© 2016. Indian Journal of Arachnology 5 (1-2): 143-171 ISSN 2278-1587 (Online)

SIGNIFICANCE OF ORIENTAL TAXA IN PHYLOGENY OF CRAB ( S.LAT. AND STIPHROPODIDAE)

Pekka T. Lehtinen Zoological Museum, FIN-20014 University of Turku, Finland [email protected]

ABSTRACT Numerous taxonomic changes in Oriental crab spiders are listed with justifications and suprageneric taxa are discussed at global level. Similar pattern of more than one inner cheliceral setae is found to be a synapomorphy for Borboropactidae, Stephanopidae and Thomisidae. The separation of Stiphropodidae as a valid family is confirmed. The polyphyly of Dietinae sensu Petrunkevitch, 1928 is confirmed and its tribes are listed in Thomisidae, Thomisinae (Dietini, Alcimochthini, and Amyciaeini) and Bomidae (Apyretini, Tagulini, Emplesiogonini, Mystariini, Porropini, and Talaini). Boliscini, new tribe of Bomidae is created for Boliscus Thorell, 1891 and Corynethrix L. Koch, 1876, and family group names based on Bomis L. Koch, 1874 without known female is discussed. A strange gnathocoxal process in all Boliscus spp. is described. The Stephanopine groups Phartini, Borboropactini and new tribes Cebrenninini, Angaeini and Geraestini are transferred to Borboropactidae Wunderlich, 2004, and Phrynarachnini, Stephanopoidini, and Hedanini to Thomisidae s.str. These transfers are based on similar structure of copulatory organs of both sexes, trichobothrial patterns of legs and several ultrastructural characters. Phrynarachne Thorell, 1869 with type species P. rugosus Latreille, 1804 is now included in the same tribe as Simon, 1864. Senoculifer Balogh, 1936 from New is transferred from Philodromidae to Thomisidae and synonymized with Demogenes Simon, 1895. Xysticus palawanicus Barrion, 1995 is synonymized with Alcimochthes limbatus Simon, 1885 (Alcimochthini). Lehtinelagia Szymkowiak, 2014 and its type species Xysticus evanidus L. Koch, 1867 (Queensland) are synonymized with Tharrhalea L. Koch, 1875 and its type species T. albipes L. Koch, 1875 (Queensland). Ocyllus Thorell, 1887 with juvenile type species O. binotatus Thorell, 1887 from is synonymized with Cebrenninus Simon, 1887; the latter is selected as valid for simultaneously published names. Ocyllus pallens Thorell, 1895 is transferred to Oxytate L. Koch, 1878 in Dietini. Taypaliito Barrion, 1995 and its type species T. iorebotco (, Palawan) are synonymized with Talaus (Microcyllus) Thorell, 1892 and its type species T. (M.) elegans (, ). Takachihoa Ono, 1985 (type species Oxyptila truciformis Bösenberg & Strand, 1906 = Xysticus nebulo Simon, 1909, syn. n.) is synonymized with Pycnaxis Simon, 1895, and Coriarachne nigrostriata Simon, 1886 is transferred to Pycnaxis. Tharrhalea maculata Kulczyński, 1911 and T. variegata Kulczyński, 1911, both from New Guinea are transferred to Massuria Thorell, 1887.

INTRODUCTION This work is a part of worldwide revisional study of Thomisidae sensu auct. It is not formally a complete modern revision, as it is based on selected important results. It is a list of results with carefully justified taxonomic changes in the Oriental taxa of this group. In this way, a lot of taxonomic results about this group can be published rather quickly. Limitation of space and time available makes the inclusion of normal complete data of true revisions impossible.

December, 2016 143 Significance of oriental taxa in phylogeny of crab spiders Pekka T. Lehtinen

Complete descriptions of treated taxa are not presented here, but taxa with new status or limitation are shortly diagnosed and reasons for the new opinion is explained. Exceptionally extensive personal field work in all tropical areas has also made possible the conclusions here, even though some of them much deviate from previously presented results.

Problems to be solved. a) Many name-bearing types of crab spiders described long ago by Simon, Thorell, L. Koch, Karsch, Workman, O. Pickard-Cambridge, and other taxonomists of the past have never before been studied after their original description, even those preserved in well known European museums in Paris, Genoa, Stockholm, London, Geneva, Oxford, and Dublin. b) Several local specialists in the Oriental region have not at all or quite seldom, at least, tried to co-operate with colleagues or at least ask for advice from well trained specialists from Europe, North America or . This is unfortunately true for most arachnologists from , Pakistan and Philippines working with crab spiders after 1960. Positive exceptions have been several current arachnologists in , , Taiwan, and , while much arachnological work for (in addition to the work in Thailand) has been recently carried out by some active European specialists. Incorrectly placed Philippine crab spiders have been checked in IRRI, Los Baños, and many of Dahl´s (1907) errors from slides in ZMB. c) Fruitful work of most European, North American and even of many local arachnologists has been prevented and/or complicated by worldwide unique policies of some museums, especially ZSI (Kolkata), preventing not only the loan of material, but also suggested work with them on site. d) After the United Nations Conference on Environment and Development in Rio de Janeiro in 1992, governmental offices in many (less developed) countries interpreted the decisions in a way, which largely stopped the real co-operation with scientists in developed countries (except for those living in countries with high economical resources), e.g., by legally prohibiting all transport of biota to foreign countries for scientific purposes. Although the policies were most radically changed in Brazil, almost similar is the situation in Philippines. This explains that the situation does not depend at all on the attitudes of local arachnologists. The results are presented in two parts. Complete description of a new Malagasy-Mauritian tribe of Stephanopidae, new limitation of Stiphropodidae and Bomidae, all changes within the traditional subfamily Thomisinae and within most true members of the tribe Hedanini will be published separately. Bond et al. (2014) and Fernandez et al. (2014) have recently used an entirely different method of molecular biology, phylogenomics, and independently obtained surprisingly similar results as the phylogenetic conclusions presented here that are based on ultrastructural analysis of cuticle, setae and sensory organs. This seems to confirm the applicability of ultrastructure as a reliable method in solving problems of phylogenetic relationships of supraspecific taxa. The aim of this study is to list and discuss taxonomical changes to crab spider taxa of the Oriental region, according to the results of personal work and give sufficient justification for them, though formal details for complete taxonomic revisions are presented only for a few selected cases. Special attention has been directed to the correction of numerous strange taxonomic interpretations of Oriental crab spiders after 1960, both by Indian and Philippine active arachnologists also working with the crab spiders.

Essential abbreviations used: * I have checked the holotype/syntypes. This refers to type material of the type species.

Indian Journal of Arachnology 5 (1-2) 144 Significance of oriental taxa in phylogeny of crab spiders Pekka T. Lehtinen

# I have checked authoritatively identified material. ¤ I have compared topotypical adults of both sexes and juvenile specimens of different age with holotype in cases, where the holotype is not adult. This is the only way to confirm the identity of the holotype in a few cases. Abbreviations for eye and spinneret patterns according to widely accepted formulae. If not otherwise noted, the source of photographed and micrographed specimens is the personal temporary collection (PTL) or ZMT for material collected before 1999. A further technical solution: for all new taxa in Barrion & Litsinger (1995), Barrion is here cited as the only author, as it has been officially confirmed that Mr. Litsinger has not participated to the taxonomic work with results published in this book. This principle should be generally applied as strictly Code-compliant (ICZN, 1999 § 50.1). References concerning the book as such should not be changed.

