a review of the spider genus boagrius simon, 1893 (araneae

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RAFFLES BULLETIN OF ZOOLOGY 2020

A review of the spider genus Boagrius Simon, 1893 (Araneae: Palpimanidae)

Sergei L. Zonstein1* & Yuri M. Marusik2,3

Abstract. The chedimine genus Boagrius Simon, 1893 (including only two species), is reviewed. The genus and its type species B. pumilus Simon, 1893, are rediagnosed and redescribed; the male of this species is described for the first time. The range of the genus is reconsidered as restricted only to South-Eastern Asia. Based on the structure of both the male palp and somatic characters, East African B. incisus Tullgren, 1910, is transferred to Scelidocteus Simon, 1907, as S. incisus (Tullgren, 1910), new combination. Additionally, B. simoni, new species, is described from the north-eastern part of Borneo.

Key words. Aranei, Chediminae, Scelidocteus, Singapore, Malaysia, Malay Peninsula, Borneo, new combination, new species

RAFFLES BULLETIN OF ZOOLOGY 68: 91–102Date of publication: 1 April 2020DOI: 10.26107/RBZ-2020-0010http://zoobank.org/urn:lsid:zoobank.org:pub:6533FC4C-8451-4309-BA10-C0502C26D2EB

© National University of SingaporeISSN 2345-7600 (electronic) | ISSN 0217-2445 (print)

Accepted by: Ang Yuchen1Steinhardt Museum of Natural History, Tel-Aviv University, 69978 Tel-Aviv, Israel; Email: [email protected] (*corresponding author)2Institute for Biological Problems of the North RAS, Portovaya Str. 18, Magadan, Russia3Department of Zoology & Entomology, University of the Free State, Bloemfontein 9300, South Africa

INTRODUCTION

Simon (1893a, b) described Boagrius as a monotypic genus with the type species B. pumilus Simon, 1893, from the Malay Peninsula. Tullgren (1910) and Berland (1920) added to Boagrius two supplementary species from Eastern Africa: B. incisus Tullgren, 1910, and B. kilimanjari Berland, 1920, respectively. Several decades after, Benoit (1974) transferred the latter species to Sarascelis Simon, 1887. Thus, this genus is considered currently to include only two species (World Spider Catalog, 2019). However, this genus has never been revised in its entirety. Males of B. pumilus were not described and their characters remained unknown until now. The structure of the male palp was depicted only for a putatively undescribed congener from South-Eastern Asia (Deeleman-Reinold, 2001: figs. 76, 77). The structure of the female copulatory organs in representatives of Boagrius remains totally unknown.

The present study, intending to fill these gaps, is based on the occurrence of a few specimens belonging to Boagrius in the collections of several European museums (listed below). After examination, some of these spiders were identified as representing B. pumilus (including one adult female and a formerly undescribed conspecific male), while the rest of the

material (also containing both sexes) was assigned to a new species described below. Additionally, an examination of the type series of B. incisus revealed this species belonging to another chedimine genus, Scelidocteus. Thus, the goal of the current study is to delimit, rediagnose and redescribe Boagrius, as well as to describe/redescribe the genus members in detail, and to provide the necessary taxonomical and nomenclatural changes for an excluded species.

MATERIAL AND METHODS

Specimens from the following spider collections were studied: the Manchester Museum, the University of Manchester (MMUM; UK); Museum für Naturkunde (MNB; Berlin, Germany); the Oxford University Museum of Natural History (OUMNH; UK); the Senckenberg Research Institute and Museum (SMF; Frankfurt-am-Main, Germany); Naturhistoriska riksmuseet (NRS; Stockholm, Sweden); Zoological Museum at the University of Turku (ZMUT; Finland).

Photographs were taken using an Olympus SZX16 stereomicroscope with a Canon-7D camera and prepared using Zerene Stacking software. Scanning electron micrographs were made using the SEM JEOL JSM-5200 scanning microscope at the Zoological Museum, University of Turku, Finland. Illustrations of scuta and endogynes were made after maceration in a 20% potassium hydroxide aqueous solution and exposure for a few minutes in an alcohol/water solution of Chlorazol Black. Endogynes were photographed on slides with either an Olympus SZX16 or an Olympus BH-2. Small pieces of cotton were used to fix the copulatory organs in the correct position. Background maps were taken from an open free source, SimpleMappr (Shorthouse, 2010).

