Subfamily Ylleninae

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APPENDICES -AP1-Simon's classification - AP2-Petrunkevich synthesis-1928 - AP3-Bonnet's list of subfamilies - AP4-Prószynski's revision of subfamilies 1976 - AP5-Prószynski - summary of results - AP6-Aelurillus black & gray - AP7-Maddison's views on Salticidae phylogeny 2014 - MS1-Omoedus synonymy
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Monograph of the Salticidae (Araneae) of the World 1995-2015. Part I: Introduction to alternative classification of Salticidae.

Jerzy Prószyński
38 - YLLENINES GROUP OF GENERA
Version July 1st, 2016.
Sources and permissions of illustrations, also references, are displayed in the IInd part of the monograph http://www.peckhamia.com/salticidae/

Index. Type genus Yllenus. Logunyllus Marusyllus Yllenus. (62 species)

Introduction

Representative species: Yllenus arenarius Simon*, 1868b.
Diagnosis. The group has relatively uniform habitus and natural history, consists of three genera (containing at present 77 species) previously classified as a single genus Yllenus. It does not resemble any other genera of Salticidae, from which they differ by somatic characters, male palps, epigyne and internal structures of epigyne. All species have scopula - brush of setae stretching from claws onto ventral surface of tarsus I. Chelicerae slender, without developed posterior median edge, there are no cheliceral teeth, except minute tip of sclerotized anterior median edge, protruding near fang in some species. Size of carapace and abdomen are comparable, color pattern camouflaging in sandy environment. Carapace highest at eyes III, from where thorax is gently sloping, there is no flattened thoracal part. Main character of palps is presence of large, often enormous conductor, splitting off the basis of embolus, embolus is hair like, details of these structures are developed differently in particular genera. Palpal femur broad, crescent shaped, often with protuberance. Epigyne with prominent median pocket, spermathecae and copulatory ducts either heavily sclerotized or, if walls of ducts thinner, then form of complicated coils, in some species even double spiral. Combinations of these characters is unknown in other groups of genera, including ground dwelling AELURILLINES.
Remarks. All species of YLLENINES dwelt on sands, in Asia, Africa and Europe (South and Central), protected by camouflage coloration, hiding under surface of sand, where they make nests, retreats and produce cocoons. Their natural history was studied in details by Bartos 2005 on Yllenus arenarius Simon, 1868, and was confirmed by sundry observations on many other species. The structure of this taxon fit definition of subfamily, with several mutual characters, including appearance and biology. Male palps, epigyne and internal structure of epigyne delimit three groups of species, here described as three genera, closely related to each other but constituting separate lines.
Remarks. Splitting off embolus from conductor in Yllenus, both arising from the mutual base, is striking in comparison with Pellenes lapponicus, as demonstrated in SEM of that species (Figure 167). I hypothesize that it represents ancestral state
* Footnote. The authorship of Yllenus arenarius is written differently in various publications, usually "Menge in Simon, 1868", including Bonnet 1959: 4904 (although he sacrificed half a page to prove that Menge himself did not use name Yllenus and described his species as Marpesia areniclor only in 1877). World Spider Catalog (version 2016) also use that authors' combination in citation of paper, but "abbreviates" authorship of species to nonexistent paper "Menge 1868". Radical proposal to limit authors quotation to authors' of publication only, illustrated by example of this very species, ( http://www.peckhamia.com/salticidae/introduction/I_About_this_database.html#different") was overlooked. I may add that the only author quoting correctly Yllenus arenarius Simon, 1868 was Roewer 1954: 1252.