MATERIAL AND METHODS My field research on tropical spiders began in the Oriental region in 1969 by a field trip to Sri Lanka (with four later expeditions). Number of visits to other Oriental areas are listed in brackets: (5), Thailand (4), Indonesia: Sumatra, Java, Kalimantan, Sulawesi, Bali, Lombok & Flores (5), (1), Philippines: Luzon, Mindoro, Cebu, Palawan & Mindanao (3), India (3), Pakistan (1), (3), China+Hongkong & Lantau Island+Macau & Coloane Island (3), and Taiwan (5). During these trips, spider material was also collected from Samoa (Savaii, Uapou & Tutuila), Tonga (Tongatapu, Eua, Vavau), Fiji (Viti Levu) (3), Micronesia (Guam), New Guinea with Goodenough Islands & Manus Island (2), Australia with Tasmania (4), (3), New Zealand (2), 19 islands of Polynesia (Cook-, Society-, Marquesas- & Tuamotu- Islands with 5 expeditions), as well from Easter Island, Baltra & Santa Cruz I. of the Galapagos Islands, and Kauai, Oahu, Maui & Hawaii in Hawaiian Islands. In addition to the personally collected material, an absolute provision for this study has been the checking of type material in numerous museums all over the world. Personal best, applied strategy: Selected own material packed while visiting museums and most of the work done by comparing own material with the types and possible additional specimens on site. Much type material was checked through loans. When types were no more extant or their site of preservation was unknown, other authoritatively identified material or material collected from the type locality was used instead of or in addition to nominal types. Useful information about the species which could not be studied personally was referred from Griswold et al. (2005). The light microscopical studies were done with an OLYMPUS SZH dissecting microscope and the measurements with ocular micrometer attached to WILD M5 microscope. Ultrastructural studies of leg surfaces, various types of setae and sensory organs, in addition to traditional study of male palpi and some epigynae were carried out by JEOL JSM-5200 SEM-device and necessary coating with BIORAD PS 3 sputter and digitized with SemAfore software. A part of SEM work was carried out in other institutions using different devices. Some SEM mounts were done from samples first cleaned with Terg-A-zyme using TRANSSONIC T 40 for vibration. The digital photographs were taken with Olympus digital camera and binocular and enhanced using the “CombineZS” software. All measurements are given in millimeters. The results gained with traditional taxonomic methods and large scale analysis of ultra-

Indian Journal of Arachnology 5 (1-2) 145 Significance of oriental taxa in phylogeny of crab spiders Pekka T. Lehtinen structural characters are carefully compared with the results of phylogenomic analyses and other molecular analyses recently carried out by other arachnologists, but the latter methods have not been personally applied. Phylogenetic analyses with simplest traditional methods (mainly Hennig 69) have been personally applied to several groups, but the main work with phylogenetic analyses has been concentrated to testing of previous results by adding new characters without excluding anything. The influence of different strategies in selection of characters and details of coding have been now thoroughly estimated for phylogenetic analysis, and they seem to be the main reasons for numerous errors in the results of cladistic analyses during the last three decades, as also was independently shown with use of phylogenomics by Bond et al. (2014) and Fernandez et al. (2014).

Acronyms for museums, present curator in parentheses, if not participated to on site work, loans or information: AMS = Australian Museum, Sydney, Dr. Michael Gray, Dr. Graham Milledge, Ms. Helen Smith. BMNH = British Museum, Natural History, London, Dr. Paul Hillyard, Dr. Janet Beccaloni. CCDL = temporary private collection of Dr. Christa Deeleman-Reinhold, Sabah/Leiden, Netherlands. CJCL = temporary private collection of the late Dr. Jean-Claude Ledoux, Aramon, . CJS = temporary private collection of Dr. Jenny Shield, La Trobe University, Victoria, Australia. HDE = Hope Department of Entomology, Oxford, Great Britain (information about type preservation). IRRI = International Rice Research Institute, Los Baños, Philippines, Dr. Alberto Barrion. IZPA = Instytut Zoologiczny, Polska Akademia Nauk, (M.Sc. Tomasz Huflejt) & M.Sc Dominika Mierzwa. MCSN = Museo Civico di Storia Naturale, Genoa, Italy, Dr. Lidia Capocaccia, Dr. Gianna Arbocco, Dr. Giuliano Doria & Ms. Maria Tavano MNHG = Muséum d´Histoire Naturelle, Geneva, Switzerland, Dr. Peter Schwendinger. MNHN = Muséum National d´Histoire Naturelle, Paris, France, the late Dr. Jacqueline Heurtault, Dr. Christine Rollard. MRJ = Museu Nacional, Rio de Janeiro, Brazil, Mr. Thiago da Silva Moreira, (Dr. Adriano Kury). MZUF= Museo Zoologico La Specola, Florence, Italy Ms. Sarah Whitman (Dr. Luca Bartolozzi) NCBN = Netherlands Centre for Naturalis, Leiden, Netherlands, Dr. Jeremy A. Miller. NHRS = Naturhistoriska Riksmuseet, Stockholm, Sweden, Dr. Torbjörn Kronestedt. PTL = temporary private collection of Pekka T. Lehtinen. SMF = Seckenberg Museum, Frankfurt am Main, Germany. ZMB = Zoologisches Museum der Humboldt Universität, Dr. Shahan Nawai, Dr. Jason Dunlop. ZMH = Zoologisces Staatsinstitut und Zoologisches Museum, Hamburg, Germany, Dr. Gisela Rack, (Dr. Hieronym Dastych). ZMT = Zoological Museum, University of Turku, Finland Dr. Pekka T. Lehtinen (Dr. Ilari Sääksjärvi).

Taxonomic publications were completed in a series named “Spiders of the Oriental -Australian region” about subfamilies Venoniinae and Zoicinae in Lycosidae, Stenochilidae, and Nesticidae, which were published and background work for many other groups was done. The first revisions for Oriental Thomisidae were completed in 2004 preceded by visits to more than 50 museums for checking the type material of almost all type species of the world genera

Indian Journal of Arachnology 5 (1-2) 146 Significance of oriental taxa in phylogeny of crab spiders Pekka T. Lehtinen of Thomisidae, to one of them (MNHN) for eleven times (1964-2002). Although some of the Neotropical type material of type species of Thomisid genera could not yet be personally checked for reasons of uncertainty of their present location, practically all types of type species of Oriental and Pacific genera of crab spiders could be studied, except Tharrhalea albipes L. Koch, 1875 and Corynethrix obscura L. Koch, 1876, which could be placed according to the original drawing. Many of the results presented here are derived from results of parts of my global revision and in these cases most of the detailed results will be published separately, based on large manuscripts which mostly have long time been more or less completed, waiting only for checking of minor details.

TAXONOMY

Key to crab spider families 1. Legs I-II strong, longer than leg IV, branched hairs lacking - male bulbus discoidal...... Thomisidae - male bulbus not discoidal -- median apophysis in male bulbus, tarsal sensory area present ...... Borboropactidae -- median apophysis lacking, tarsal sensory setae not in well restricted area...... Stephanopidae

2. Legs I-II & IV more or less of same length, branched hairs present, male bulbus not discoidal - tarsi and metatarsi with rigid fusion...... Stiphropodidae - tarsi and metatarsi as a movable unit...... Bomidae s.lat.

Notes concerning family taxa in the key. Thomisidae was first treated as an independent family by Sundevall (1833), Stephanopidae by O. Pickard-Cambridge (1871), Bomidae by Dahl (1907), Stiphropodidae by Lehtinen (2007) and Borboropactidae by Wunderlich (2014). As it is obvious, all these groups have independently evolved from the now extinct groups of Araneomorph spiders and not from each other, it is justified to treat these five groups as families, in spite of the fact that they are currently treated mostly as subfamilies of a single family Thomisidae. Besides this, the relationship of Bomidae and Stiphropodidae to the remaining three families is uncertain and the assigning of some tribes to Bomidae or Thomisidae is still uncertain and under continued analysis. Although Aphantochilidae Thorell, 1873 has been accepted as a family by several arachnologists, it is now generally accepted that representatives of this group are close relatives of the Thomisid subfamily Strophiinae. Philodromidae was long included in Thomisidae, but it is now known to belong to different line of evolution.

Most important morphological characters in crab spiders

Presence/absence of median apophysis in male palp: The most important morphological character in major classification of crab spiders is the presence or absence of a median apophysis as a separate sclerite articulating to the tegular surface. A median apophysis is lacking in Stephanopidae,

Indian Journal of Arachnology 5 (1-2) 147 Significance of oriental taxa in phylogeny of crab spiders Pekka T. Lehtinen and in Thomisid subfamilies Strophiinae, and Thomisinae, but always present in Borboropactidae. In contrast to many other families, there are no problems of palpal homology in crab spiders. Borboropactidae with normal Thomisid patterns of tarsal and metatarsal trichobothria (Angaeini, Cebrenninini & Epidiini) are suggested to consist the most plesiomorphic group of Thomisidae. The median apophysis has disappeared at least in two lines of evolution of crab spiders; one leading to Thomisinae, a large worldwide group and the other to Stephanopidae (with its new limitation used here) with a range from and Mauritius through eastern parts of the Oriental region, Australia and New Zealand to tropical and subtropical parts of South and Central America. There are three totally allopatric tribes, but only Stephanopini is found within Oriental region.

Trichobothrial patterns of tarsi and metatarsi. All groups of revised Thomisinae and Strophiinae have the basic pattern: one row in tarsi and at least one straight row in metatarsi, although there are sometimes a few additional subdistal trichobothria in metatarsi. All Stephanopidae (in its revised sense) have very complex patterns on tarsi, metatarsi, and tibiae. Some Borboropactidae in revised sense (Borboropactini & Cebrenninini) have more or less complex patterns on tarsi and metatarsi, although quite different from those of Stephanopidae, while the trichobothrial patterns of the three remaining tribes (Phartini, Angaeini, and Geraestini is more or less similar to the simple thomisine pattern. These characters seem to lead later to grouping of the Borboropactid tribes to subfamilies, but it has not yet been carried out here, as the reciprocal relationships of the five tribes are not yet sufficiently known.