Taxonomy & Systematics

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Zonstein & Marusik: A review of Boagrius

Measurements were made to an accuracy of 0.01 mm. Lengths of leg and palp segments were measured on the dorsal side, from the midpoint of the anterior margin to the midpoint of the posterior margin. The sequence of the segment measurements is given for legs I–IV as follows: total length (femur + patella + tibia + metatarsus + tarsus), while for palp it is presented in the following order: total length (femur + patella + tibia + tarsus/cymbium). All measurements are given in millimetres.

The following abbreviations (except for some of them, encoded in the text) are used in the article: ALE, anterior lateral eyes; AME, anterior median eyes; CH, carapace height; CL, carapace length; CW, clypeus width; CyL, clypeus length; PLE, posterior lateral eyes; PME, posterior median eyes; TL, total length of body in dorsal view.

TAXONOMY

Family Palpimanidae Thorell, 1870

Subfamily Chediminae Simon, 1893

Boagrius Simon, 1893

Boagrius Simon, 1893a: 405; Deeleman-Reinhold, 2001: 52, figs. 76, 77; Jocqué & Dippenaar-Schoeman, 2006: 196, fig. 75b, c.

Type species. Boagrius pumilus Simon, 1893, by monotypy.

Diagnosis. By possessing a similar state of some characters (a bipartite thoracic fovea, relatively long and slender legs I–IV, a weakly swollen femur I, and a poorly developed leg scopula), the considered genus resembles members of Levymanus Zonstein & Marusik, 2013, distributed in the eastern part of Africa and in the Near East (Figs. 1B, 1J, 2A–H cf. Zonstein & Marusik, 2013: figs. 1, 2, 10, 11, 26–31). All other chedimine genera, except for those discussed here, embrace species possessing a pit-like or a slit-like thoracic fovea, a stouter femur I, and a better developed leg scopula (see Zonstein & Marusik, 2013: figs. 1–9; 2017a: figs. 3–8, 11–15, 19, 40–42; 2017b: fig. 1A–E; Zonstein et al., 2016: figs. 1–6; 2018: figs. 1–4; Marusik & Zonstein, 2018: figs. 1–5, 12, 13).

Boagrius can be clearly distinguished from Levymanus by: a) having a more compact cephalothorax with shorter and narrowed anterior and posterior parts; b) the presence of an elevated hump in the median part of the carapace; c) the relatively smaller AMEs (which are two to three times as large as other eyes); d) a shorter distance between the thoracic sulci; and e) a shorter abdomen with a considerably shorter petiolar tube (Figs. 1A, B, 2A–H, 3A–C, 3E–L). In species of Levymanus, the carapace is longer and narrower anteriorly and posteriorly, the thoracic hump is not as conspicuous, the AMEs four to five times as large as other eyes, the distance between the thoracic sulci is noticeably broader, and the abdomen is longer with an unusually long

petiolar tube (cf. Zonstein & Marusik, 2013: figs. 10–15, 33–34; Zonstein et al., 2017: fig. 1A–F).

Description. Small bicolour chedimine palpimanids with body length 2.1–2.4 mm in males and 2.6–2.8 mm in females. Carapace domed and oval, slightly narrowed and truncated anteriorly and posteriorly, densely and finely corrugated (Fig. 1A), with elevated hump between eye group and thoracic fovea. Eight eyes. AME largest, about two to three times larger than other eyes, which are subaequal in size. ALE and PLE almost touching each other. PME widely spaced from each other, as well as from AMEs and from PLEs. Clypeus about two times higher than AME diameter. Chilum inconspicuous. Thoracic fovea short and bipartite, with two separate sulci closely located side by side (Fig. 1B). Posterior notch deep subtriangular (Fig. 1C). Chelicerae relatively small, only slightly longer than clypeus; stridulatory ridges absent; cheliceral fang serrate; cheliceral furrow without true or peg teeth (Fig. 1D–F). Sternum shield-like, widest between coxae I and II, with fine reticulation; labium about as broad at base as it is long. Maxillae short and moderately wide (Fig. 1G, H).

