Few species are well adapted to eremic environments (Minelli and Golovatch 2013), occasionally in atypical habitats such as forest canopy (Lewis 1982; Phillips et al. 2020) or tropical rivers (Siriwut et al. 2016). Although less common than lithobiomorphs, scolopendromorphs may occur in caves, where they are represented with some highly adapted species, mainly from the family Cryptopidae. Other families are only marginally recorded in caves: Scolopocryptopidae (genera Thalkethops Crabill, 1960 and Newportia Gervais, 1847 with several species from American caves, including several troglobites), Plutoniumidae (genera Plutonium Cavanna, 1881 and Theatops Newport, 1844) in European caves and Scolopendridae (genus Otostigmus Porat, 1876; O. cooperi Chamberlin, 1942 inhabits Chilibrilo caves in Panama (Chamberlin 1942); Otostigmus troglodytes Ribaut, 1914 found in a cave near Tanga, Tanzania (Ribaut 1914)). The genus Cryptops is by far the most frequent in the caves worldwide with some 18–20 species found in caves in South Europe (Spain, France, Italy, Greece), Canary Islands, Cuba, Brazil, Australia and Africa. Troglomorphic species are known from the nominate subgenus, and the subgenera Trigonocryptops and Paracryptops (see Table 6).

Several morphological characters traditionally used in centipedes taxonomy could be subject to intraspecific variation related to postembryonic development, animal life stage and ecology (Akkari et al. 2017). This might render species identification problematic in some cases and generates taxonomic errors. This is also true for such a highly variable and widely distributed species as C. anomalans. In fact, nine species and subspecies were hitherto synonymised with this species (see Krapelin 1903; Verhoeff 1931; Crabill 1962; Zapparoli 2002). Three subspecies are still listed as valid for it (Chilobase 2.0). Now the identity of these taxa and the presence of any possible cryptic species within C. anomalans could only be revealed via an integrative study combining morphological and molecular markers. Whereas clear molecular differences are here indicated by the different markers and the high interspecific distance between C. anomalans and the newly described species C. speleorex sp. nov., the morphological comparison was not as straightforward since both species show several similarities, including an overlapping in size. While several of the differences observed between both species (Table 2) could be understood as a clear indication of troglomorphism in C. speleorex sp. nov. such as the elongation of appendages, a few other characters including the number of saw teeth on tibia and tarsus 1 of the ultimate legs, number of coxal pores and the shape of spiracles were diagnostic to separate both species.

Intraspecific distance between the two sequenced Cryptops speleorex sp. nov. specimens is relatively high in comparison to the detected interspecific variation (Tables 3–5) raising a question whether these two specimens could actually be interpreted as two separate species. However, this variation is only shown in the two mitochondrial markers – there are no morphological differences (or any difference in their nuclear 28S marker) between the C. speleorex sp. nov. specimens. As Morgan-Richards et al. (2017) well explains, cryptic speciation should never be used as a null hypothesis in the absence of phenotypic or nuclear data supporting it. Instead, “the origin of the divergent mtDNA haplogroups might result from complex biogeographical scenarios or they might simply represent normal, stochastic processes of mutation and extinction of a non-recombining locus within a large population”.

Taxonomic and evolutionary implications of C. speleorex sp. nov

The type locality of C. anomalans is unknown and therefore it is impossible to conclude which part (if any) of the studied population is the actual C. anomalans described by Newport (1844). Before this study, only a handful of C. anomalans specimens from a limited geographic range had been sequenced (Spelda et al. 2011; Vahtera et al. 2013; Wesener et al. 2016). We acknowledge that describing C. speleorex sp. nov. as a new species leaves C. anomalans paraphyletic and that monophyly is violated by this taxonomic act. However, we view this as an inevitable consequence of speciation with a particular evolutionary implication, i.e., that C. speleorex sp. nov. evolved within what is currently known as C. anomalans. It is worth noting that the closest evolutionary relatives of C. speleorex sp. nov. appear to be the C. anomalans specimens from Serbia (excluding the sample number 4) and Romania (Figs 8, 9). This means that they are most closely related to each other than either of them is to the rest of the studied C. anomalans populations. The current situation with C. anomalans should not be seen as a failed taxonomy but as a natural consequence when new data from a widespread species is obtained.

Figure 9. 

Map of Europe showing geographic distribution of Cryptops specimens analyzed herein. Asterisk – C. speleorex sp. nov. Dot – other Cryptops spp. used in the study (see Table 1 for details).

We are especially grateful to Serban M. Sarbu (Adjunct Faculty, California State University Chico) for calling our attention to this interesting material and for committing samples from Movile Cave for study. Stefan Baba (“Emil Racoviță” Institute of Speleology & Faculty of Biology, University of Bucharest, Romania), Dragan Antić and Dalibor Stojanović (both from University of Belgrade – Faculty of Biology) committed further specimens of C. speleorex and C. anomalans from Serbia and Romania for study. The study was partially funded by project #KP-06-H21/1-17.12.2018 of the National Science Fund, Ministry of Education and Science of the Republic of Bulgaria to PS and by Helsinki Entomological Society to VV. We thank G.D. Edgecombe, C. Martínez-Muñoz, A. Schileyko and Ivan H. Tuf for their constructive comments that greatly benefited the manuscript.