Important ultrastructural characters in Thomisidae s.lat. or Thomisoidea Inner surface of the trichobothrial pit. This is more or less similar in most groups of Thomisidae, and also in most families in traditional Araneomorpha (except Filistatidae), consisting of parallel horizontal ridges along the entire inner wall. The strongly deviating inner surface of trichobothrial pit in Filistatidae is not discussed here, as the position of this family as a suborder of its own (based on numerous macromorphological, ultrastructural, and biological characters) will be separately published. The other exception is the crab spider family Stephanopidae in its revised limitation, where soft scales cover the pit with general structure. The scales are different in three tribes, but only the nominate tribe Stephanopini is present in the Oriental Region. The ultrastructure of the outer face of the trichobothrial plate is quite variable within true crab spiders (Thomisidae, Borboropactidae and Stephanopidae). This pattern is sometimes phylogenetically significant, e.g., as a differentiating character between Phrynarachnini and Diaeini, but most details still need further study and hence they are not further discussed here.

Diagnosis of Thomisidae A meaningful discussion about the of currently accepted subfamilies of Thomisidae and crab spiders in general is possible first, when convincing arguments for the limitation of Thomisidae are available. It is widely known that the presence of strong and long two anterior pair of legs is true for majority of the crab spiders in the northern hemisphere. However, this character is not distinct in some groups restricted to the tropical areas. According to the present study, Thomisidae s.lat. (including Stephanopidae and Borboropactidae, but not finally revised

Indian Journal of Arachnology 5 (1-2) 148 Significance of oriental taxa in phylogeny of crab spiders Pekka T. Lehtinen

Figure 1 a-h. Synapomorphy of crab spiders (Thomisidae, Borboropactidae, Bomidae and Stephanopidae): > 1 cheliceral setae on the inner face of the cheliceral furrow, a: Thomisus onustus, Turkey, Yamanlar Dagi, b: Cebrenninus sp., Myanmar, Prome, c: Angaeus sp. Taiwan Wushikan, d: Xysticus audax Finland, Ylöjärvi, e: Thomisops pupa, South , Badplaas, f: Holopelus bufoninus Thailand, Doi Suthep, g: Phrynarachne decipiens, Myanmar, Prome, h: Stephanopis n. sp. Philippines, Luzon, Caliraya, Siniloan.

Indian Journal of Arachnology 5 (1-2) 149 Significance of oriental taxa in phylogeny of crab spiders Pekka T. Lehtinen

Bomidae), the dominant crab spider superfamily Thomisoidea (Thomisidae, Borboropactidae & Stephanopidae) has one very convincing synapomorphy: The inner cheliceral margin has a row of 2-3(-numerous) setae, while all other Araneomorph families, also Stiphropodidae, have 0-1 setae only in this position.

Thomisoidea Suprageneric groups of crab spiders The concepts of taxonomic groups within the “crab spiders” have been rather variable since the time of Latreille (1829 in Cuvier: Le Règne ), when all the spider groups with laterigrade pattern of legs were still included in “Araignées latérigrades”. With that principle all Sparassid, Selenopid, and Philodromid groups were united to Thomisids. The present author (Lehtinen, 2007) first emphasized the deviating position of some traditional Thomisid groups in relation to confirmed Thomisidae. This can mainly be explained by the fact that during the active analysis of Araneomorph spiders with phylogenetic analysis only, current representatives of Stiphropodidae and Bomidae were not included in the matrices or when some of them were included, the list of characters did not include any of the synapomorphies of these families or the coding was carried out inappropriately for getting reliable relationships. Benjamin & Clayton (2016) recently included both Stiphropus and Pagida to their phylogenetic matrix, and correctly found their reciprocal relationship and also their combined position as a sister group of Apyretinini, but all Stiphropodid synapomorphies were not included in their analysis, leaving this complex still within Thomisidae. Simon (1895a) first created a classification of crab spiders, which has for many parts, constituted the frame for all later classifications. He listed six subfamilies for the accepted family Thomisidae: Aphantochilinae, Strophiinae, Stiphropodinae, Stephanopinae, Misumeninae, and Philodrominae. However, according to the current Code (ICZN, 1999 § 23.1; 35.5) the name of a suprageneric taxon must be derived from the generic name used for the highest category of the family group names, in this case on Thomisus, not Misumena. Similar problems were still frequent at the time, when the requirements of the Code were not regularly applied and even were not well known among leading taxonomists of the past. Dahl (1907) split crab spiders to quite many families (Stephanopidae, Misumenidae, Amyciaeidae, Mystariidae, Cymbachidae, Diaeidae, Platythomisidae, Xysticidae, Urarachnidae, Bomidae, Strophiidae), but this action has not been accepted by other authors, although all his names have nomenclatoral validity. Dahl (1907) did not name the subdivisions of traditional Stephanopinae, and actually there are no older family group names available for Borboropactidae, Wunderlich, 2004, although unfortunately the tribe Borboropactini represents a group with non-typical habitus for the majority of this group which must anyway be taken out from the limitation of Stephanopidae. It is quite surprising that the strongly polyphyletic Stephanopinae has not been split to the named groups, although many cladistics analyses have found distinct clades within it. Petrunkevitch (1928) transferred numerous tribes of Thomisidae to a new subfamily Dietinae, and unfortunately this incorrect classification has still been widely used, although it is just based on parallel adaptation of several groups to move on smooth surfaces, but without any synapomorphies. The monophyly of the crab spiders has been repeatedly discussed between arachnologists within

Indian Journal of Arachnology 5 (1-2) 150 Significance of oriental taxa in phylogeny of crab spiders Pekka T. Lehtinen the last forty years. Thomisidae was long classified confamilial with Philodromidae, but they have been proved to be non-relatives independently by Homann (1975), Ono (1988), and Lehtinen (1996). Some Philodromid taxa have been erroneously included in Thomisidae. Here is a previously unknown case:

Demogenes Simon, 1895 Demonax Thorell, 1881, junior homonym of Demonax Kinberg, 1867 (Polychaeta). Demogenes Simon, 1895a: 1066 (replacement name); Lehtinen, 2004: fig. 16 in p. 154. Oxyptila Berland, 1934: 133 figs 10-13, non Oxyptila Thorell, 1869: Demogenes heterophthalmus Berland, 1934 comb. n. ex Oxyptila. Senoculifer Balogh, 1936: 239, syn. n. and transfer from Philodromidae to Thomisidae; Chrysanthus, 1964: 91 figs. 21-23. Ozyptila, Ono, 1985: 29, fig. 26-32, in part (#O. nipponica only); Ono, 1988: 129 fig. 135-140 (O. matsumotoi), non Ozyptila Simon, 1864. , Tikader, 1980: 159 f. 225, in part, M. andamanensis only), non Misumenops F.O. Pickard-Cambridge, 1900 (see Lehtinen & Marusik, 2008). Type species of Demonax Thorell, 1881 by original designation and monotypy D. lugens Thorell, 1881 (*MCSN) from New Guinea; type species of Senoculifer by original designation S. dentibulbis Balogh, 1936:239 fig. 1-2 (type preservation unknown) from New Guinea = Demogenes lugens, syn.n.

Levi (1982) and Lehtinen (1996) used the superfamily name Thomisoidea with a single family, Thomisidae, including also Aphantochilidae Thorell, 1873. Homann (1975) and Ono (1988) placed Aphantochilinae as a subfamily of Thomisidae. In my present opinion, this Neotropical group is close to Strophiinae or even a part of it, and just represents maximal exaggerated development of their cheliceral modifications. Benjamin et al. (2008) claimed that his cladistic analysis proves the monophyly of Thomisidae. Unfortunately, this is true only for the groups which were included in his matrices. All genera of Stiphropodidae (raised to family status by myself in 2007) were lacking in his analysis, although species of both subfamilies, Stiphropodinae and Pagidinae are present in Sri Lanka. Recently, Pagida was included in his matrix (Benjamin & Clayton, 2016) and was found to be a sister taxon of Stiphropus, but still they were left within Thomisinae, obviously because of highly unbalanced choice of characters. It is also worth mentioning here that, in his recent analysis of Dionychan spiders, Ramírez (2014) did not study representatives of genera originally included in Stiphropodidae, neither any real Bominae, but Boliscus, regarded here to represent a new tribe in Bominae, was included in his discussions and some cladograms, but obviously only a few characters were studied, as Boliscus was placed as a sister group of Thomisus by him. (op. cit. figs. 214 A-B on p. 315 and figs. 215 A-D on p. 317). Wunderlich (2004) created the family Borboropactidae for a part of traditional Stephanopinae, but did not include all other groups which are more closely related to Borboropactus than to Stephanopis. Borboropactidae is accepted here, but with largely different limitation than the original one. It is easily differentiated from Stephanopidae by a plesiomorphic character, presence of a separate median apophysis in the male palpal bulbus.