Figures 164-171. Diagnostic characters of informal group of genera YLLENINES. 164-165 - enormous conductor in Yllenus, (here represented by Y. zyuzini Logunov & Marusik, 2003), lateral and ventral views, 166 - forked embolus and conductor, reduced in size, in (Logunyllus saliens (O. Pickard-Cambridge, 1876) comb. n., 167 - for comparison: conductor reduced to a sheath hiding hair-like embolus, common in many Salticidae genera (here in Pellenes lapponicus -SEM of dissected bulbus and embolus, dorsal view), 168 - conductor and embolus in Marusyllus hamifer Simon, 1895, 169 - scopula on tarsus I in M. coreanus Prószynski, 1968, 170 - body shape and color pattern in Logunyllus salsicola (Simon, 1937), comb. n., 171 - cheliceral dentition in L. vittatus (Thorell, 1875) comb. n. + Yllenus arenarius (Simon, 1868)- epigyne, spermathecae and ducts. 164-167 - from Logunov & Marusik 2003, ©Atrhropoda selecta, 2003; 168-172 - from Prószyński 1968, ©Annales zoologici. By courtesy.

Figures 172-174. Logunyllus univittatus (Simon, 1871) , comb. n., © Photo J. Lissner -172, Logunyllus vittatus (Thorell, 1875), comb. n., male and female, © Photo J. Nigl -173-174. Both by courtesy.

Guide to identification of genera by palps and internal structure of epigyne

Guide to identification of genera by epigyne

Guide to identification of genera by color pattern

Gen. Logunyllus Prószyński, 2016 gen. n. (35 species)
[partial synonym for the genus Yllenus (Yllenus albocinctus group Prószynski, 1968)].

Yllenus (albocinctus group) Prószynski, 1968e: 455,
Yllenus (albocinctus group) Logunov, Marusik 2003b: 23.
Type species Yllenus albocinctus Simon, 1876
Etymology. Named in honor of prominent arachnologist - Dr. D.V. Logunov [LOGUN-ov-YLL-US], author of a large number of publications on Salticidae, including big monograph of the genus Yllenus. Grammar gender - masculine.

Diagnosis. General characters shared with remaining genera of YLLENINES - Marusyllus Prószyński, 2016 gen. n. (see below) and Yllenus Simon, 1868, but palps and epigyne are "normal" - that is male cymbium and embolus are relatively short, conductor is slender, while bulbus is relatively large. The articulating apparatus consists of complicated processes on cymbium and tibia. The epigyne of female is relatively strongly sclerotized, the copulatory canals are straight, or form simple loops, the latter often consist of distal - stronger sclerotized, and proximal - less sclerotized parts. Drawings below constitute integral part of the genus definition.
Remarks. The description of this genus upgrades, without other changes, previously described group of species Yllenus (albocinctus group) (Prószynski, 1968: 455, Logunov, Marusik 2003b: 23).

The following species are transferred from Yllenus Simon 1868 to Logunyllus gen. nov.: Logunyllus albifrons-1 (Lucas, 1846), comb. n., L. albifrons-2 (Caporiacco, 1933), comb. n.,, L. albifrons-3 (Logunov &, Marusik, 2003)[= L. & M., 2003], comb. n., L. albocinctus (Kroneberg, 1875), comb. n., L. algarvensis (L. & M., 2003), comb. n., L. arabicus (Prószynski, 1993), comb. n., L. auriceps (Denis, 1966), comb. n., L. bactrianus (Andreeva, 1976), comb. n., L. bakanas (L. & M., 2003), comb. n., L. bucharaensis (L. & M., 2003), comb. n., L. caspicus (Ponomarev, 1978), comb. n., L. dalaensis (L. & M., 2003), comb. n., L. gavdos (Logunov, Marusik, 2003), comb. n., L. guseinovi (L. & M., 2003), comb. n., L. halugim (L. & M., 2003), comb. n., L. improcerus (Wesolowska, van Harten, 1994), comb. n., L. knappi (Wesolowska, van Harten, 1994), comb. n., L. logunovi (Wesolowska, van Harten, 2010), comb. n., L. mirabilis (L. & M., 2003), comb. n., L. mirandus (Wesolowska, 1996), comb. n., L. nigritarsis (L. & M., 2003),, comb. n., L. nurataus (L. & M., 2003), comb. n., L. pavlenkoae (L. & M., 2003), comb. n., L. pseudovalidus L. & M., 2003), comb. n., L. ranunculus (Thorell, 1875), comb. n., L. saliens (Pickard-Cambridge O., 1876), comb. n., L. salsicola (Simon, 1937), comb. n., L. shakhsenem (L. & M., 2003), comb. n., L. somonensis (Prószynski, 1982), comb. n., L. squamifer (Simon, 1881), comb. n., L. tamdybulak (L. & M., 2003), comb. n., L. tschoni (Caporiacco, 1936), comb. n., L. univittatus (Simon, 1871), comb. n., L. validus (Simon, 1889), comb. n., L. vittatus (Thorell, 1875), comb. n., L. zaraensis (Logunov, 2009), comb. n., L. zhilgaensis (L. & M., 2003), comb. n.

