Further notes on the Harmochireae of Africa (Araneae, Salticidae, Pelleninae)

Four new spiders, Bianor paulyi sp. n. (♂♀; from Madagascar and the Comoros), Microbianor deltshevi sp. n. (♂♀; from Madagascar), M. madagascarensis sp. n. (♂♀; from Madagascar), and Sibianor anansii sp. n. (♂♀; from Botswana), are diagnosed, illustrated and described. A new combination is suggested: Sibianor proszynskii (Zhu et Song, 2001) comb. n. (ex Harmochirus). New faunistic records from Africa are given for Bianor albobimaculatus, B. kovaczi, Harmochirus luculentus, Modunda staintoni, Neaetha oculata, and Sibianor kenyaensis. A brief discussion of the current state of knowledge of the Harmochireae is also provided.


Introduction
Originally (Simon 1903), the group Harmochireae consisted of a single genus Harmochirus.Referring to the atlases of diagnostic illustrations by Prószyński (1984Prószyński ( , 1987)), Żabka (1991: p. 12, table 1) commented that the Harmochireae was a group of randomly selected genera, of which four genera (Bianor, Harmochirus, Neaetha and Modunda) were similar; yet, he discussed neither its diagnostic characters, nor its taxonomic status.Later, Logunov (1996Logunov ( , 2001) ) provided a diagnosis of the Harmochireae and included seven genera in it (see Table 1).Based both on somatic morphology and on the structure of the copulatory organs, all the salticid genera currently included in this group were diagnosed, (re)described and discussed in detail by Logunov (2000Logunov ( , 2001)).
Th e group Harmochireae is a member of the subfamily Pelleninae (sensu Maddison and Hedin 2003;Maddison et al. 2008).Its congeners can be readily distinguished from the rest of the Pelleninae genera by the absence of the compound terminal apophysis and cymbial dorso-lateral projections in males and of the epigynal fl aps in females (for further details see Logunov 2001).Th e group Harmochireae most probably deserves tribal status within the Pelleninae, but this matter is outside the scope of the present work.It is worth mentioning though that Bianor and Sibianor, the two genera of Harmochireae sampled by Maddison et al. (2008) for their phylogenetic analysis, formed a separate clade within the Pelleninae (Op.cit.: fi g. 9).Th e taxonomic status and composition of the group Harmochireae require further attention in the future, when more genera (particularly, Harmochirus, Microbianor and Neaetha) have been sampled for DNA sequence data.
Since the latest review of Harmochireae (Logunov 2001) only a few new species have been described (e.g., Jastrzębski 2007), with one of them, Bianor biguttatus, described from north Africa (Arabian Peninsula) (see Wesołowska and van Harten 2002).In the present work, I have considered new African material of various Harmochireae genera (10 species altogether) which became available to me in 2008.Four new species are described, and new faunistic records are reported for six other species.
* Th e male mentioned by Jastrzębski (2007: p. 27) (Strand, 1907) C and E Africa, Madagascar.Th e taxonomic status and validity of this species are uncertain (Logunov 2001).H. duboscqi (Berland et Millot, 1941) Ivory Coast, Senegal.Likely to be a junior synonym of H. luculentus (Logunov, 2001).H. rufescens Caporiacco, 1940 Somalia; nomen dubium (Wesołowska 1994;Logunov 2001).Modunda Simon, 1901M. ghigii Caporiacco, 1949 Kenya.Th is species, listed by Platnick (2009) under the genus Bianor, is of uncertain taxonomic status, as the ♀ holotype (see Caporiacco 1949: fi g. 96) is not a member of the Harmochireae (no central blind-ending pocket!) and belongs elsewhere.M. orientalis Dönitz et Strand, 1906 Japan.Th is species, listed by Platnick (2009) under the genus Bianor, was described from a single ♀ (Bösenberg and Strand 1906: table 8, fi g. 123); as the original illustration is insuffi cient to assign the species to any of the currently known Harmochireae genera, it is safer to treat the species as being of uncertain taxonomic status.Neaetha Simon, 1884 N. aegyptiaca Denis, 1947 Egypt.On the basis of the ♂ palp illustrated nicely by Denis (1947: plate V, fi gs 14, 15), it is easy to conclude that N. aegyptiaca is not a member of Neaetha but belongs in Evarcha Simon, 1902.