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Oriental genera transferred from Stephanopidae to Thomisidae s.str. Although the classification of crab spiders by Dahl (1907) has been largely criticized by all later authors and most details of it have not been accepted by any later taxonomist, neither by myself, it was the first attempt to relimit Stephanopid genera to those, which are here regarded to consist the revised Stephanopinae. Origin of the results below refer to my research program of suprageneric revision of the world Thomisidae, in which several large revisions will be separately published and for those, only the most essential new results are shortly discussed here, with groups including only Ethiopian and Neotropical taxa predominantly excluded.

Position and relimitation of the tribe Phrynarachnini. The tribe Phrynarachnini has up till now listed with two widely accepted genera, Phrynarachne Thorell, 1869 s.lat., with several Old World species and the hitherto monotypic Trichopagis Simon, 1886 from , which is not further discussed here, but it is not related to Phrynarachne. The New Guinean Cladonotula Strand, 1929 is here regarded as valid sister group of Phrynarachne and most probably it is not monotypic. Cladonotula is a nomen novum for Cladonotus Thorell, 1877, preoccupied by Cladonotus Saussure, 1812 in Orthoptera. The synonymy of the Oriental Ornithoscatoides O. P.-Cambridge, 1884 (type O. ceylonica O. P.-Cambridge, 1884 #♂♀ PTL) with Phrynarachne has been widely accepted. The small Malagasy species Phrynarachne clavigera Simon, 1903 (*♀ MNHN) and P. pusiola Simon, 1903 (*♀ MNHN) must be transferred to an undescribed genus related to the Oriental -Melanesian Demogenes Simon, 1895. Phrynarachne Thorell, 1869 is a junior synonym of Botryogaster Doleschall, 1859, but much more used and an application to validate the younger name is in preparation. These genera were incorrectly placed to Stephanopinae as a separate tribe Phrynarachnae by Simon (1895a), only based on the type of gnathocoxae and cheliceral armature, both characters which have a rather low value in Thomisid taxonomy. This phylogenetic placing and limitation of the tribe Phrynarachnini has later been used by all authors, although this tribe is now found to be a much larger group, including Synema Simon, 1864 and its African relatives Pyresthesis Butler, 1879, Lampertia Strand, 1907, Firmicus Simon, 1895 and Rimania Dahl, 1907 which are all transferred here from Diaeini to Phrynarachnini. As their carapace and abdomen are not strongly modified as in Phrynarachne Thorell, 1869, the new subtribe Synemina is created for these genera without strongly modified body of Phrynarachnina. All Synemina have a pair of symmetrical light spots on abdomen, a character shared by Camaricini Simon, 1895 with somewhat similar type of copulatory organs and possibly the latter could be included in Phrynarachnini as well. Both male and female copulatory organs of Phrynarachne most closely resemble those of the well known Synema s.str. as limited by myself now with radically decreased number of species, and no species at all in the Neotropical and Pacific regions (unpublished data). The most probable reason for the fact that a suprageneric name derived from the generic name Synema has never been suggested is possibly due to the strong polyphyly of the concept “Synema” in all catalogs. Many of the species still listed in Synema by Platnick (2014) are members of the tribe Diaeini. The exceptional habitus of Phrynarachne spp. with numerous tubercles both on carapace and abdomen seems to be an autapomorphy for this genus and Cladonotula only. Besides this, the structures of the trichobothrial base and the trichobothrial patterns of legs are practically

Indian Journal of Arachnology 5 (1-2) 152 Significance of oriental taxa in phylogeny of crab spiders Pekka T. Lehtinen identical to those of Synema. There is an excellent character separating Phrynarachnini from Diaeini. The upper face of the trichobothrial base in Phrynarachnini is smooth, while it has several parallel laminae separated by transverse furrows in Diaeini (cf. plate 3 figs. k, l & o). No family group name has been based on Synema or its close relatives, as they have been traditionally included in a very large, but strongly polyphyletic tribe Diaeini, often also including Xysticus s.lat. The Code (ICZN, 1999 § 23.1.) provides application of priority, and therefore the tribe including Phrynarachne must be called Phrynarachnini Simon, 1895. Many of Simon’s large tribes of Thomisinae must be relimited, as they were based on striking adaptive characters of habitus with low phylogenetic value. Obviously no taxonomist describing new species in Phrynarachne within the last hundred years have doubted Simon’s authority in placing of this genus to Stephanopinae in the classification. According to most authors since Simon (1895a), discussing the diagnostic characters of Thomisid subfamilies, latest Ono (1988), the presence of more than one tooth on the retromargin of chelicerae has been regarded as the most important “synapomorphy” for Stephanopinae, However, this is a misinterpretation of Simon’s original characterization. Actually, he (Simon, 1895a: 1037) described several patterns of cheliceral and gnathocoxal modifications, but some of them were just deviating patterns of hairs, including heavy hairiness on the inner cheliceral groove. As a result study of three phylogenetically most important character groups, details of male and female copulatory organs, patterns of leg trichobothria and type of the trichobothrial base of the type species of Phrynarachne Thorell, 1869 and Thomisus rugosus Walckenaer in Latreille, 1819, it is obvious that Phrynarachne must be transferred to Thomisinae, where it is closely related to Synema with its revised limitation, including only the globosum-group and genera distinctly related to it. Simon (1895a: 1038), when creating this subfamily Stephanopsinae, divided it in the key according to the type of gnathocoxae to two main groups: - Stephanopseae: “laminae subrectae, extus vix emarginatae” (gnathocoxae with almost straight margin, laterally hardly bordered) - Hedaneae, Stephanopoideae & Phrynarachneae: “laminae maxillaries fere Misumeneae, leviter convergentes et extus, pone insertionem trochanteries, leviter emarginatae” (gnathocoxae almost as in Misumena, weakly convergent and on the outer face, behind the insertion of trochanterae weakly bordered). Besides this, he emphasized that leg I is longer than leg II in his Stephanopseae, while in the other three groups the second pair is either the longest or at least as long as the first pair.

My ongoing revision of the limitation of Stephanopinae has distinctly shown that all three groups of the second alternative in the key must be definitively transferred out from true Stephanopinae, where all taxa are especially characterized by very complex trichobothrial patterns on tarsi, metatarsi and tibiae, and also by the ultrastructure of the inner wall of the trichobothrial base which is widely different from that of Borboropactidae and Thomisidae. The final position of Hedanini is still unclear, as it could even represent an unnamed subfamily related to Thomisinae, but characterized by a unique type of tutaculum at the base of the male cymbium. The strictly Neotropical Stephanopoidini seems to be a tribe within Thomisinae.

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Figure 2 Phylogeny of Phrynarachnini. a-d: habitus dorsally. a: Phrynarachne rugosa ♀ Mauritius, b: P. rugosa ♂ Madagascar, c: Synema globosum ♂ Turkey, Yamanlar Dagi, d: Camaricus maugei ♀ China, Yunnan; e-h: male palpi. e. Phrynarachne rugosa Madagascar, Maroantsetra (ventral view), f: same specimen (lateral view), g: (SEM-micrograph) Cladonotula jobiensis Papua New Guinea, Goodenough Island, h: Synema globosum Turkey, Karsiyaka; i-j: female epigyne, i: Phrynarachne rugosa, Maroantsetra, j: Synema globosum Karsiyaka; k-n: SEM-micrographs of trichobothria, k: Phrynarachne decipiens Myanmar (basal plate), l: Cladonotula jobiensis (tarsal row), m: Diaea (s.lat.) sp. Madagascar, Taolagnaro, basal plate of metatarsal trichobothrium, n: Synema globosum, Karsiyaka (basal plate of tarsal trichobothrium); o-p. tibial trichobothria, o: Synema globosum Karsiyaka, p: Phrynarachne decipiens Myanmar.