Palps moderately short, with thickened distal segments in females (Fig. 1I); legs I–IV long and slender. Leg formula: 1423. Leg cuticula fairly smooth. Femur I rather long and weakly swollen; patella I thin and long (longer than tibia). Tibia and metatarsus I with weakly developed prolateral scopula (Fig. 1J). Leg tarsi straight (not curved), with two narrow unidentate claws and ascopulate (Fig. 1K, L). Claw tufts weakly developed.

Abdomen ovoidal, slightly extended anteriorly and obtuse posteriorly, without dorsal pattern (uniformly pale coloured) in unsclerotised part. Abdominal scuta conforming a rather short pedicel tube; dorsal portion of scutum relatively small. Small spinneret group set on low mound. AMS small, cylindrical, two-segmented; PMS and PLS reduced to a few sessile spigots in females and absent in males.

Male palp (Figs. 3C, D, 4A–I, 5A–L). Femur longer than wide (ca. 2.3), equal in length to tibia and cymbium; patella more than two times shorter than femur; tibia strongly swollen, ca 1.6 thicker than femur; cymbium and bulb not embedded into the tibia; cymbium with sparse ‘scopular’ setae; bulb longer than cymbium, with two outgrowths, the tegular apophysis (Ta) and embolus (Em); tegular apophysis long (subequal to the length of tegulum), straight, with bent and bifurcated tip; embolus well sclerotised, relatively long (equal or ca. 1.5 times shorter than tegular apophysis), partly screwed (bent around the axis), with thin apophysis (Ea).

Female copulatory organ (Fig. 6A–F). Posterior edge of epigastric scutum bow-like, shallowly concave in median part, posterior edge of epigastric furrow (postgastric area) with two or three thin scuta, two lateral and one median (not distinct in B. pumilis), all these scuta almost invisible maceration. Endogynes differ in two species, wider than long (ca. 3 times in the type species and ca. 1.5 in the new

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Fig. 1. Scaled SEM micrographs of somatic structures of Boagrius pumilus (subadult female from Bukit Fraser, Malaysia). A, carapace, anterior view; B, thoracic fovea, dorsal; C, posterior notch of carapace; D–F, chelicera, anteroventral, ventral, and posteroventral view, respectively; G, H, palpal endites, ventral; I, palp, prolateral; J, leg I, tibia to tarsus, prolateral; K, apex of tarsus I, showing tarsal claws and tufts, anterior view; L, same, leg IV, retrolateral view.

species); each receptacle (Rl + Rt) with three grape-like glands (Gg); receptacles in type species with pore gland field (Pg) anteriorly.

Composition. Boagrius currently includes two named species: B. pumilus Simon, 1893, and B. simoni, new species.

Distribution. South-East Asia: Singapore and Malaysia (mainland part and Borneo).

Boagrius pumilus Simon, 1893(Figs. 1A–L, 2A–D, 3A–F, 4A–D, 5A–F, 6A, B, 8)

Boagrius pumilus Simon, 1893a: 405; 1893b: 211 (female); Murphy & Murphy, 2000: 240, fam. 45 fig. 1; Murphy & Roberts, 2015: 140, pl. 238 (female).

Holotype. Female, SINGAPORE, no other data (MNHN; borrowed to date by another museum, thus not examined).

Material examined. SINGAPORE, 1 male, 1 female, 1 juvenile, Bukit Timah, 100 m, 20–25 February 1988, coll.

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Fig. 2. A–D, habitus of Boagrius pumilus; E–H, habitus of B. simoni, new species; A, B, E, F, males; C, D, G, H, females; A, C, E, G, dorsal view; B, D, F, H, lateral view. Scale bar = 1 mm.

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Fig. 3. A–F, cephalothorax of Boagrius pumilus; G–I, cephalothorax of B. simoni, new species; A–D, G–I, males; E, F, J, K, females; A, E, G, J, dorsal view; B, F, H, K, ventral view; C, D, I, anterior part of cephalothorax in dorsal, ventral, and anterior view, respectively. Scale bar = 0.25 mm.

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Fig. 4. A–D, details of male palp of Boagrius pumilus; E–I, details of male palp of B. simoni, new species; A, E, entire palp, retrolateral; B, F, same, prolateral; C, D, G, distal part of palp, ventral; H, I, same, proventral. Scale bar = 0.25 mm.