Genera and species
Distribution, comments and references N. alborufula Caporiacco, 1949 Kenya.Th e original illustration by Caporiacco (1949: fi g. 91) leaves no doubt that this taxon is not a member of Neaetha, as the fi gured ♂ palp is more similar to that of Hispo Simon, 1885 rather than to any Bianor-type taxa.N. catula Simon, 1886 E and S Africa.Th is species is of uncertain taxonomic status: the ♀ illustrated by Caporiacco (1949: fi g. 90) does not belong to the Harmochireae, as it does not have the central pocket.N. fulvopilosa (Lucas, 1846) Algeria, Tunisia.Th is species is of uncertain taxonomic status, known only from the original description.N. ravoisiei (Lucas, 1846) Algeria, E Africa.Th is species is of uncertain taxonomic status.Th e record by Strand (1907: p. 720) was based on an immature specimen.Stichius Th orell, 1890 S. albomaculatus Th orell, 1890 Sumatra; nomen dubium (Logunov 2001).Velloa Peckham et Peckham, 1903 V. modesta Peckham et Peckham, 1903 S Africa; nomen dubium (Wesołowska 1994, Logunov 2001).

Bianor Peckham et Peckham, 1885
Bianor is the largest genus of the Harmochireae, consisting of 21 valid species (Table 1), with the majority of them occurring in the Ethiopian or Oriental Regions.Six additional species have uncertain taxonomic status or are invalid (  (2001).Yet, such reconsideration will only be possible pending collection of a series of both sexes from the type locality of B. rusticulus, given on the original label as 'Clanwilliam, Cape Colony'.With a few exceptions (e.g., B. biocellosus Simon, 1902), species of Bianor cannot be reliably diagnosed from the female copulatory organs alone because of a wide range of intraspecifi c variation (for details, see Logunov 2001: p. 222).Th erefore, descriptions based on single females are hardly suffi cient for proper diagnoses of Bianor species.Yet, such descriptions continue to appear (Jastrzębski 2007;Wesołowska and van Harten 2002).
Comments.B. albobimaculatus is the most common and widespread species of the genus Bianor, reported from South Africa, northward throughout the entire continent to the Mediterranean, the Arabian Peninsula and the Near East, and then northeastward to Central Asia (Logunov 2001).However, in the light of new records of B. kovaczi from Botswana (see below), whose females are practically identical to those of B. albobimaculatus, some of the records of the latter species from South Africa should be revised.

Bianor kovaczi Logunov, 2001 Figs 1-6
Identifi cation.Wesołowska and Tomasiewicz (2008:  Comments.To date, this species has been recorded from Ethiopia only (Logunov 2001;Wesołowska and Tomasiewicz 2008).New records here from Ivory Coast and Botswana signifi cantly extend our knowledge on the distribution of B. kovaczi, which seems to be a common Afrotropical species.Males of B. kovaczi can easily be distinguished from the remaining Bianor species by the proportions of the tegulum and especially by the relatively large promarginal teeth of the chelicerae (see Wesołowska and Tomasiewicz 2008: fi g. 14).Th e latter character varies to some extent, which is Unfortunately, females of all African Bianor species cannot be reliably diagnosed by their copulatory organs; males are always required to identify a species.Paratypes.MADAGASCAR: 2♂5♀ (MRAC,225.263;[13][14],together with the holotype; 1♂2♀ (MRAC,201.744,206.734),same locality, Tamatave prison, on sand and masonry walls, 09-11.19946), reported from several African countries (see above), but can be reliably distinguished from it by the cheliceral dentition in males: i.e., the promarginal tooth being rather large and bicuspid (Fig. 12).Th e conformation of the epigyne and spermathecae in both species (cf.Mt pr 1ap, rt 2ap.Colouration.Carapace russet, punctured-reticulate, sparsely covered with white scales (Fig. 7); clypeus and 'cheeks' russet, 'naked' (with sparse black hairs).Sternum yellowish brown, covered with white protruding hairs.Labium, maxillae and chelicerae russet.Abdomen (Fig. 7): dorsum brown, with pairs of white spots and without scutum; sides and venter yellowish grey.Book-lung covers and spinnerets yellow, tinged with grey.Legs I stronger and longer than others, russet, Tb, Mt and Tr covered with protruding light hairs.Legs II-IV yellowish brownish, with all tarsi contrastingly yellow.Palps yellowish-brownish; palpal structure as in Figs 10-11.9), but diff ers as follows: carapace much more densely covered with white adpressed scales; clypeus and chelicerae anteriorly covered with white hairs and scales; dorsum with poorly marked or no pattern of white spots; palps entirely yellow.Epigyne and spermathecae as in Figs 13-14.

