Review of the genus Hasarius ( Araneae : Salticidae )-a taxonomic fiasco 1 *

The genus Hasarius Simon, 1871, is revised following methodology recommended by “Pragmatic classification” of Prószyński (2017). Structure of the genus is insufficiently known, containing at present single cosmopolite species and a few of uncertain congeners. The paper introduces the following nomenclatorical corrections: Hasarius adansoni: Jastrzebski, 2010b: 321, f. 1, 4-5 (female only) = Hasarius tropicus Jastrzebski, 2010 correction of identification. Species: kweilinensis Prószyński, 1992, orientale Zabka, 1985, dactyloides Xie, Peng & Kim, 1993 listed variably in combinations with generic names Habrocestum, Habrocestoides, Hasarius and Chinattus seem to be misplaced in these genera and deserve transfer to own new genus. Qualification of 6 species of Hasarius as "nomina dubia" by the WSC (ver. 18.5) after Roewer's (1954[1955]: 1523-1524) is changed to "pending revision" because of existence of preserved type specimens. Synonymy of the genera Gedea Simon, 1902 and Meata Zabka, 1985 proposed recently by Maddison (2015) without published documentation are not recognized here until proof will appear printed.


Introduction
History of research on genus Hasarius Simon, 1871 reflects changing approach to taxonomic research.Wanless (1983Wanless ( [1984]]: 471) writes that Hasarius comprises "about 45 species", Prószyński (1971: 412) lists 38 "species" labeled as Hasarius in various collections of spiders, the Salticidae Database of Prószyński (2016b: 16) gives lists of 32 species and of 70 synonyms, which is not much different from joint number of 98 species and synonyms given by the WSC (ver. 18.5, assessed November 2017) but in the last paper Prószyński (2017b: 16) gives number of species as 16. and stretching anteriorly beneath eyes lateral, its optical frontal extension is white mane on palps, usually held horizontally by a male Hasarius (Fig. 1B, E, I, L, N1, P).Abdominal pattern consists of semilunar white line along anterior edge, broad median lighter belt along the whole length of abdomen, accompanied in posterior third of length by a pair of minute, contrasting pair of white dots (Figs 1A,2B).That color pattern may appear variable, be either striking or barely discernible, white lines may be thin or broadened along their course, in preserved specimens may partially disappear (Fig. 1G).There seems to be difference in the hue of the background body coloration, which typically may be relatively light (Fig. 1A) or almost black (Fig. 1L) or fawn (Fig. 1D).Color pattern of face in males may be very significant, with presence of red, red and white, or yellow rings of scales around eyes I, red lower lateral bands entering eye field (Fig. 1N1), or a pair of small red dots beneath eyes I lateral and small white triangle above touching point of eyes I median (Fig. 1P).The taxonomic significance of these markings deserves revision, which however cannot be undertaken without photographic documentation.It should be noted that shape of embolus and its basis is somewhat variable and deserves more attention (Figs 1B, J, M, 2C, F-H).
Distribution.H. adansoni is commonly accepted as cosmopolite in warmer parts of the world and as an invader of hothouses in cooler countries.It is now clear, that majority of species placed at some times in Hasarius, were misplaced, and should be transferred elsewhere (Fig. 3).While placement of H. tropicus (Figs 1N1-O) is uncertain (male palp indicates rather Chinattus), discovery of unnamed "tadpole eating" Hasarius species (Figs 1P-Q) in S India suggest that distributional center (a "home land") of the genus may be located in tropical Asia, different coloration of specimen from Kenya (Figs 1L, but note difference in embolus -1M) may ultimately be recognized as another separate species.So there is a possibility of discovery of other local species in other warm areas of the Old World.

Hasarius insularis Wesolowska & van
Harten, 2002 -shape of thick walled spermatheca and ducts (Fig. 2M) corresponds with the same structures in H. adansoni (Fig. 2D), but no other diagnostic characters are known in females of this genus, confirmation of classification of this species must be delayed until male will be known.
H. tropicus Jastrzebski, 2010 from Phuntsoling in Bhutan was described from male specimen, presumably because of typical white mane on palps, in spite of different body color pattern, especially striking red coloration of lower anterior angle of carapace (Fig. 1N1).The original description is illustrated, however, by very different palps (Fig. 2I), not resembling other species of Hasarius.It was discovered recently that palpal white mane occurs also in some species of Euophrys C. L. Koch, 1834(E. frontalis (Walckenaer, 1802), E. herbigrada (Simon, 1871) and E. sulphurea (L.Koch, 1867) -for documentation see Prószyński (2018cin press).Therefore the white hairs on palps may be less useful character than it was expected.The matter should be checked on other specimens and species, documented by both color photos of habitus and drawings/photos of palps, epigyne and spermathecae.Female collected from the same locality and by the same expedition was misidentified as H. adansoni from which differs by color pattern (Fig. 1O) and details of epigyne (Fig. 2J), unfortunately spermathecae were not documented -it is assumed provisionally that it is matching the male H. tropicus and tentatively transferred to that species.