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F. Murphy & J. Murphy (MMUM G7572.3055); 1 subadult female, same collecting data but 15 August 1992 (MMUM G7572.3112); 1 juvenile, Upper Pierce Reservoir, in litter, 19 February 1988, coll. F. Murphy & J. Murphy (MMUM G7572.3258). MALAYSIA, 1 male, Pahang State, Bukit Fraser, wet moss and litter, 18 November 1984, coll. P. Lehtinen (ZMUT); 1 subadult female, same collection data, 1,420 m, roadside ferns and bush, 14 March 1998, coll. P. Lehtinen (ZMUT); 1 female, same locality, Jeriau Waterfall, 3°43′26.07″N, 101°42′36.22″E, 1,040 m, disturbed primary forest, leaf litter (sieving), 18 June 2013, coll. P. Jäger (SMF).

Diagnosis. The species can be distinguished from B. simoni, new species, by the lighter carapace (scarlet red vs. carmine red) and by the structure of the male and female copulatory organs: a terminal part of the tegular apophysis with a fairly developed bifurcation (Figs. 3C, D, 4C, D) vs. a shallow, almost indistinct depression in B. simoni, new species (Fig. 4G–I); the endogyne is ca. 3 times wider than long and the pore glands are located anteriorly (Fig. 5A, B), while in B. simoni, new species, the endogyne is about 1.5 times wider than long and the pore gland is located posteriorly.

Description. Male (from Bukit Frazer, ZMUT). Habitus as in Fig. 2A, B. Colour in alcohol: carapace, sternum, labium, chelicerae, maxillae, and dorsal abdominal scutum bright scarlet red; coxae and femora I–IV, and entire palp light orange; other segments of legs I–IV, and unsclerotised part of abdomen including spinnerets milky white.

Measurements: TL 2.36, CL 1.14, CW 0.87, CH 0.64, CyL 0.20, femur I L/W 3.44 (0.93/0.27). Carapace with moderately coarse granulations (Fig. 3A). Eye sizes and interdistances: AME 0.14, ALE 0.05, PLE 0.04, PME 0.05, AME–AME 0.04, AME–ALE 0.04, AME–PME 0.09, PLE–PME 0.11, PME–PME 0.14.

Palp and leg measurements: palp 0.99 (0.34 + 0.12 + 0.24 + 0.29), leg I 2.91 (0.93 + 0.64 + 0.60 + 0.31 + 0.43), leg II 2.51 (0.76 + 0.43 + 0.50 + 0.43 + 0.39), leg III 2.34 (0.67 + 0.39 + 0.46 + 0.46 + 0.36), leg IV 3.15 (0.94 + 0.43 + 0.71 + 0.64 + 0.43).

Palp (Figs. 3C, D, 4A–D, 5A–F). Tegular apophysis (Ta) long (subequal to the length of tegulum), straight, with bent and bifurcated tip; embolus (Em) well sclerotised, relatively long (ca. 1.5 times shorter than tegular apophysis), partly screwed (bent around the axis), with thin apophysis (Ea).

Female (from Bukit Frazer, SMF). Habitus as in Fig. 2C, D. Colour in alcohol as in male.

Measurements: TL 2.65, CL 1.21, CW 0.91, CH 0.6, CyL 0.21, femur I L/W 2.97 (1.07/0.36). Carapace: longer, with less coarse granulations than in male (Fig. 3E). Eye sizes and interdistances: AME 0.11, ALE 0.06, PLE 0.04, PME 0.06, AME–AME 0.06, AME–ALE 0.06, AME–PME 0.06, PLE–PME 0.11, PME–PME 0.13.


Copulatory organ (Fig. 6A, B). Posterior edge of epigastric furrow (postgastric area) with a pair of lateral (Ls) scuta, median scutum indistinct, lateral scuta almost invisible after maceration; endogyne almost three times wider than long; atrium (At) large; lateral parts (receptacles?) suboval, well sclerotised, receptacles seem fused anteriorly; anterior part with pore-gland field (Pg); grape-shaped glands very large, about 1/6 of endogyne width.

Note. Endogyne was lost during preparation of the figure.

Distribution. South part of the Malay Peninsula: Singapore and Malaysia (Pahang State). See Fig. 8.