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Relimitation of Hedanini Simon (1895a) diagnosed Hedaneae in comparison with Stephanopodeae and Phrynarachneae by stating that lateral eyes occupy almost the whole frontal area, and included in his Hedaneae three genera with Oriental species: Hedana L. Koch, 1874 (included by him, supposed synonym Cetuma Simon, 1885), Tharrhalea L. Koch, 1875 (included by him, supposed synonym Cerinius Thorell, 1877), Pycnaxis Simon, 1895, and purely Neotropical Erissus Simon, 1895. Later, following genera with Oriental species have been added to this tribe: Ebrechtella Dahl, 1907, Reinickella Dahl, 1907 and Sinothomisus Tang, Yin, Griswold & Peng, 2006, but also Neotropical Erissoides Mello-Leitão, 1929. Lehtinen (2004) revised Ebrechtella and Erissoides, and transferred them to Thomisinae: Misumenini. Tharrhalea is discussed below in more detail, while Hedana and Erissus are found to be polyphyletic too, but are not revised here. Taxonomy of Pycnaxis has been mostly solved, but due to its complex results, it is revised here.

Tharrhalea L. Koch, 1875 Thomisus L. Koch, 1865: 869, in part (evanidus only) non Thomisus Walckenaer, 1805 Xysticus L.Koch, 1867: 218, non Xysticus C.L. Koch, 1835 Diaea L. Koch, 1874: 565; Rainbow, 1904: 105; Kulczyński, 1911: 137; Strand, 1913: 118; Chrysanthus,1964: 41, non Diaea Thorell, 1869 Tharrhalea L. Koch, 1875: 604 fig. 47; Chrysanthus, 1964: 89; non Tharrhalea Simon, 1886, Simon, 1895 a&b, Kulczyński, 1911, or Barrion & Litsinger, 1995 Lehtinelagia Szymkowiak, 2014: 416, syn.n.

Tharrhalea has been a real problem for several reasons. There are eleven species listed in Tharrhalea, but none of the later described ones are congeneric with the type species. The present location of Bradley collection seems to be unknown and the genus has never been revised. Most misplacements were done by Simon (1895a). This was realized after careful observation of Thomisid descriptions and figures of “Die Arachniden Australiens”. 1. Tharrhalea is a senior synonym of Lehtinelagia Szymkowiak, 2014. When he revised Australian Diaea spp., the holotype of T. albipes was not even tried to locate, as it was not described in Diaea – and most of its congeners were originally in Diaea, therefore a new genus was created. 2. Tharrhalea was unfortunately placed to Hedaneae by Simon (1895a) and therefore listed by all taxonomists later in Stephanopinae. Tharrhalea has characters of Misumenini and Diaeini and could be placed in either of them; here Misumenini is selected, mainly because of the dark annulations of male legs I-II, present in all Misumenini, but absent or obscure in Diaeini. It is the dominant genus of Thomisidae in Polynesia and quite common in Melanesia and in Australia outside the desert areas. 3. It is very difficult to understand why the remaining species in catalogs have been placed to Tharrhalea – may be because the type material of the type species was not available to the authors of these species. Tharrhalea praetexta (L. Koch, 1865) (#ZMH, PTL), comb. n. ex Diaea was described from Samoa, but has later found to be a very widespread and common Polynesian-Melanesian species. It was not revised by Szymkowiak (2014), as it does not live in Australia.

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Figure 3 a-d. Massuria angulata. a-b: Thailand, Doi Suthep, a: carapace, b: abdomen, c-d: Malaysia. Perak, Tapah, c: metatarsal trichobothrium, d: tibial trichobothrial pattern.

Misplaced Tharrhalea 1. Cerinius Thorell, 1877 (a genus of true Hedanini) was erroneously regarded a junior synonym of Tharrhalea by Simon (1895a); C. fuscus Thorell, 1877(*MCSN) and C. irroratus Thorell, 1881 (*MCSN) were later erroneously listed in Tharrhalea. 2. Tharrhalea superpicta Simon, 1886, T. cerussata Simon, 1886, and T. semiargentea Simon, 1895 from Madagascar & Mauritius (all *MNHN) constitute together a new genus in a new tribe of Stephanopidae – the only African genus of that family. 3. T. bicornis Simon, 1895 (*MNHN) from Luzon is probably not even a thomisid and could not yet be exactly placed – but it is not related to Tharrhalea. 4. Tharrhalea luzonica (Karsch, 1880) was originally placed to Cerinius and Tharrhalea mariae Barrion, 1995 (*IRRI) also belongs there, although both are still listed as Tharrhalea in catalogs. 5. Females and juvenile specimens of a species closely related to #Tharrhalea maculata Kulczyński, 1911 and Massuria angulata Thorell, 1887 is rather common in Malaysia and Thailand, but I have never visited this area when males are adult. It is certainly congeneric with Massuria watari Ono, 2002 from Japan. T. variegata Kulczyński, 1911 is also congeneric with T. maculata as originally described.

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There was an unfortunate error in Lehtinen, 2004 concerning the concept of Massuria Thorell, 1887. During my visit to MCSN, several samples were checked from the Oriental region identified by Thorell as Massuria spp., none of the checked specimens were congeneric with the type species M. angulata Thorell, 1887, but appear to represent a species of Diaeini. The type material of Massuria angulata was unfortunately not seen, hence a figure of a large Diaeini sp. from Sumatra as a figure of Massuria was published (Lehtinen, 2004: fig. 14). Specimens of Massuria from Malaysia and Thailand in my collections have been thought to belong to some other thomisid genus. The first figure of a true male Massuria was first published by Ono (2002 & 2009).

Pycnaxis Simon, 1895 Coriarachne Simon, 1886: 145, non Coriarachne Thorell, 1870 Pycnaxis Simon, 1895b: 440; Simon, 1895 a: 1041 Oxyptila, Bösenberg & Strand, 1906: 259, non Ozyptila Simon, 1864 Xysticus, Simon, 1909: 142; Bristowe, 1931: 1410, non Xysticus C.L. Koch 1835 Takachihoa Ono, 1985: 29, syn.n.; Zhang, Zhu & Tso, 2006:82; Tang & Li, 2010: 84

Type species of Pycnaxis: P. guttata with juvenile holotype from Philippines, Luzon (¤ MNHN), numerous topotypical males, females and juveniles of different stages compared side by side with the type, type species of Takachihoa: Oxyptila truciformis Bösenberg & Strand, 1906 from Japan. Pycnaxis was placed to Stephanopinae Hedaneae by Simon (1895a) and later always listed there, e.g., still by Ono (1988). Listed here in Coriarachnini as a typical group. Male palpal bulbus of Pycnaxis and Takachihoa have similar curved ridge on tegulum and the habitus of species of both genera are strikingly similar. If the male of Pycnaxis guttata had been known during the description of Takachihoa, that genus hardly would have been erected. The comparison of juveniles of different stages, together with adults of both sexes from the type locality with the juvenile type is an instance of necessary “detective work” that is sometimes necessary in placing a species with juvenile holotype. In this situation, similar work was carried out in cases requiring correct assignation of juvenile types of Ocyllus binotatus and Pharta bimaculata.

Numerous transfers: Takachihoa Ono, 1985, syn.n.: type species #Pycnaxis truciformis (Bösenberg & Strand, 1906)/Japan, comb.n. ex Ozyptila = Xysticus nebulo Simon, 1909 (*MNHN) /Vietnam, syn.n., Pycnaxis nigrostriatus Simon, 1886 *MNHN)/ Vietnam, comb.n. ex Coriarachne. Pycnaxis krakatauensis (Bristowe, 1931) (*BMNH) /Krakatau, comb.n. ex Takachihoa/ Xysticus, Pycnaxis onoi Zhang, Zhu & Tso, 2006/Taiwan, Pycnaxis lamellaris Tang & Li, 2010/China, and Pycnaxis tumida Tang & Li, 2010/China, all comb.n. ex Takachihoa. There is at least one undescribed species from #Papua New Guinea.

Oriental genera transferred from Stephanopidae to Borboropactidae Wunderlich, 2004 Confused by the common tendency for strong external modifications of the body and legs, Simon (1895a) unfortunately did not notice any of the essential diagnostic characters of two entirely different groups of his Stephanopinae, which he placed to his tribe Stephanopseae. Actually, the key to his genera of Stephanopseae was primarily based on eye patterns, a character

Indian Journal of Arachnology 5 (1-2) 157 Significance of oriental taxa in phylogeny of crab spiders Pekka T. Lehtinen

Figure 4 a-r. Pycnaxis; a-f: dorsal habitus, a:♀ & c:♂ Pycnaxis guttata Philippines, Luzon, Mabitac, b: P. truciformis ♀ Vietnam, Hanoi, d: Pycnaxis n. sp. ♂ Malaysia, Pahang, Bukit Fraser, e: P. nigrostriata ♂ Malaysia, Pulau Pinang, Pinang Hill, f: P. krakatauensis ♀ Malaysia, Cameron Highlands, Bringchang; g: P. guttata ♂ Mabitac tarsus I dorsally; h: P. onoi Taiwan, Taichung, Tzuyou (basal part of embolus); i-k: epigyne, i: P. guttata Mabitac, j: P. krakatauensis Bringchang, k: P. nigrostriata Pinang Hill; l-o: SEM-micrographs of male palpi, l: Pycnaxis guttata Mabitac, m: P. krakatauensis Bukit Fraser, n: P. nigrostriata Pinang Hill, o: P. onoi Tzuyou; p-r: SEM-micrographs of embolic structures, p: P. krakatauensis Bringchang; embolus, q: P. onoi, Tzuyou distal part of embolus, r: P. nigrostriata Pinang Hill, embolic tip.

