Neobisium babinzub sp. n. and Neobisium tarae sp. n. (Neobisiidae, Pseudoscorpiones) from the central Balkan Peninsula (Serbia)

Biologia, Bratislava, 62/1: 78—83, 2007 Section Zoology DOI: 10.2478/s11756-007-0013-8

Neobisium babinzub sp. n. and Neobisium tarae sp. n. (Neobisiidae, Pseudoscorpiones) from the central Balkan Peninsula (Serbia) ´ ´, Rajko N. Dimitrijevic ´, Vladimir T. Tomic ´ & Bojan M. Mitic ´ Božidar P.M. Curči c Institute of Zoology, Faculty of Biology, University of Belgrade, Studentski Trg 16, 11000 Belgrade, Serbia; e-mail: [email protected]

Abstract: Two new pseudoscorpion species, Neobisium babinzub sp. n., and N. tarae sp. n. (Neobisiidae, Pseudoscorpiones), were collected in soil and leaf-litter on Mt. Stara Planina and Mt. Tara, Serbia, diagnosed and thoroughly illustrated. The main morphological characteristics of the two taxa clearly demonstrate their adaptation to life in soil. Key words: pseudoscorpions; Neobisiidae; Neobisium babinzub; Neobisium tarae; soil fauna; Serbia

Introduction

Material and methods

The family Neobisiidae and its nominal genus, Neobisium J.C. Chamberlin, are widely distributed through the Balkan Peninsula. In fact, research shows that soil fauna differs from the epigean fauna both by its general composition and by its marked endemism. The endemic pseudoscorpions found in the Dinaric Karst represent 80% of the total number of known species in the ´ region (Curči´ c et al. 2004); in the Carpatho-Balkanic Arc such a percentage varies between 60% and 65% ´ (Curči´ c et al. 2004). Moreover, endemism itself is not limited to the species, but extends to the subgenera and genera. In fact, there are autochthonous forms differentiated entirely within one site, as well as archaic species extinct elsewhere. The extraordinary richness of the Balkan Peninsula in pseudoscorpions (about 400 ´ ´ species and subspecies; Curči´ c 1986, 1988a; Curči´ c et al. 2004) shows clearly the advanced degree of differentiation of species. Furthermore, the close relationship between many of these endemic organisms and fossils from the Tertiary period indicates their origin, and relict nature (Schawaller 1978, 1980; Poinar et al. 1998). During the study of some Serbian pseudoscorpions from two small collections made by D. Pavi´cevi´c (Belgrade) and S. Peši´c (Kragujevac), we found two species pertaining to the genus Neobisium J.C. Chamberlin (Neobisiidae), respectively. The first species, N. babinzub sp. n., was found on Mt. Stara Planina (= Mt. Balkan) in Eastern Serbia, and the second one, N. tarae sp. n., on Mt. Tara, in Western Serbia. This paper presents distributions of both species and compares these with their phenetically closest congeners. Furthermore, some taxonomic and biogeographical features of these species are briefly discussed.

Two samples of epigean (soil) pseudoscorpions from Serbia have been subjected to a thorough analysis. The specimens analysed were dissected, illustrated and diagnosed. These are kept in gum-chloral medium (on separate slides) in the collection of the Institute of Zoology, Belgrade. Drawings in Indian ink were completed by the use of Ergaval and Axioscope stereomicroscopes (Carl Zeiss, Jena). ´ Setal designations follow Beier (1932) and Curči´ c (1977).