Harmochirus Simon, 1885
Th e genus Harmochirus accommodates six valid species (Table 1) plus three species names of uncertain taxonomic status (Table 2).Th e genus is largely restricted to the Oriental Region, with a few species (e.g.H. insulanus and H. zabkai) also known from southern areas of the Palaearctic Region and one (H.luculentus) from the Ethiopian Region.

Microbianor Logunov, 2000
Th is is a small, recently described genus with fi ve valid species (Table 1).Originally, the genus was considered endemic to the Seychelles (Logunov 2000), but after the discovery of M. saaristoi in Réunion (Ledoux 2007) it became clear that the genus was more widespread.Th e fi ndings of two new Microbianor species from Madagascar support this conclusion.It is interesting to note that after the original description of Microbianor (Logunov 2000) all new discoveries of its species have been made in the islands of the Indian Ocean rather than continental Africa.Furthermore, both new Malagasy species came from the NE shore of Madagascar.Th us, it is reasonable to assume that the distribution of Microbianor may refl ect the Asian faunal affi nities of Madagascar, which have been reported for a number of animal groups including scorpions and spiders (e.g., Legendre 1972;Lourenço 2003;Griswold 2003).If so, more new species of Microbianor might also be found in Sri Lanka and India.Paratypes.MADAGASCAR: 12♂4♀1juv (MRAC,200.164;,together with the holotype; 2♂1♀ (MRAC, 177.809), same locality, 11.1993, AP;1♀ (MRAC, 177.904), same locality, Asplenium-forest, 12.1993, AP;1♂ (MRAC, 201.769), Madagascar, Tamatave, Foulpointe, forest on clay, 07.1994, AP;1♀ (MRAC, 200.225), same locality and habitat, 12.1993, AP;1♂ (MRAC, 205.810), same locality, yellow trap on upper beach margin, under Scaevola taccata, 11.1995, AP;1♂ (MRAC, 200.064),Madagascar, Foulpointe, sea shore forest, sieving litter, 11.1993, AP.

24 27
Diagnosis.Compared to all the described species of Microbianor (see Logunov 2000), M. madagascarensis sp.n. diff ers in having the best-developed and most clearly observable fossae in females (Fig. 31; in the rest of Microbianor species the fossae are undeveloped or poorly marked).By conformation of their palp, males of M. madagascarensis sp.n. are similar to those of M. nigritarsis (cf.Logunov 2000: fi gs 11-12), from which they can be distinguished by yellow tarsi tinged with brown in their basal halves (Fig. 26; black in M. nigritarsis).Females of both species are quite distinct in the conformation of their epigyne and spermathecae (cf.rest of carapace .Clypeus russet, with sparse long white hairs forming a triangle and overhanging the chelicerae.Sternum yellowish-brownish.Maxillae, labium and chelicerae russet.Abdomen greyish-yellowish; dorsum, entirely covered with brownish shining scutum and with a pair of transverse white spots (sometimes poorly marked; Fig. 24).Book-lung covers and spinnerets yellow, slightly tinged with brown.All legs yellow, with no brown rings, but tibiae I and the basal half of tarsi I brownish (Fig. 26).Palps yellow to brownish, but cymbium always yellow; palpal structure as in

Modunda Simon, 1901
Th is is a very small genus consisting of two valid species, of which one (M.aeneiceps) remains known only from the type locality (Sri Lanka: Colombo).Th e second species mentioned below is a widespread Afrotropical species.

Neaetha Simon, 1884
Th is is a relatively small genus consisting of 7 valid species that are known exclusively from the Mediterranean and Africa (Table 1).Five additional species names are of uncertain taxonomic status (Table 2).A new record of a Neaetha species given below has extended the distribution of the genus as far south as Botswana.
Comments.Th is is the fi rst record of N. oculata from southern Africa.To date the species was known from the Eastern Mediterranean, Egypt and the Arabian Peninsula (Wesołowska and van Harten 1994;Logunov 1996).Th e species N. irreperta Wesołowska et Russell-Smith, 2000, described recently from Tanzania, displays a strong similarity to N. oculata.On the basis of published accounts of both species, it is impossible to decide whether they are conspecifi c or not.Th e matter requires further attention in the future.