LOGUNYLLUS syn. Yllenus (albocinctus group) albocinctus : Proszynski J. 1968e. Ann. zool. 26: 463-466, f 25, 38, 52, 66, 136-143 + Proszynski 1982. from . Ann. hist. nat. Mus. hung., Budapest, 74: 290, f 48. By courtesy. .

LOGUNYLLUS syn.Yllenus (albocinctus group) albifrons +b) ranunculus: Proszynski 1968e. Ann. zool. 26: 463-466, f 25, 38, 52, 66, 136-143 + Punda H. 1975. Ann. zool. 33: 35-36, f 1-2 (sp?-Libya near Gialo) + Prószyński 2008 - Internet (comparison) +b) Punda 1975. Ann. zool., 33: 37, f 5. By courtesy.

LOGUNYLLUS syn. Yllenus (albocinctus group) saliens : Proszynski 1968e. Ann. zool. 26: 476-479, f 8, 26, 42, 73, 161-167 + Punda 1975. Ann. zool. 33: 36-37, f 3-4 + Logunov, Marusik 2003b.: 63-66, f 44, 54, 220-234 - comparison. By courtesy.

LOGUNYLLUS syn. Yllenus (albocinctus group) salsicola : Proszynski 1968e. Ann. zool. 26: 455-456, f 121-123, 125 + 2003: 174, f 703-708.

LOGUNYLLUS syn. Yllenus (albocinctus group) squamifer : Proszynski 1968e. Ann. zool. 26: 460-463, f 129-135. By courtesy.

LOGUNYLLUS syn. Yllenus (albocinctus group) shakhsenem +b) tamdybulak +c) zhilgaensis : Logunov, Marusik 2003b.: 68-69, f 153-156 +b) 71-72, f 213-215 +c) 83-85, f 292-299. By courtesy.

LOGUNYLLUS syn. Yllenus (alb. group) univittatus : Proszynski 1968e. Ann. zool. 26: 467-470, f 7, 16, 31, 40, 53, 68, 144-150+ ©Photo Lissner.

LOGUNYLLUS syn. Yllenus (albocinctus group) validus + (=?) somonensis : Proszynski 1968e. Ann. zool. 26: 470-472, f 28, 41, 151-154 + + Logunov, Marusik 2003b.: 77-79, f 82, 272-284 - comparison+b) Proszynski 1982. Ann. hist. nat. Mus. hung., 74: 292-293, f 50-52. . By courtesy.

LOGUNYLLUS syn. Yllenus (alb. group) vittatus : Proszynski 1968e. Ann. zool. 26: 472-476, f 27, 39, 54, 67, 75, 155-160
+ ©Photo M. Freudenschuss + J. Nigl. By courtesy.

LOGUNYLLUS syn. Yllenus (alb. group) algarvensis +b) auriceps + c) bakanas +d) bucharaensis : Logunov, Marusik 2003b.: 30-31, f 110-114 +b) 156, f 501-503 +c)33-35, f 121-127. By courtesy.