c 2007 Institute of Zoology, Slovak Academy of Sciences


Neobisiidae J.C. Chamberlin, 1930 Neobisium J.C. Chamberlin, 1930 Neobisium babinzub sp. n. (Figs 1–8, Table 1) Description. Anterior carapacal margin slightly convex medially, with a small knob-like epistome (Figs 3, 4). Anterior eyes well-developed, posteriors reduced and twice as smaller as the former. Carapacal setal formula: 4 + 6 + 6 + 8 = 24. One preocular microseta on each carapacal side. Carapace reticulate throughout. Setation of abdominal tergites I–X: 9-11-12-14-1414-13-12-11-11. Male genital area: sternite II with 12 clustered setae medially and posteriorly, forming two barely distinguished groups. Sternite III with 11 anterior, 20 posterior, and four suprastigmal small setae on either side. Sternite IV with no posterior setae and three microsetae along each of the stigma. Sternites V– X with 18-19-17-18-17-14 setae. Female genital area: unknown. Twelfth abdominal segment with two pairs of small setae. Pleural membranes granulostriate. Cheliceral spinneret (galea) (Fig. 5) prominent (ducts not visible). Fixed cheliceral finger with 16 contiguous and small teeth, approximately of the same size. Movable cheliceral finger with six small distal denticles,

New false scorpion species from the Balkan Peninsula

79

Figs 1–8. Neobisium babinzub sp. n., holotype male from Serbia: 1 – pedipalpal chela; 2 – pedipalp; 3 – epistome; 4 – carapace; 5 – chelicera; 6 – leg IV; 7 – flagellum; 8 – male genital area. Scales 0.25 mm (Figs 3, 5, 7, 8) and 0.50 mm (Figs 1, 2, 4, 6).

followed by a median larger tooth (above the level of the galeal seta), and by a series of six small contiguous teeth, which end immediately below the insertion site of the seta gl (diminishing towards the base of the finger) (Fig. 5). Flagellum eight-bladed, characteristic of the genus (Fig. 7). Cheliceral palm with six setae, movable cheliceral finger with one seta only (Fig. 5). Apex of pedipalpal coxa with four or five long and acuminate setae. Pedipalpal segments yellowish-brown, smooth, and as in Figs 1 and 2. Internal excavation of chelal patella shallow and short (Fig. 2). Fixed chelal finger with 38 teeth; the distalmost teeth are triangular and interspaced, followed by some low triangular and finally by some contiguous and flattened teeth (Fig. 1). Movable chelal finger with 28 teeth; distally these are triangular and interspaced. They give way to small and almost triangular and only slightly interspaced teeth proximally. Pedipalpal chela with four and eight trichoboth-

ria on its movable and fixed finger, respectively (Figs 1, 2). Pedipalpal femur 4.12 times as long as wide and 1.45 times longer than carapace. Pedipalpal patella 2.49 times as long as broad, pedipalpal chela 3.44 times longer than broad. Chelal fingers considerably longer than chelal palm. Pedipalpal femur only slightly longer than chelal fingers (Figs 1, 2; Table 1). The ist trichobothrium closer to the finger tip than to ib. Coxa I with a short lateral and median corner bearing small spines. Tibia IV and basitarsus IV with one long sensitive seta each, tarsus IV with two such setae (Fig. 6). Subterminal tarsal setae furcate, each branch with few spinules. Morphometric measurements and ratios are presented in Table 1. Material examined. Holotype: male, E Serbia, Mt. Stara Planina, Babin Zub Peak, 1,600 m a.s.l., from leaflitter (22◦ 50 E, 43◦ 23 N), 13.VII.1994, leg. D. Pavi´cevi´c. The type specimen is deposited in the collection of the Institute of Zoology in Belgrade (IZB 822-1).

´ ´ et al. B.P.M. Curči c

80 Table 1. Linear measurements (in millimeters) and morphometric ratios in Neobisium babinzub sp. n. and N. tarae sp. n. N. babinzub