Boagrius simoni, new species(Figs. 2E–H, 3G–K, 4E–I, 5G–L, 6C–F, 8)

Holotype. Male, MALAYSIA, Sabah State (Borneo Island), Lahad Datu District, Ulu Segama Forest Reserve, 4°57.9′N, 117°48.1′E, 200 m, pitfall trap, 1–2 November 2005, coll. D.J. Mann, E.M. Slade & J.F. Villanueva (OUMNH-2006-051a).

Paratypes. 1 female, collected together with the holotype, coll. E.M. Slade & J.F. Villanueva (OUMNH-2006-051b); 1 male, same collecting data but 24 November 2005, coll. D.J. Mann (OUMNH-2006-051c); 1 male, same collecting data but 4°57.7′N, 117°50.2′E, 250 m, pitfall trap, March 2005, coll. E.M. Slade & J.F. Villanueva (OUMNH-2005-062).

Etymology. The specific name is a patronym in honour of Dr. Eugene Simon (1848–1924), the outstanding French arachnologist who made an immense contribution to the study of spiders and described several thousand spider taxa, including the genus Boagrius and its type species.

Diagnosis. The species can be distinguished from B. pumilus by a darker carapace (carmine red vs. scarlet red in the latter species) and by the structure of the male and female copulatory organs: the tegular apophysis with a shallow bifurcation vs. a stronger developed bifurcation; the endogyne about 1.5 times wider than long, and the grape-shaped glands being relatively small (1/7 of the endogyne width) vs. the endogyne which is three times wider than long with larger glands (ca. 1/6 of the endogyne width).

Description. Male holotype (OUMNH-2006-051a). Habitus as in Fig. 2E, F. Colour in alcohol: carapace, sternum, labium, chelicerae, maxillae, and dorsal abdominal scutum carmine red; coxae and femora I–IV, and entire palp light orange; other segments of legs I–IV, and unsclerotised part of abdomen including spinnerets pale brownish white.

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Fig. 5. Scaled SEM micrographs. A–F, male palp and copulatory organs of Boagrius pumilus; G–L, male palp and copulatory organs of B. simoni, new species; A, G, entire palp, prolateral and retrolateral view, respectively; B, distal palpal sclerites, retrolateral; C, H, K, same, ventral; D, J, same, prolateral; E, F, I–L, same, anterior view. Abbreviations: Ea, thin apophysis; Em, embolus; Ta, tegular apophysis.

Measurements: TL 2.10, CL 1.10, CW 0.83, CH 0.27, CyL 0.19, femur I L/W 3.10 (0.90/0.29). Carapace: with coarse granulations (Fig. 3G). Eye sizes and interdistances: AME 0.14, ALE 0.05, PL 0.05, PM 0.04; AME–AME 0.04, AME–ALE 0.06, AME–PME 0.04, PLE–PME 0.10, PLE–PLE 0.14.


Palp (Figs. 4E–I, 5G–L). Tegular apophysis with shallow bifurcation. Embolus (Em) almost as long as tegular apophysis, distinctly screwed, with complex tip.

Female paratype (OUMNH-2006-051b). Habitus as in Fig. 2G, H. Colour in alcohol as in male, but entire body and legs I coloured slightly darker.

Measurements: TL 2.78, CL 1.29, CW 0.93, CH 0.64, CyL 0.22, femur I L/W 2.82 (0.96/0.34). Carapace: longer, with slightly less developed granulations than in male (Fig. 3K). Eye sizes and interdistances: AME 0.10, ALE 0.06, PLE 0.06,

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Fig. 6. A, B, female copulatory organs of Boagrius pumilus; C–F, female copulatory organs of B. simoni, new species; A, endogyne, receptacles, and accessory structures, ventral view; B, D, same, posterior view; C, E, F, same, dorsal view. Scale bar = 0.25 mm. Abbreviations: At, atrium; Ft, fine thread; Gg, grape-shaped gland; Pg, pore gland; Rl, Rt, longitudinal and transversal parts of receptacle, respectively.

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Fig. 7. A–D, habitus and structures of Scelidocteus incisus (Tullgren, 1910), new combination, male lectotype; E–G, habitus and structures of Scelidocteus incisus (Tullgren, 1910), new combination, female paralectotype; A, E, habitus in dorsal view; B, F, cephalothorax, dorsal; C, D, distal sclerites of palp, retrolateral and ventral view, respectively; G, cephalothorax in ventrolateral view. Scale bar = 1 mm (A, E); 0.5 mm (B, F, G); 0.1 mm (C, D).