Hasarius adansoni (Audouin, 1826)
Type specimen -collected [presumably by Etienne Geoffrey Saint-Hilaire] from unspecified locality in "Egypt and Syria" (which, at the time of Napoleonic Invasion 1798-1801, extended to the present day Israel) documented on original drawings (Fig. 2A).Present location of type specimen is unknown, presumed lost.

Designation of neotype.
Due to diversity shown in existing documentation (Figs 1A-N, 2A-H) of the cosmopolite species assumed to be Hasarius adansoni (being simultaneously type species of the genus Hasarius), there is particular need to designate neotype, to stabilize understanding of the species and to replace lost original type specimen.Good candidate for that is male specimen from Israel [possible terra typica, or adjacent to it], documented by Prószyński (2003: 68, f. 256-257) (Fig. 2B-C), which agrees with characters shown in the original drawing (Fig. 2A).The proposed neotype specimen is accompanied by matching female (Fig. 2D) from the same area.The proposed neotype specimen is preserved in the Israel National Arachnid Collection at Hebrew University, Givat Ram Campus, Jerusalem, Israel.Documentation studied.Summary of world's literature provided in "Monograph of Salticidae (Araneae) of the World 1995-2016", part I & II, by Prószyński (2016a, b, and current literature).
Remarks.The diagnoses, descriptions and graphic documentation of Hasarius adansoni, available in the literature, contain characters pertaining, in fact, to the whole genus, but insufficient to subdivide it into possible partial species.Solution of that hypothesis and writing eventual differentiating diagnoses must be delayed until next revision.

Placement and relationship of the genus Hasarius
There are two different proposals of placement of the genus Hasarius (Figs 5 and 6 below).Pragmatic classification proposed by Prószyński (2016aPrószyński ( , 2017b) ) points at similarity of palps, and in a lesser degree of epigynes.That methodology is based on comparison of morphological characters, and is aimed primarily at identification of genera and species of Salticidae, it is not a phylogeny guide but may help in understanding phylogeny of particular taxa as a side effect.It requires: 1) usage of existing, or preparation of new, diagnostic graphic documentation, of high quality and in a standardized way; 2) charting that documentation into comparative plates containing all relevant recognizable species (or genera, or super generic taxa, whichever are needed for particular case), always checked with type species of relevant genera; 3) all findings should be transparent and open to permanent scrutiny at all time.
Molecular phylogeny and classification, as developed in publications on Salticidae by papers of Maddison et al. (since 2003), could be understood as a system permitting quick establishing phylogenetical relationships among large number of taxa, a system based on selected fragments of DNA, processed with appropriate computer programs.Its visible output are phylogeny trees, grouping genera according to highly sophisticated indices.The morphological premises for these relationships of Hasarius were quoted but not documented, it may be suspected that the arrangement presented includes component of previous knowledge of morphology, applied intuitively in part.Valuable comments to tribus Hasariini, pertaining to Hasarius and related genera, are give by Maddison (2015, Journal of Arachnology 43: 246-247 and 278), compilation of diagnostic characters of these genera are shown in the present paper on Figs 6-7.Due to different approach, comparison of these characters are not fully compatible with conclusions drawn from morphological studies presented in Fig. 5.Further research may, perhaps, resolve these discrepancies.It is not clear which mutual properties are shared by genera illustrated on these plates.

Conclusion
Analysis of literature data on the genus Hasarius does not yield a coherent picture of a biological taxon, a constellation of diverse but related species, having own summary areal consisting of ranges of particular species, displaying similar biological and ecological properties, own history of evolution and spreading, different from majority of other biological units of a genus rank.Apart from poor state of knowledge of fauna of Salticidae, this is probably a result of general conviction that Hasarius adansoni is a cosmopolite species and assumption that every Hasarius-like species is H. adansoni.The present paper confirms existence of at least two other distinct species: "H".[?] tropicus in Bhutan (Fig. 1N1), and undescribed "tadpole-eating" species (Fig. 1P-Q) photographed in the West Ghats Range in India.Enclosed photographs show black bodied specimen photographed in Kenya (Fig. 1D), brownish specimens from Singapore (Fig. 1D), lighter colored immigrant in Slovakia (Fig. 1A, C) -are these really conspecific?Palp of Israel specimen (Fig. 2B) is slightly different from that in Slovakian specimen (Fig. 1B), and manifestly different from that of Bhutani species (Fig. 2I) (dismissed as mismatched -but is that sure?).These are problems demanding answer from future revision.

Figure
Figure 1A-N.Habitus of Hasarius adansoni and related species from various areas of the World.A-C -Exemplary documentation of H. adansoni from Slovakia (A-B -male habitus and palp, C -female habitus and cleared epigyne), D -H. adansoni male from Singapore, E -H. adansoni from Borneo: Brunei, F -H. adansoni from Japan, G-K -H.adansoni from Philippines: Manila-Park, L-M -H.adansoni from Kenya, N -H.adansoni female from India.SOURCES: A-C -©Photo A, Sestakova, D -© Photo H.K. Tang, E -©Photo Koh J. 2013, F -© Ono, Ikeda, Kono.Salticidae of Japan, G-K -©Photo Freudenschuss, L-M -© Phot J. Holstein, N -©Photo by Marashetty Seenappa.All ©copyrights are retained by the original authors and copyright holders, used here by their courtesy.

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