N. tarae

Character/species

Body Length (1) Cephalothorax Length (2) Width (2a) Ratio 2/2a Abdomen Length Chelicerae Length (3) Width (4) Length of movable finger (5) Ratio 3/5 Ratio 3/4 Pedipalps Length with coxa (6) Ratio 6/1 Length of coxa Length of trochanter Length of femur (7) Width of femur (8) Ratio 7/8 Ratio 7/2 Length of patella (tibia) (9) Width of patella (tibia) (10) Ratio 9/10 Length of chela (11) Width of chela (12) Ratio 11/12 Length of chelal palm (13) Ratio 13/12 Length of chelal finger (14) Ratio 14/13 Leg IV Total length Length of coxa Length of trochanter (15) Width of trochanter (16) Ratio 15/16 Length of femur + patella (17) Width of femur + patella (18) Ratio 17/18 Length of tibia (19) Width of tibia (20) Ratio 19/20 Length of metatarsus (21) Width of metatarsus (22) Ratio 21/22 Length of tarsus (23) Width of tarsus (24) Ratio 23/24 TS ratio – tibia IV TS ratio – metatarsus IV TS ratio – tarsus IV

HM

HM

PM

2.99

3.36

3.085

0.71 0.68 1.04

0.73 0.68 1.07

0.74 0.67 1.10

2.28

2.63

2.435

0.49 0.26 0.315 1.555 1.88

0.52 0.25 0.34 1.53 2.08

0.56 0.275 0.35 1.60 2.04

4.79 1.60 0.69 0.52 1.03 0.25 4.12 1.45 0.76 0.305 2.49 1.79 0.52 3.44 0.79 1.52 1.00 1.265

4.39 4.35 1.31 1.41 0.62 0.66 0.46 0.48 1.01 0.89 0.23 0.23 4.38 3.87 1.38 1.30 0.71 0.74 0.285 0.305 2.49 2.43 1.59 1.58 0.43 0.44 3.70 3.59 0.75 0.77 1.74 1.75 0.835 0.805 1.11 1.045

3.64 0.49 0.45 0.19 2.37 1.03 0.315 3.27 0.77 0.15 5.13 0.40 0.11 3.64 0.50 0.09 5.555 0.39 0.15 0.24

3.21 0.42 0.39 0.16 2.41 0.89 0.25 3.56 0.75 0.13 5.77 0.35 0.10 3.50 0.41 0.08 5.125 0.38 0.15 0.40

3.40 0.45 0.41 0.17 2.44 0.93 0.305 3.05 0.79 0.13 6.08 0.37 0.09 4.11 0.45 0.08 5.625 0.31 0.16 0.34

HM – holotype male, PM – paratype male.

Etymology. After its type locality. Differential diagnosis. From its phenetically close congener, Neobisium fuscimanum (C.L. Koch, 1843), N. babinzub sp. n. is easily distinguished by the form of the epistome (almost absent vs. knob-like), by the pedipalpal femur length to width ratio (4.30 vs. 4.12; Beier