Indian Journal of Arachnology 5 (1-2) 158 Significance of oriental taxa in phylogeny of crab spiders Pekka T. Lehtinen with minimal phylogenetic value in spiders with great modifications in the shape of carapace and especially in the head region. Wunderlich (2004) created Borboropactidae as a family to include several fossil crab spiders and only Borboropactus of the extant genera. Here, this group is treated as a family of crab spiders with entirely new limitation. The best diagnostic character for this group is presence of a median apophysis articulated with tegulum. My Borboropactidae includes five recent groups, Phartini (= Epidius-clade of Benjamin et al., 2008), three new tribes Angaeini, Geraestini, Cebrenninini, and Borboropactini Wunderlich, 2004. The three first tribes consist of the most primitive groups with several characters, especially trichobothrial patterns of legs and ultra- structure of the trichobothrial base even shared with Thomisinae.

Borboropactini The tribe Borboropactini has the most complex ultrastructure of legs and also strongly tuberculate surface of the whole body and legs. The tarsal trichobothrial pattern includes a large sensory area with modified setae around the trichobothria, while metatarsi have a pair of lateral trichobothria. All digital figures and SEM-micrographs in this study have been taken from the same widespread Oriental species. It is most probably Borboropactus cinerascens (Doleschall, 1859), but no type material of this species is known to be preserved - my male specimens were compared with the holotype of B. bangkongeus Barrion, 1995. There are many females collected together with these males.

Phartini This tribe actually has no suggested tribal name as I called it simply Pharta-group (Lehtinen, 2004), while Benjamin et al. (2008) treated it as Epidius-clade in cladistic analysis. I strongly prefer Phartini as a tribal name, referring to a genus without striking autapomorphies, while Epidius represents a genus with strongly deviating, and for the group atypical habitus. Epidius Thorell, 1877 with synonyms Pothaeus Thorell, 1895 (*P. armatus Thorell, 1895 from Myanmar *BMNH, syn. n.), and Cupa Strand, 1906 (Cupa typica Bösenberg & Strand, 1906 *S♂♀ from Japan) and Pharta Thorell, 1891 [with synonym Sanmenia Song & Kim, 1992, type species Cupa zhengi Ono & Song, 1986 #PTL] consist the known genera. Epidius is a widespread Old World genus with the majority of species in the Oriental region. Male palp is sometimes very long due to long basal segments.

Cebrenninini There is a nomenclatural problem in the name of the nominal genus. Exact time of publication is not known for the two generic names published in 1887, but it is suggested here that the name Cebrenninus Simon, 1887 should be validated, as it is much used, while Ocyllus Thorell, 1887 has not been used outside catalogs after its description. The reason is that the holotype of Ocyllus binotatus Thorell,. 1887 (*MCSN) is juvenile and has not been checked before my visit to MCSN. However, a comparison of juveniles of the same stage undoubtedly prove that this juvenile specimen from Myanmar is congeneric with material of the type species of Cebrenninus, C. rugosus Simon, 1887 from Indonesia. Ono (1988) is the only author, who has presented an opinion about Ocyllus and placed it to Coriarachnini. The later described Ocyllus pallens Thorell, 1895 (*MCSN) from Myanmar is not congeneric with the type species,

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Figure 5 Borboropactus cinerascens (Doleschall, 1859) (= *B. bangkongeus Barrion, 1995!) Philippines, Luzon, Tuba, a-h. a: lateral habitus ♂ b: ♂ palp lateral, c: epigyne, d: tarsal sensory area, e: tarsal trichobothrium in sensory area, f: ♂ palp ventral, g: metatarsus I lateral, h: metatarsal lateral trichobothria. but belongs to Dietini. A side-by-side comparison has not been carried out, but it seems to be a junior synonym of either Oxytate parallela Simon, 1880 or of some other species of Oxytate described much later. Cebrenninini includes the Oriental genera *Cebrenninus Simon, 1887, *Ascurisoma Strand, 1928, Ibana Benjamin, 2014 and one undescribed genus, where Cupa kalawitana Barrion, 1995 (*IRRI) from Mindanao also must be included, together with an (# PTL) undescribed species. Simon (1887) originally listed Cebrenninus in Clubionidae s.lat. and Thorell (1890), his synonymous Libania in Heteropodoidea, but this genus has been listed in Thomisidae Stephanopinae since Simon (1895). In spite of exceptional leg pattern, this group can be included in Borboropactidae because of presence of a median apophysis, representing perhaps the most plesiomorphic group of crab spiders.

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Angaeini Angaeini includes only the single, but large and widespread genus Angaeus Thorell, 1881 with the type species A. pudicus Thorell, 1881 (*MCSN) from Ceram Island, Moluccas. Quite long time no males were known of other, mainly Oriental species (cf. Platnick, 2000), and females were even placed in Stephanopis (S. weyersi Simon, 1899) (independently synonymized by Benjamin, 2013) and juvenile specimens were even described as Misumenoides kripalaniae (Tikader, 1963) (see Lehtinen, 2004). Wunderlich (2008) included Angaeus to his Borboropactidae as the second extant genus, but did not discuss the other groups.

Angaeus Thorell, 1881 Angaeus Thorell, 1881: 346; Benjamin, 2013: 71-80 Stephanopis Simon, 1899: 98 (only S. weyersi), non Stephanopis O.Pickard-Cambridge, 1869 Misumenoides, Tikader, 1963: 257 fig. 8 a-b, non Misumenoides F.O. Pickard-Cambridge, 1900 Paraborboropactus Tang & Li, 2009, syn. Benjamin, 2013

Angaeus was long very imperfectly known, as males were known only for the type species A. pudicus Thorell, 1881 from Indonesia, Moluccas, Ceram Island, and as usual for Thorell, no figures were published for it. Several species of Angaeus belong to the dominant species of spiders in the ground layer of east and southeast Asia. They are identifiable to generic level even as small juveniles because of the characteristic dark triangular figure on abdominal dorsum.

Geraestini new tribe This group consists of at least three African genera, but is not further discussed here.

Dietinae Simon, 1895 This is based on Dieteae Simon, 1895, but first used as a subfamily name by Petrunkevitch, 1928. The existence of a subfamily Dietinae has been a much disputed taxonomic problem during almost a hundred years. Polyphyly of Dietinae was confirmed by Lehtinen (2001 lecture), due to parallel adaptation for possible movements on smooth surfaces by ultrastructure of scopular setae, which is even structurally different in different groups. In spite of that, the concept of Dietinae has quite recently been used in taxonomic papers published within the Oriental region, possibly trusting on the authority of Ono (1988, 2009), where the original limitation of Petrunkevitch, 1928 has been accepted. The Oriental tribe Dietini was transferred to Thomisidae Thomisinae as a sister group of Misumenini & Diaeini, while the tribes Alcimochthini and Amyciaeini are related with Tmarini (Lehtinen, 2001). Apyretini, Emplesiogonini, Mystariini, and Tagulini were still listed as tribes of Thomisidae: Thomisinae by Lehtinen (2004), but here they are regarded to consist together the Apyretinini s.lat. in Bomidae or Stiphropodidae. A majority of species of these genera live outside Oriental region.

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Figure 6 a-j. Angaeus; a, c-d, j: A. leucomenas. Philippines, Luzon, Caliraya, b: Angaeus sp. 2 Philippines Luzon, Caliraya, e, h-i Angaeus sp. Taiwan, f-g: A. rhombifer, Malaysia, Pulau Pinang; a: female dorsally, b-c: male palp, d: median apophysis of male palp, f: epigyne, g: tarsal organ, h: tarsal trichobothrium, i: specialized setae on metatarsal surface, j: a pair of subdistal metatarsal trichobothria.