LOGUNYLLUS syn. Yllenus (alb. group) caspicus +b) dalaensis + c) gavdos +d) guseinovi +e) mirabilis : Logunov, Marusik 2003b.: 37-40, f 128-136 +b) 40-43, f 137-146 +c) 43-46, f 147-152 +d) 46-48, f 157-164 +e) 51-53, f 177-182, By courtesy.

LOGUNYLLUS syn. Yllenus (alb. group) mirandus : Wesolowska W. 1996. Arthr. Sel. 5 (1/2): 46-48, f 41a-e. By courtesy.

LOGUNYLLUS syn. Yllenus (alb. group) nigritarsis +b) nurataus + c) pavlenkoae +d) pseudovalidus : Logunov, Marusik 2003b.: 55-53, f 196-200 +b) 56-58, f 216-217 +c) 58-60, f 201-212 +c)33-35, f 121-127. By courtesy.

+c)

LOGUNYLLUS syn. Yllenus (alb. group) arabicus +b) bactrianus : Proszynski 1993. F. Saudi Ar. 13: 48, f 37-40 +b) Andreeva 1976. 91, f 127-128 + Logunov, Marusik 2003b.: 31-33, f 115-120 + c) One or two species? Comparison of palps of L. arabicus and saliens (by Prószyński), misidentification of female L. saliens (by Prószyński and Logunov). By courtesy.

LOGUNYLLUS syn. Yllenus improcerus +b) knappi + c) logunovi : Wesolowska, van Harten 1994, 86: 78-81, f 157-158 [redrawn] +b).Wesolowska, van Harten 1994. 86: 81-83, f 159-160 +c) Wesolowska, van Harten. UArab Emirat. 2010. By courtesy.

LOGUNYLLUS syn. Yllenus tschoni +b) halugim : Lectotype - drawn Proszynski. In: Punda 1975. Ann. zool., 33: 37-38, f 6-7 +Logunov, Marusik 2003b. 72-74, f 253-262 + Wesolowska, van Harten 2010. UArab Emir. 3: 60, f 92-97 +b) Logunov, Marusik. 2003b. 48-50, f 174-176. By courtesy.

Gen. Marusyllus Prószyński, 2016 gen. n. (21 species)
[partial synonym for the genus Yllenus (hamifer group Prószynski, 1968)].

Yllenus (hamifer group) Prószynski, 1968e: 430.,
Yllenus (hamifer group) Logunov, Marusik 2003b: 118.

Type species Yllenus hamifer Simon, 1876.
Etymology. Named in honor of prominent arachnologist - Dr. Y.M. Marusik [MARUS-ik-YLL-US], co-author of the big monograph of the genus Yllenus, the editor of many of my publications and personal friend. Grammar gender - masculine.
Remarks. The description of this genus upgrades, without other changes, previously described group of species Yllenus (hamifer group) (Prószynski, 1968: 430., Logunov, Marusik 2003b: 118).

Diagnosis. General characters shared with remaining genera of YLLENINES (see above), but palps and epigyne are very special. Tip of cymbium striking by being drawn into long, anterior "tail", arching above bulbus and serving as guide to embolus, the latter's hair thin and very long. Bulbus relatively small. Conductor very unusual, crescent shaped, broad and robust, in majority of species larger than bulbus. Palpal femur broad, crescent shaped, tibia short with large tibial apophysis, articulating with corresponding plate-like outgrowth of the cymbium's postero-vental angle. Spermathecae set transversally, flattened oval, small and simple but with developed posterior part, often coiled and set parallel to epigyne's axis. Copulatory ducts thin walled, long but twisted into tight and compact, double spiral. Length of ducts seem to parallel length of embolus. Integral part of the definition are drawings below.