1963) (Table 1), by the chelal palm length to width ratio (1.50–1.60 vs. 1.74–1.75; Beier 1963) (Table 1), by the pedipalpal chelal length to pedipalpal femur length ratio (femur longer than chelal finger vs. femur length almost equal to chelal finger length), by the form of pedipalpal segments (Fig. 10 vs. Fig. 107B; Beier 1963), by the number of setae on tarsus IV (one vs. two), by the presence/absence of a small exterior slit on femur ´ IV (present vs. absent; Curci´ c 1977), and by the general distribution (from the Austrian Alps to the Balkans and Turkey vs. Eastern Serbia). Biology and distribution. This species was collected from beech leaf-litter at an altitude of about 1,600 m a.s.l. under Babin Zub Peak on Mt. Stara Planina (= Mt. Balkan) in E Serbia. It appears that N. babinzub sp. n. is an euedaphic form; it is also suggested that it may represent an endemic form in the highland studied. Neobisium tarae sp. n. (Figs 9–16, Table 1) Description. Anterior carapacal margin slightly convex medially, with a pointed and triangular epistome (Figs 12, 13). Anterior eyes well-developed, posteriors somewhat reduced (and smaller than the anterior eyes; Fig. 13). Carapacal setal formula: 4 + 6 + 6 + 8 = 24 setae (Fig. 13). No preocular microsetae are developed. Carapace reticulate throughout. Setation of abdominal tergites: 6-8-12-13-13-14-1312-12-11 (holotype male), and 6-9-11-13-13-14-13-1313-11 setae (paratype male). Male genital area: sternite II with 8–10 clustered setae medially; sternite III with 21–25 anterior and posterior setae and three or four suprastigmatic microsetae on either side; sternite IV with 13 posterior setae and three small setae along each stigma (Fig. 15). Sternites V–X with 23-22-22-2118-14 (holotype) and 23-22-22-21-17-16 posterior setae (paratype). Female genital area: unknown. Twelfth abdominal segment with 2 + 2 microsetae. Pleural membranes granulostriate. Galea (cheliceral spinneret; Fig. 16) prominent. Fixed cheliceral finger with 15 or 16 contiguous teeth, movable cheliceral finger with 11 or 12 small teeth (Fig. 16). Flagellum eight-bladed, characteristic of the genus (Fig. 16). Cheliceral palm with seven long setae, movable cheliceral finger with one seta. Apex of pedipalpal coxa with five long and acuminate setae. Pedipalpal segments not elongated, but smooth (Figs 9, 10). Pedipalpal femur with two distal and interior tubercles (Fig. 10). Internal excavation of chelal patella shallow and short (Fig. 10). Fixed chelal finger with 63–67 small and close-set teeth; movable chelal finger with 54–60 small and contiguous teeth (Fig. 9). Distally, teeth on both fingers are asymmetrically pointed; these give way to small, apically rounded teeth; on both fingers, teeth are close-set (not interspaced; Fig. 9). Pedipalpal chela with four and eight trichobothria on its movable and fixed finger, respectively (characteristic of the genus; Fig. 9). The ist trichobothrium is

New false scorpion species from the Balkan Peninsula

81

Figs 9–16. Neobisium tarae sp. n., holotype male from Serbia: 9 – pedipalpal chela; 10 – pedipalp; 11 – leg IV; 12 – epistome; 13 – carapace; 14 – flagellum; 15 – male genital area; 16 – chelicera. Scales 0.25 mm (Figs 12, 14–16) and 0.50 mm (Figs 9, 10, 11, 13).

closer to ib than to the finger tip (Fig. 9). Pedipalpal femur 4.17–4.38 times as long as broad; patella 2.43–2.49 times longer than broader; pedipalpal chela 3.59–3.70 times longer than broader. Chelal fingers only slightly longer than chelal palm (Table 1). Coxa I with a small and acute lateral spine. Tibia IV, basitarsus IV, and tarsus IV each with a long sensitive seta (Fig. 11). Subterminal tarsal setae furcate, each branch with few spinules. Morphometric ratios and measurements are presented in Table 1. Material examined. Holotype: male, W Serbia, Mt. Tara, beech forest, from under stones (19◦ 32 E, 43◦ 55 N), 4.VII.1995, leg. S. Peši´c. Paratype: male, same data as for

the holotype. The type specimens are deposited in the collection of the Institute of Zoology in Belgrade (holotype: IZB 441–1; paratype: IZB 441–2).

Etymology. After Mt. Tara, its type locality. Differential diagnosis. From its morphologically similar (phenetically close) ancestor, N. simile (L. Koch, 1873), N. tarae sp. n. differs in many important respects, such as the size of the epistome (small vs. prominent), the form of the galea (inconspicuous vs. prominent), the granulation of the pedipalpal femur [“mediodorsal fest in der ganzen L¨ange mit einer Reihe von Tuberkelchen” (Beier 1963: Fig. 95) vs. only two