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Boliscini, new tribe with notes on Bominae auct. Up till now, Boliscus Thorell, 1891, with type species Boliscus segnis Thorell, 1891 (*NHRS) from continental Malaysia has been included in the tribe Bomini Simon, 1895 and it has also been thought to include only a few species, one of them, the type species regarded as very widespread. Because of this misunderstanding, the valid name of the type species has been thought to be B. tuberculatus (Simon, 1886) by Ono (1984), described from . As long as the holotype of B. tuberculatus has not been compared with authoritatively identified B. segnis; I will use the specific name segnis for all specimens compared with the holotype of B. segnis. B. duricorius (Simon, 1880) from New Caledonia and B. decipiens O. P.-Cambridge, 1899 from Sri Lanka are known by females only, which may cause some problems in their revision. During the present study, numerous samples of personally collected Boliscus have been studied and compared with each other as well as with species of other genera traditionally listed in Bomini. Figures 7 a-c easily reveal that some species of Boliscus are also quite different for their patterns of abdominal modifications and color, as well as color patterns of carapace, not only in the details of the copulatory organs. The large genus Boliscus waits for revision, preferably complemented with molecular analysis. I have not studied the type material of Boliscodes amaenulus Simon, 1909 from Vietnam. The results convincingly show that Boliscus is not closely related to most genera of true Bomini (excluding Corynethrix and possibly Boliscodes) and also that there are several Oriental species in this genus, which urgently need a taxonomic revision, preferably also with molecular analysis. Unfortunately, no material of Corynethrix L. Koch, 1876 has been available for this study. The type species C. obscura L. Koch, 1876 from Queensland can be easily differentiated from Boliscus by shape of the epigynal hood, which has a posterior cavity (L. Koch, 1876 Pl. 69 fig. 8a), while that of Boliscus is faced anteriorly. Although Corynethrix seems to be a relative of Boliscus, its final phylogenetic position must wait for study of the actual material. Simon (1895:1004) listed Boliscus as a genus different from Corynethrix L. Koch,1876 and listed the latter as genus ignotus. Ono (1988) listed both, but personally studied only Boliscus from his subfamily Bominae. Boliscus is not closely related with other traditional Bominae because of several differences in phylogenetically significant structures, and also the male palp does not closely resemble that of the other genera of Bomini. Firstly, the male palpal tibial armature is not simple as in most true Bominae, but consists of both RTA and VTA. In Bomini, VTA is present; also in Avelis Simon, 1895 and only in one species, T. sanmen Song, Zhang & Zheng, 1992 from Zhejiang and Hubei, China (Song et al., 1999) in the widespread Old World genus Thomisops Karsch, 1879. The RTA of Boliscus has its distal surface modified rasp-like with parallel ridges (Figure 7k), as in the Oriental Stiphropodid genus , and also, e.g. the genus Runcinia Simon, 1875 in Thomisinae, Misumenini, but there is no VTA in Boliscus nor in any species of Bomini studied by me. The gnathocoxal strongly ramose apophyses (Figure 7o) of all studied specimens of Boliscus are unique among all spiders known to me. These structures are exactly similar in all studied specimens in all populations and thus, cannot be regarded as deformed structures as easily thought at first sight. Without molecular analysis it is still a matter of opinion whether Boliscini is listed as a group of Stiphropodidae, Bomidae or Thomisidae. Lack of spines, leg proportions and type of trichobothrial bases are in favor of Stiphropodidae; normal tarsi, lack of branched hairs, and the copulatory structures with some

Indian Journal of Arachnology 5 (1-2) 163 Significance of oriental taxa in phylogeny of crab spiders Pekka T. Lehtinen

Figure 7 Boliscus spp. a-c: habitus dorsally, a: Boliscus segnis Malaysia, Pulau Pinang, Pinang Hill, b: B. tuberculatus sensu Ono, Indonesia, Sumatera Barat, Payakumbuh, c: Taiwan, Tengchich 2040 m, d-f: B. segnis, female copulatory organs, d: large epigyne, e: small epigyne, f: vulva of large epigyne; g-k: male palpi, g: B, tuberculatus sensu Ono, Payakumbuh ventral, h: the same lateral, i: Boliscus sp. Tengchich ventral, j: the same lateral, k: Boliscus sp. Singapore, Bedok SEM-micrograph ventral; l: Boliscus sp. tarsus I dorsal SEM; m-n: tarsal trichobothrium, m: Boliscus sp. Bedok, n: B. segnis, Pulau Pinang; o: distal gnathocoxal processes, B. segnis, Pulau Pinang; p-r. SEM-micrographs of Boliscus sp. Bedok, p: two distal setae on inner cheliceral face, q: tarsal organ of leg I, r: distal metatarsal surface.

Indian Journal of Arachnology 5 (1-2) 164 Significance of oriental taxa in phylogeny of crab spiders Pekka T. Lehtinen kind of epigynal hood refer to relation with Bomidae or Thomisidae. Most branched setae on legs of Bomidae s.str. have been modified to thick setae with serrate margins, and it seems to be the sister group of Stiphropodidae. There is also a difference of habitat preference between Boliscus and Bomini, as Boliscus is frequently found in the vegetation layers, also in bushes and trees, while the majority of true Bomini are typically inhabitants of the ground layer. There is an exceptional phenomenon in the evolution of Boliscus. In several populations studied so far, e.g., in Malaysia, Pulau Pinang, Pinang Hill, Indonesia, Sumatera Barat, Payakumbuh, and Taiwan, Tengchich (2040m above sea level) there are two different size classes of adults without any specimens of intermediate size, manifested also in two size classes of copulatory organs. Both in Taiwan and in Sumatra the RTA of male palp is slightly different in large and small palp, suggesting that the two size classes obviously have evolved independently and do not just represent specimens of the same taxon with different size. My material of B. segnis from Pinang consists of females only and no essential differences have been found in structure of epigynes of different size. Ramírez (2014) did not study any real Bomini, but included Boliscus in his discussions and some cladograms, but only a few characters were studied, as Boliscus was placed in his cladograms as a sister group of Thomisus (op. cit., fig. 214 A-B on p. 315 and fig. 215 A-D on p. 317). This result is very difficult to understand as neither me nor any other arachnologist has ever found any similarities between Boliscus and Thomisus. This seems to be a further instance about serious problems in the use of traditional cladistics analysis in comparison of supraspecific taxa. The lack of all ultrastructural characters in matrices of Ramírez (2014) may also partly explain these odd results, as I have repeatedly emphasized the superiority of ultrastructural characters because they are mainly not affected by adaptations of macromorphological characters (Lehtinen, 1976: p. 187-188; 1996: p. 400), but see also abstracts in unpublished ISA congress lectures (Lehtinen, 2001, 2004, 2007, 2010, 2013). The phylogeny and limitation of Bomini is not discussed here in more detail. Nomenclatural problem for Bomidae Simon, 1895. Although I have here accepted Dahl’s (1907) family status for Bomidae, there are serious problems connected with that name, when used in any family group taxon (tribe, subfamily, or family). Bomis L. Koch, 1874 is exceptionally imperfectly known to serve as a type genus for a suprageneric taxon. It’s epigyne or vulva have never been described, and neither there are any correctly identified and published females of this genus in any collection, as the type species B. larvata from Queensland was described according to a subadult female (*ZMH). The male of the type species (L. Koch, 1876 p. 798 pl. 69, fig.4) has a palp that is fundamentally different from all species of the well known Bomine species from Africa and the Oriental region. The relationship of Bomis to well known other Bomini has never been confirmed and as no cheliceral peg teeth were mentioned in two descriptions by L. Koch, it is even possible that the only Australian genus of Bomini belongs to some other group of small crab spiders. If this would be true, some of the better known genus of the traditional Bomini, Holopelus, Thomisops, etc. should serve as the nominal genus for a still unnamed tribe. Unfortunately, three Indian species have been misplaced to Bomis (cf. Tikader, 1980), but they must be transferred to other genera, Thomisops khajuriai (Tikader, 1980) comb.n., and Holopelus bengalensis (Tikader, 1962), comb.n., both ex Bomis. A female only has been described for Bomis calcuttaensis Biswas & Mazumder, 1981, for which no type material in

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ZSI has been available, but it could probably be a senior synonym of Thomisops sanmen Song, Zhang & Zheng, 1992 from East China. Besides this, it is highly probable that the African species of Holopelus (Dippenaar-Schoeman, 1986) are not congeneric with the two known Oriental species, for which I have succeeded to collect some specimens. A Code-compliant solution for this nomenclatural problem must be found before a family-group name now based on Bomis, will become more widely used.