The following species are transferred from Yllenus Simon 1868 to Marusyllus gen. nov.: Marusyllus aralicus (Logunov, Marusik, 2003), comb. n., M. auspex (Pickard-Cambridge O., 1885), comb. n., M. bajan (Prószynski, 1968), comb. n., M. bator (Prószynski, 1968), comb. n., M. coreanus (Prószynski, 1968), comb. n., M. elegans (Kroneberg, 1875), comb. n., M. gregoryi (Logunov, 2010), comb. n., M. hamifer (Simon, 1895), comb. n., M. kalkamanicus (L. & M., 2003), M. karnai (L. & M., 2003), M. kotchevnik (L. & M., 2003), M. maoniuensis (Liu, Wang, Peng X., 1991), M. mongolicus (Prószynski, 1968), comb. n., M. murgabicus (L. & M., 2003, comb. n., M. namulinensis (Hu, 2001), comb. n., M. pamiricus (L. & M., 2003), comb. n., M. pseudobajan (L. & M., 2003), M. robustior (Prószynski, 1968), comb. n., M. tuvinicus (L. & M., 2003), comb. n., M. uzbekistanicus (L. & M., 2003), comb. n.

MARUSYLLUS syn. Yllenus (hamifer group) hamifer :Proszynski J. 1968e. Ann. zool. 26: 430-434, f 2, 17, 32, 45, 57, 78-84. By courtesy.

MARUSYLLUS syn. Yllenus (hamifer group) bajan : Proszynski J. 1968e. Ann. zool. 26: 440-444, f 95-97. By courtesy.

+b)

MARUSYLLUS syn. Yllenus (hamifer group) coreanus + elegans : Proszynski 1968e. Ann. zool. 26: 438-440, f 19, 34, 72, 76-77, 90-91 + Logunov, Marusik 2003b: 131-134, f 12-16, 26, 30, 32 + 51-55, 446-455 +c) Kroneberg 1875: 50, t 5, f 37. By courtesy.

MARUSYLLUS syn. Yllenus (hamifer group) robustior : Proszynski J. 1968e. Ann. zool. 26: 435-438, f 18, 33, 58, 85-89. By courtesy.

MARUSYLLUS syn. Yllenus (hamifer group) mongolicus:Proszynski J. 1968e. Ann. zool. 26: 450-454, f 5, 13, 22, 36, 48, 62, 111-118. By courtesy.

MARUSYLLUS syn. Yllenus (hamifer group) auspex (a+b+c) + d) bator :Proszynski J. 1968e. Ann. zool. 26: 445-450, f 108-110, 98-100 +b) comparison of paralectotypes of M. baltistanus and M. auspex - Prószyński 2015 online +c) Logunov. Arthropoda Selecta 20: 239, f 16-18+d) Logunovet al. 2011 .Arthr. Sel. 20: 239, f 16-18. By courtesy. REMARK. Synonymy of M. auspex and baltistanus is sustained by comparison of epigyne of both paralectotype specimens, misidentification of other specimens is irrelevant.

MARUSYLLUS syn. Yllenus (hamifer group) aralicus + kalkamanicus : Logunov, Marusik 2003b: 119-121, f 71, 406-415 + 1999: 275-277, f 44-47.

MARUSYLLUS syn. Yllenus (hamifer group) karnai +b) kotchevnik +c) murgabicus + pamiricus : Logunov, Marusik 2003b: 138-139, f 466-471 +b) 139-141, f 472-475, +c) 144-146, f 485-492 +d) 146-149, f 505-511. By courtesy.

MARUSYLLUS syn. Yllenus (hamifer group) maoniuensis + namulinensis: Logunov, Marusik 2003b: 151, f 493-496 + 136, f 497-498. By courtesy.

MARUSYLLUS syn. Yllenus (hamifer group) pseudobajan + tuvinicus + uzbekistanicus: Logunov, Marusik 2003b: 150, f 435-437 + 152-154, f 85, 519-522 + 154, f 523-534. By courtesy.

MARUSYLLUS syn. Yllenus gregoryi: Logunov 2010. Bull. Br. arachnol. Soc.: 15 (3), 91–92, figs 1-2.4. By courtesy.