Sibianor Logunov, 2001
Th is genus, with its 15 described species, is almost entirely confi ned to the Palaearctic and Afrotropical Regions (Table 1).Th e only exception is S. aemulus (Gertsch, 1934), known from the Nearctic Region (Canada and USA; Ubick et al. 2005).A new species described below is one of the three species known to date from tropical Africa.It is safe to assume that the real diversity of the Afrotropical species remains largely undescribed.Diagnosis.Of the African species of Sibianor, the new species is most similar to S. victoriae (cf.Logunov 2001: fi gs 292, 293), described from a single male from Kenya.S. anansii sp.n. diff ers in having a wider tibial apophysis, which is also of a diff erent shape (Fig. 39).Th e female of S. anansii sp.n. diff ers from all the known Sibianor  Comments.Th is is the fi rst record of S. kenyaensis following its original description and a new record outside the type locality (Kenya: Kilifi ).

Discussion
At present, 57 recognized species from seven genera are included in the group Harmochireae (Table 1).Of these, 20 species are known only from their respective type localities, and a few remain unrevised and/or of uncertain taxonomic status.An additional 17 species names are excluded from the list of valid names for various reasons (see Table 2).
Of the Neotropical genera of the Pelleninae, Havaika Prószyński, 1992, an endemic genus to the Hawaiian Archipelago, is rather similar to members of Harmochireae in the conformation of its copulatory organs, especially in the thin, whip-shaped embolus (see Prószyński 2008: fi gs 21-36).Yet, according to the available molecular phylogeny of Salticidae (Maddison and Hedin 2003: fi g. 5;Maddison et al. 2008: fi g. 9), within the Pelleninae, Havaika is closer to Pellenes Simon, 1876 andHabronattus F. O. Pickard-Cambridge, 1901 than to Bianor and Sibianor, i.e. it does not belong with the group Harmochireae as it is currently diagnosed (see Introduction, and Logunov 2001).
Harmochireae is predominantly an Old-World group, as over 90% of its described species are restricted to the Palaearctic, Ethiopian or Oriental Regions (Table 3).In the Neotropics, the only described species is Bianor biocellosus, known from the type locality in Brazil ('le Para' ;Logunov 2001: p. 234).A single species, Sibianor aemulus, has been described and recorded from the Nearctic Region (Logunov 2001;Ubick et al. 2005).Yet, there are three Bianor species described from the Australian Region, including New Zealand, but two of them (B.compactus and B. concolor) remain unrevised and known from the original descriptions and only from their type localities.Th e taxonomic status of both of them is uncertain, as these names might be junior synonyms of B. maculatus.With the above four exceptions, the remaining Harmochireae species have only been recorded from the Old World.Within the Old World, the Palaearctic and Ethiopian regions contain a similar number of recorded species and genera: 23 vs. 20 species and fi ve vs. six genera, respectively.Four genera (Bianor, Harmochirus, Modunda and Sibianor) have their representatives in all three zoogeographic regions of the Old World.Of these genera, Modunda is represented by a single widespread species M. staintoni, distributed throughout Africa from Botswana, north-eastward to Afghanistan and Punjab.Five of the six described species of Harmochirus are confi ned to the Oriental Region.Th e only widespread Afrotropical species is H. luculentus.Th e genus Sibianor is most diverse in the Palaearctic Region, with 10 of its 15 species recorded there.All Sibianor species have recently either been revised or been described (Logunov 2001;Zhu et Song 2001, etc.), therefore to date the species composition of Sibianor seems to be the most reliable in terms of taxonomy.Th e genus displays a clear modern centre of diversity in the Manchurian area, to which at least six species are confi ned.Bianor is the most common genus of Harmochireae in the Ethiopian and Oriental regions.In the Palaearctics, the only known, widespread species is B. albobimaculatus, confi ned to southern regions below the 40-45° latitude.
Two genera display a rather limited distribution.Napoca is a monotypic genus known to date only from Israel.All the species of Microbianor have been described from islands in the Indian Ocean, but new fi ndings from Sri Lanka and India are possible as has been discussed above (p.276).

Table 1 .
Valid species of the Harmochireae and their distribution.

Table 2 .
A list of existing species names of the Harmochireae excluded from discussion.

Table 2
).In the light of fi ndings of a new Bianor species from Madagascar and of new records of B. kovaczi from Botswana (see below), of which females are practically identical to those of B. albobimaculatus (cf.Figs 13-14 and fi gs 19-27, 36-46 in Logunov 2001), it is worth reconsidering the taxonomic status of B. rusticulus Peckham et Peckham, 1903 (known from the ♀ holotype), which was synonymized with B. albobimaculatus by Logunov fi gs 13-20).

Table 3 .
Number of described species of the Harmochireae in main zoogeographical regions*.