´ ´ et al. B.P.M. Curči c

82 laterodistal tubercles; Fig. 10], the pedipalpal femur length [1.10 mm (Beier 1963) vs. 0.92–0.96 mm; Table 1], the pedipalpal chelal palm length [0.85 mm (Beier 1963) vs. 0.75–0.77 mm; Table 1], the chelal finger length [1.15–1.20 mm (Beier 1963) vs. 0.805–0.835 mm; Table 1], the form of the pedipalps [compare Beier (1963; Fig. 95) with Fig. 10 of the present study], and the distribution area (W Europe vs. SE Europe). Biology and distribution. The new species inhabits the slopes of Mt. Tara in W Serbia. To all circumstances, it is an ancient form of the early Tertiary origin. This taxon is probably an endemic and relict species. Conclusion The soil fauna of forests, especially in temperate regions, is primarily mesophilic, just as the two newly described neobisiid pseudoscorpions. Under natural conditions, microhabitat diversity is affected not only by the diversity of organic substrates, but also by the types of climate. In regions with irregular topography, mosaics of moist and dry regions are formed. Apart from these horizontal mosaics, vertical ones (e.g., the type localities of both N. babinzub sp. n. and N. tarae sp. n.) – where different organic horizons differ in moisture content – also exist (especially in highly structured profiles appearing in some mountainous areas). Certain habitats are especially suitable for “preparation” of some pseudoscorpions for their future permanent residence in caves: the soil, the edge of long-lasting snow covers (which occur mainly on high mountains), humus, and mosses (Juberthie et al. 1980). Edaphism or euedaphism (strict adaptation to life in deeper layers of soil), to be more precise, represents the adaptive response of epigean or humicolous species in many faunistic groups (including pseudoscorpions) to survival under conditions of the Mediterranean climate. As already emphasized, such species arose in regions or geological epochs characterized by constant climatic conditions, especially with regard to humidity (Juberthie et al. 1980). A contribution to specific structure of the relict and autochthonous fauna of endogean pseudoscorpions of the Balkan Peninsula, like N. babinzub sp. n. and N. tarae sp. n., was also made by the complex processes involved in development of lacustrine systems, whose continuity was determined by varying duration of marine transgression and regression (Stevanovi´c 1951). It is evident that the majority of epigean and all troglobitic pseudoscorpion species inhabit areas that were formed around the shorelines of former freshwa´ ter basins (Curči´ c 1988b). It can be concluded from this fact that the presence of water (or moisture) is of fundamental significance in development of the phenomenon of edaphism that is the changeover from strict adaptation to life in deep humus and soil layers to a more cryptic way of life (in caves and pits). As far as the origin of N. babinzub sp. n. is concerned, one may suppose that this species presently in-