Gen. Yllenus Simon,1876 (sensus Prószyński, 2015) (19 species)
[meaning restricted to Yllenus (arenarius group) Prószynski,1968]

Yllenus (arenarius group) Prószynski, 1968e: 479,
Yllenus (arenarius group) Logunov, Marusik 2003b: 85.
Type species Yllenus arenarius.
DIAGNOSIS. Genus consists of species remaining from the original genus Yllenus, left after separation of new genera Logunyllus and Marusyllus (see above), sharing general characters of YLLENINES but differing in characters of palps and epigyne. Male palps with robust cymbium and bulbus, conductor very robust but elongated and flattish, embolus indistinctly longer than bulbus, hairlike but thick. Tibial apophysis varies from prominent to enormous, articulating with corresponding structures of cymbium's wall. Epigyne with median pocket developed as a distinct plate at the posterior edge, in some species bending ventral wards. Copulatory ducts thick walled, usually short and gently bent, spermathecae thick walled. Drawings below constitute integral part of the genus definition.

Yllenus (arenarius group) arenarius: Proszynski J. 1968e. Annales zoologici, 26: 488-492, f 9, 29, 44, 56, 71, 178-184 + Logunov, Marusik 2003b. KMK Sci.Press: 85-89, f 56 + ©Photo M. Bartos, 2000. By courtesy.

Yllenus horvathi :Proszynski J. 1968e. Ann. zool. 26: 481-488, f 10, 30, 43, 55, 70, 170-177 + © Photo: T. Szuts. By courtesy.

Yllenus arenarius group)flavociliatus :Proszynski J. 1968e. Ann. zool. 26: 479-481, f 168-169. + Punda 1975. Ann. zool.
33, 3: 38, f 8 + Logunov, Marusik 2003b: 97-100, f 80, 338-344.. By courtesy.

Yllenus arenarius group) kulczynskii + b) staregai: Punda 1975. Ann. zool.: 33, 3: 39-41, f 9-12 + b) 39-41, f 13-14 + c) curageous identification and matching of the above species (without examination of types): Logunov 1992. Arthropoda selecta 1(2): 67-70, f 8a-g, 9a-d. By courtesy.

Yllenus (arenarius group) zyuzini : Logunov, Marusik 2003b: 116-118, f 45, 53, 55, 69-70, 394-405. By courtesy.

Yllenus (arenarius group) desertus : [The same paratype specimen, as pictured by:] Wesolowska W. 1991. Stutgarter Beitr. Naturk., A, 465: 4, f 10-12 and Logunov, Marusik 2003b: 89-90, f 306-308 + 90-92, f 312-313. By courtesy.

Yllenus (arenarius group) charynensis + : Logunov, Marusik 2003b: 89-90, f 306-308 + 90-92, f 312-313. By courtesy.

Yllenus (arenarius group) dunini + b) gajdosi +c) karakumensis : Logunov, Marusik 2003b: 92-94, f 64-65, 67, 320-324
+b) 100-101,f 309-311 +c) 103-105, f 353-357. By courtesy.

Yllenus (arenarius group) erzinensis : Logunov, Marusik 2003b: 94-94, f 325-335. By courtesy.

Yllenus (arenarius group) kononenkoi +b) lyachovi + c) marusiki : Logunov, Marusik 2003b: 105-107, f 358-365 +b) 107-110, f 372-383 +c) 110-111, f 314-316. By courtesy.

Yllenus arenarius group) rotundiorificus : Logunov, Marusik 2003b: 111, f 336-337. By courtesy.

Yllenus arenarius group) turkestanicus +b) uiguricus +c) zaraensis : Logunov, Marusik 2003b: 111-114, f 79, 384-388
+b) 114-116, f 389-393 +c) Logunov 2009. Turk. Contrib. Nat. History 12: 917-918, 20-21. By courtesy.