Stiphropodidae The results of worldwide supraspecific revision show that this group with its recent limitation is restricted to tropical and subtropical parts of Old World, but is also lacking in Australia and Pacific archipelagoes. Although the specific revisions of African and Asiatic species of “Stiphropus” s.lat. have not yet been completed, the species groups in these continents are more or less allopatric. Ono (1980) revised the Asiatic species, but put all species to Stiphropus. Some new genera in Africa might be necessary, while the Oriental species are easily grouped to south-eastern Casturopoda O. P.-Cambridge, 1885 (allopatric species in Sri Lanka, Malaysia, and Luzon) and Himalayan Apsectromerus Simon, 1885. The subfamily Pagidinae is represented in the Oriental region by three species: the type species of Pagida, P. salticiformis O. P.-Cambridge, 1883 in Sri Lanka, P. pseudorchestes (Thorell, 1890) in Sumatra, and a still unnamed species in the Indian Himalaya. Besides this, representatives of Nyctimus Thorell, 1877 have been found in Malaysia and Sumatra, new record from Sulawesi. Zametopias Thorell, 1892 with type species Z. speculator Thorell, 1892 (*MCSN) from Sumatra is a junior synonym of Nyctimus Thorell, 1877 with type species N. binotatus Thorell, 1877 (*MCSN; #PTL) from Sulawesi. They were described according to different sexes. In my opinion, Stiphropodidae with nine distinct synapomorphies is not a sister group of Thomisidae, but rather a family of Zodarioidea. The previous thomisid subfamily Stiphropodinae Simon, 1895, together with Pagidini Simon, 1895 from his Thomisinae was treated as a valid family by Lehtinen (2007), then with the following nine synapomorphies within Thomisidae sensu auct. (= crab spiders without Philodromidae). 1. all legs completely spineless (1, 2, 7, 9 & partly 3 also valid for Bominae) 2. legs IV not essentially shorter or similar in length with legs I & II, metatarsi and femora IV not short in comparison to Thomisidae s.str. 3. hair covering consists of branched setae (not simple, serrate, plumose or feathery sensu Lehtinen 1967 & 1996, non Jocqué & Dippenaar-Schoeman, 2006) 4. tarsi and metatarsi I & II, at least, have no movable joint (= tarsometatarsus), tarsi of male leg I, at least, distinctly thickened 5. tips of tarsi IV structurally different from tarsi I-II (claw structure and/or setae) 6. serrula proper either partly reduced or with exceptional sensory setae 7. male palpal tegulum semiglobular to rounded oval, not disc shaped 8. epigyne without hood, but a rounded or transversely oval pit is present 9. anterior cheliceral margin with a row or group of peg teeth The limitation of Stiphropodidae most probably must be much widened because of careful examination of various poorly known and seldom studied tropical groups of crab spiders, many of them concentrated to or at least common in the Oriental Region. However, these results will be presented with a detailed argumentation later.

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Phylogenetic summary This study was intended to solve a lot of main problems of the phylogeny of crab spiders. A lot of relimitations and redefinitions of previously described suprageneric taxa of crab spiders have been successful by study of own Oriental material compared with the holotypes from numerous museums with Oriental spider material in Europe, Asia and Australia. According to the results of the present study, the crab spiders of the world (and the Oriental region), where even almost all tribes of the main groups Thomisidae s.str., Borboropactidae s.lat, Bomidae (with still somewhat uncertain content) and Stiphropodidae are present, only the radically revised Stephanopidae s.str. is represented in the oriental region by a few species of Stephanopini, while the two other tribes are totally lacking. Several new tribes in Borboropactidae (Phartini, Angaeini, Cebrenninini) and Bomidae (Boliscini) could be created with sufficient argumentation, while the most important reasons to reject the polyphyletic Thomisid subfamily Dietinae were gathered. Parallely, the nine convincing synapomorphies for Stiphropodidae (derived from my congress lecture in Brazil, 2007) were repeated here. It could also be preliminarily stated that the phylogenetic placement of several deviating tribes of Simon, which have been always cataloged in Thomisinae, must be carefully analyzed, as it seems that the limitations of Bomidae and possibly also Stiphropodidae must be re-estimated.

Concluding remarks For study of spider taxonomy in general and also for taxonomy, evolution, and phylogeny of crab spiders of the Oriental Region is possibly the most interesting area in the world. It is still possible to find really enchanting spiders in this region all the time, and it is also important to note that many areas rich in undescribed taxa can also be found, even in the groups where relatively much collection has been done during the last 150 years by various specialists and also by collectors of more general interest. Oriental region has a mixture of Australian, Pacific, Palaearctic and Ethiopian taxa, it just awaits for new challenges. For me, Oriental region has a very special position, as I made my first tropical collecting trip here as well as most of the later ones, and I hope to have the possibility to still publish a lot of my taxonomic and phylogenetic results based on this material of the last more than 50 years study of the Oriental spider fauna, 47 years of it is also based on personally collected Oriental spider material. However, there is still enormous work left for spider taxonomists of the future, hopefully applying the most advanced methods, including phylogenomics.

ACKNOWLEDGMENTS The kind invitation of the late Dr. Ganesh Vankhede for myself and my daughter Heli to participate the 3rd Conference of Asian Society of Arachnology in Amravati, India has made possible to gather together my longtime results of the study of Oriental crab spiders and his action deserves my deep gratitude. The staff of the organization committee has helped me in all ways, including exceptional, but necessary visits to local hospitals during the conference. Ms Priyanka Hadole should be specially thanked for all her efforts, during the finishing of this manuscript. As this work has been an exceptionally large and time consuming project with minimal external funding support, it is practically impossible to list all persons and institutions which have made these results possible. I have tried to list all those who have been of pivotal help and I am also grateful to all of

Indian Journal of Arachnology 5 (1-2) 167 Significance of oriental taxa in phylogeny of crab spiders Pekka T. Lehtinen them, whose names might have been left out here. Dr. Yuri Marusik, Magadan, Russian Far East, has been an extremely important collaborator during his repeated visits to our museum by taking most of the digital photographs, helping in finishing of the plates with his technical experience, making available much material collected by him from different parts of Asia, and by giving new ideas and also important critics during numerous discussions about problems in Arachnology. Help in computers and its programs has been given by many staff persons in our university, but especially by the technician Veikko Rinne, Zoological Museum and Laboratory Engineer, Mr. Pasi Mustalahti, Computer Center. The most important persons during the actual work have been the curators and other staff of the museums, where the comparison of types with my own material and checking of other types has been carried out during my repeated visits to these museums under curatorships of several persons, all remembered here with gratitude, especially in Muséum National d´Histoire Naturelle, Paris, France, Museo Civico di Storia Naturale “Giacomo Doria”, Genoa, Italy, Naturhistoriska Riksmuseet, Stockholm, Sweden, British Museum, Natural History, London, United Kingdom, International Rice Research Institute, Los Baños, Philippines, Zoologisches Museum der Humboldt Universität, Berlin, Germany, Muséum d´Histoire Naturelle, Geneva, Switzerland, but also in other museums with less visits and listed in the main text. A part of the study of type material has been realized through loans of many of these museums. Dr. Francesco Ballarin from Verona, Italy, now in Chinese Academy of Sciences, Beijing, is especially thanked for arranging the loan of types of Dr. L. di Caporiacco from MZUF, Florence, during his research visit to our museum. Without careful analysis of the type material the results gained here could not have been possible. Besides this, many colleagues have kindly offered important material for this project, some collected by them, as well as fruitful discussions about the problems of crab spider taxonomy and also general problems of Arachnology. For Oriental material of crab spiders, Dr. Christa Deeleman-Reinhold, Malaysia, Sabah/Netherlands, Ossendrecht and Australian material essential for this study, the late Dr. Victor Hickman, Hobart, Tasmania and Dr. Jenny Shield, Bundoora, Victoria, are also specially thanked here. The list of persons essentially helping in the field work in the Oriental region is also very long and only a few are mentioned here. The list even includes ambassadors of Finland in Vietnam, Philippines, China, and Venezuela, as well as the former High Commissioner of in Australia, Mr. Joseph K. H. Koh. More detailed lists of help of numerous persons can be found in my publications dealing with spiders of the Oriental -Australian region. I am especially grateful to the late Chancellor of the University of Helsinki, Prof. Ernst Palmén, who made possible the beginning of my field work in the Oriental Region by organizing the initial project funding through Herman Rosenberg Foundation 40 years ago, and also by helping and encouraging in all ways for my projects dealing with taxonomy of tropical . Most field work in later phases was realized through self-paid excursions from congress sites after congress trips paid by the Academy of Finland or Universities of Helsinki and Turku, as taxonomy as such has never been included in preferred disciplines in funding by official sources in Finland. I am also especially grateful to my wife Maija Lehtinen, who has accepted the strange arrangement that during my long period of active retired scientist I could use the major part of my pension for funding of my continued field work in the tropics, as all economical support to retired scientists from official sources is by definition impossible in Finland, including also participation to international congresses. She has also participated to single field trips to Sri Lanka, India, Taiwan, Thailand and Malaysia, as well as to single expeditions to Society Islands (Tahiti & Moorea), Marquesas Islands, Australia, and New Caledonia, also taking care of many important factors in the planning and realization of the non-scientific aspects of the expeditions.

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