habits its primordial distribution area; its close relationships with N. fuscimanum supports the conclusion that both species originated from a common ancestor; in either case, it is clear that N. fuscimanum inhabits lower and hilly areas, while N. babinzub sp. n. lives in mountainous habitats (higher than 1,500 m a.s.l.). Furthermore, the two close species, N. tarae sp. n. and N. simile probably originated from a common ancestor. Subsequently, their primordial population was locally exposed more or less to destruction, under the influence of different geotectonic events, climatic changes, or competition with immigrants (Kosswig & Battalgil 1943). It seems that only the westernmost (N. simile) and the easternmost forms have survived; due to a huge disjunction of their present ranges, it is easy to explain their divergent differentiation into separate species. This fact is further supported by a shared plesiomorphic character (presence of unusual tubercles on the pedipalpal femur) (Furon 1959; Deeleman-Reinhold ´ 1978; Curči´ c 1988a, 1998a, b). Acknowledgements We appreciate the help of Dr. S. Peši´c, who collected the specimen of N. tarae sp. n. The financial help of the Serbian Ministry of Science and Environment Protection (Grant No. 143053) is gratefully acknowledged. References Beier M. 1932. Pseudoscorpionidea. I. Subordn. Chthoniinea et Neobisiinea. Das Tierreich 57: 1–258. Beier M. 1963. Ordnung Pseudoscorpionidea (Afterskorpione), Bestimmungsbücher zur Bodenfauna Europas, Lieferung 1, Akademie-Verlag, Berlin, 313 pp. ´ Curči´ c B.P.M. 1977. Uporedno-morfološka obeležja – njihov značaj i primena u klasifikaciji taksona porodice Neobisiidae (Pseudoscorpiones, Arachnida). PhD. Thesis, Fac. Sci., Univ. Belgrade, Belgrade, 186 pp. ´ Curči´ c B.P.M. 1986. On the origin and biogeography of some pseudoscorpions of the Balkan Peninsula. Biologia GalloHellenica 12: 85–92. ´ Curči´ c B.P.M. 1988a. Cave-Dwelling Pseudoscorpions of the Dinaric Karst. Acad. Sci. Art. Slov., Cl. IV, Hist. Nat., Opera 26, Inst. Biol. Ioannis Hadži, 8, Ljubljana, 192 pp. ´ Curči´ c B.P.M. 1988b. Les Pseudoscorpions cavernicoles de la Yougoslavie, developpement historique et implications biogéographiques. Rev. Arachnol. 7: 163–174. ´ Curči´ c B.P.M. 1998a. The cave fauna in Serbia: origin, historical development, and diversification, pp. 75–83. In: Djurovi´c P. (ed.), Speleological atlas of Serbia, Serb. Acad. Sci. Arts, Jovan Cviji´ c Geogr. Inst., Inst. Prot. Nat. Serbia, Fac. Geogr., Univ. Belgrade, Fac. Biol., Univ. Belgrade, Belgrade. ´ Curči´ c B.P.M. 1998b. The hypogean fauna in Serbia: from surface to soil to caves, pp. 59–73. In: Djurovi´c P. (ed.), Speleological atlas of Serbia, Serb. Acad. Sci. Arts, Jovan Cviji´c Geogr. Inst., Inst. Prot. Nat. Serbia, Fac. Geogr., Univ. Belgrade, Fac. Biol., Univ. Belgrade, Belgrade. ´ Curči´ c B.P.M., Dimitrijevi´ c R.N. & Legakis A. 2004. The Pseudoscorpions of Serbia, Montenegro, and the Republic of Macedonia. Monographs 8, Inst. Zool., Fac. Biol., Univ. Belgrade, Hellenic Zool. Soc., Committee for Karst and Speleology, Serb. Acad. Sci. Arts, Inst. Nat. Conserv. Rep. Serbia, Belgrade-Athens, 400 pp. Deeleman-Reinhold C.L. 1978. Revision of the cave-dwelling and related spiders of the genus Troglohyphantes Joseph (Linyphiidae), with special reference to the Yugoslav species. Acad.

New false scorpion species from the Balkan Peninsula Sci. Art. Slov., Cl. IV, Hist. Nat., Opera 23, Inst. Biol. Ioannis Hadži, 6, Ljubljana, 221 pp. Furon R. 1959. La Paléogéographie, Essai sur l’évolution des continents et des oceans. Ed. Payot, Paris, 405 pp. Juberthie C., Delay B. & Bouillon M. 1980. Extension du milieux souterrain en zone non-calcare: description d’un nouveau milieu et de son peuplement par les Coléopt`eres troglobies. Mémoires de Biospéologie 7: 19–52. Kosswig C. & Battalgil F. 1943. Beitr¨ age zur türkischen Faunengeschichte. Comptes Rendus de la Société Turque, Sciences Physiques et Naturelles 8: 18–63. ´ Poinar Jr.G.O., Curči´ c B.P.M. & Cokendolpher J.C. 1998. Arthropod phoresy involving pseudoscorpions in the past and present. Acta Arachnol. 47: 79–96.

83 Schawaller W. 1978. Neue Pseudoscorpione aus dem Baltischen Bernstein der Stuttgarter Bernsteinsammlung (Arachnida: Pseudoscorpionida). Stuttgart. Beitr. Naturkd. 42 B: 1–22. Schawaller W. 1980. Bibliographie der rezenten und fossilen Pseudoscorpionidea (Arachnida) 1890–1979. Stuttgart. Beitr. Naturkd. 338 A: 1–61. Stevanovi´c P. 1951. Donji pliocen Srbije i susednih oblasti. Posebna izdanja, Srpska Akademija Nauka i Umetnosti, Geografski Institut, 2, Beograd, 361 pp. Received February 8, 2006 Accepted May 9, 2006