PME 0.04, AME–AME 0.06, AME–ALE 0.06, AME–PME 0.07, PLE–PME 0.13, PME–PME 0.14.


Copulatory organs (Fig. 6C–F). Postgaster with three sclerites, two lateral and one median. Endogyne about 1.5 times wider than long; receptacles long, bent and with horizontal (transversal, Rt) and vertical (longitudinal, Rl) parts, horizontal part about 1.5 times shorter than vertical, well sclerotised in comparison to membranous vertical part; grape-shaped glands (Gg) and fine threads (Ft) originate

from anterior part of receptacle; grape-shaped glands with long stem, as long as horizontal part of receptacle, head large, their diameter subequal to diameter of horizontal part of receptacle or 1/7 of endogyne width; pore glands (Pg) seem present on horizontal part of receptacle.

Distribution. Known only from the type locality (Fig. 8).

MISPLACED SPECIES

Scelidocteus incisus (Tullgren, 1910), new combination(Fig. 7A–G)

Boagrius incisus Tullgren, 1910: 112, pl. 1 fig. 28a–g (male, female).

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Fig. 8. ●: distribution of Boagrius pumilus. ▲: distribution of B. simoni, new species. Open circle: type locality listed in the text. Solid symbols: new records listed in the text.

Lectotype. Male (designated here), TANZANIA: SW Mt Kilimanjaro, Kibongoto (“Kibongo”, as labelled; 3°11′S, 37°06′E), 1,000–1,200 m, June 1905, Y. Sjöstedt (RMS; examined).

Paralectotype. 1 female (designated here), same collecting data (MNB 13846; examined).

Notes. This species, originally placed in Boagrius, possesses many characters which are dissimilar to those observed in B. pumilus and B. simoni, new species. Tullgren (1910) mentioned the type series of B. incisus consisting of seven adult and non-adult females, and one adult male. However, only one adult female and the above-noted male have been located. Although Tullgren (1910) described one of the syntype females first, we prefer to choose the conspecific syntype male to designate as the lectotype. This male was obviously used in the original description, while the same cannot be claimed regarding the only available female. The examination of both these type specimens revealed that this East African taxon, when compared with the South-Eastern Asian Boagrius spp., differs from them by: a) a considerably larger body size (TL 6.9–7.8 vs. 2.1–2.8 mm); b) the presence of a slit-like (not a bipartite) thoracic fovea; c) femur I considerably swollen in both sexes (vs. an almost unmodified one); d) corrugated (not smooth) segments of leg I, at least in the male; e) dense leg scopula (vs. a poorly developed one); and f) the configuration of the male copulatory organs possessing accessory structures twisting around each other (see Fig. 7A–G). According to the listed characters, this African species can be considered as belonging to the purely African chedimine genus Scelidocteus Simon, 1907. Hence, we place this species in the latter genus as S. incisus (Tullgren, 1910), new combination. Its further position will be specified in the course of a planned revision of Scelidocteus.

ACKNOWLEDGEMENTS

We thank Zoe Simmons (OUMNH), Dmitri Logunov (MMUM), Peter Jäger and Julia Altmann (SMF), Jason Dunlop (MNB), Seppo Koponen (ZMUT), and Torbjörn Kronestedt (NRS) for providing us with the material used for this study. Special thanks go to Ilari Sääksjärvi and Seppo Koponen (ZMUT) for providing us with museum facilities. English of the final draft was edited by Sarah Crews (Oakland, California, USA). We also appreciate the reviewers for their constructive comments. This study was supported in part by the Ministry of Absorption, Israel.

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Zonstein SL, Marusik YM & Omelko MM (2016) Redescription of the type species of Diaphorocellus Simon, 1893 (Araneae, Palpimanidae, Chediminae). African Invertebrates, 57: 93–103.

Zonstein SL, Marusik YM & Omelko MM (2018) Redescription of the monotypic genus Scelidomachus Pocock, 1899 (Aranei: Palpimanidae) and its type species. Arthropoda Selecta, 27: 53–56.

a review of the spider genus boagrius simon, 1893 (araneae

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