A revision of the spider genus Orthobula Simon, 1897 (Araneae: Trachelidae) in the Afrotropical Region. I. Continental species.

The continental Afrotropical species of the spider genus Orthobula Simon, 1897 are revised. The three currently known species, O. calceata Simon, 1897 (West Africa and western Congo Basin), O. milloti Caporiacco, 1949 (Kenya) and O. radiata Simon, 1897 (southern Africa), are redescribed, including the first descriptions of the males of O. calceata and females of O. radiata. Three new species are described from both sexes: O. aethiopica sp. nov. from Ethiopia, O. arca sp. nov. from South Africa, and O. marusiki sp. nov. from West Africa and the western Congo Basin. All of the species are obligate ground-dwellers, being particularly common in the leaf litter of woody plants, except for O. arca sp. nov., which prefers open grasslands.

The guardstone spiders of the genus Phonotimpus Gertsch & Davis (Araneae: Phrurolithidae) from northeastern Mexico.

The American genus Phonotimpus Gertsch & Davis, 1940 (Araneae, Phrurolithidae) was described on the basis of two species from northern Mexico. Recently, four species were described from Central and South Mexico and one species was transferred to this genus. Here we describe the males of Phonotimpus separatus Gertsch & Davis (the type species) and P. eutypus Gertsch & Davis, both previously known only from female specimens. Moreover, we describe 25 new species from Northeast Mexico (Nuevo León, Tamaulipas, and San Luis Potosí): P. ahuacatlan sp. nov., P. arcitos sp. nov., P. boneti sp. nov., P. calenturas sp. nov., P. chipinque sp. nov., P. cielo sp. nov., P. cima sp. nov., P. cuauhtemoc sp. nov., P. cumbres sp. nov., P. elviejo sp. nov., P. escondida sp. nov., P. farias sp. nov., P. frio sp. nov., P. gertschi sp. nov., P. llera sp. nov., P. perra sp. nov., P. pozas sp. nov., P. puente sp. nov., P. revilla sp. nov., P. sanpedro sp. nov., P. vacas sp. nov., P. valles sp. nov., P. taman sp. nov., P. tetrico sp. nov. and P. xilitla sp. nov. Furthermore, we propose five species groups that include almost all of the new species and all the species described to date.

Capobula gen. nov., a new Afrotropical dark sac spider genus related to Orthobula Simon, 1897 (Araneae: Trachelidae).

A new genus of the spider family Trachelidae L. Koch, 1872 from the Afrotropical Region is described. Capobula gen. nov. is represented by five species, known from South Africa and Lesotho only. Adults of both sexes of Orthobula infima Simon, 1896a, which is widely distributed in the Western Cape, South Africa, are described for the first time, and this species is transferred to Capobula gen. nov. as its type species. Four new species are described: C. capensis spec. nov. and C. neethlingi spec. nov. (South Africa: Western Cape), C. montana spec. nov. (Lesotho and South Africa: Eastern Cape, Free State and KwaZulu-Natal) and C. ukhahlamba spec. nov. (South Africa: KwaZulu-Natal). A phylogenetic analysis based on the cytochrome oxidase subunit I (COI) gene, including 14 genera of Trachelidae, one genus of Clubionidae Wagner, 1887 and three genera of Phrurolithidae Banks, 1892, supports the placement of Capobula gen. nov. in Trachelidae, with Orthobula Simon, 1897 as its likely closest relative.

The spider tree of life: phylogeny of Araneae based on target-gene analyses from an extensive taxon sampling.

We present a phylogenetic analysis of spiders using a dataset of 932 spider species, representing 115 families (only the family Synaphridae is unrepresented), 700 known genera, and additional representatives of 26 unidentified or undescribed genera. Eleven genera of the orders Amblypygi, Palpigradi, Schizomida and Uropygi are included as outgroups. The dataset includes six markers from the mitochondrial (12S, 16S, COI) and nuclear (histone H3, 18S, 28S) genomes, and was analysed by multiple methods, including constrained analyses using a highly supported backbone tree from transcriptomic data. We recover most of the higher-level structure of the spider tree with good support, including Mesothelae, Opisthothelae, Mygalomorphae and Araneomorphae. Several of our analyses recover Hypochilidae and Filistatidae as sister groups, as suggested by previous transcriptomic analyses. The Synspermiata are robustly supported, and the families Trogloraptoridae and Caponiidae are found as sister to the Dysderoidea. Our results support the Lost Tracheae clade, including Pholcidae, Tetrablemmidae, Diguetidae, Plectreuridae and the family Pacullidae (restored status) separate from Tetrablemmidae. The Scytodoidea include Ochyroceratidae along with Sicariidae, Scytodidae, Drymusidae and Periegopidae; our results are inconclusive about the separation of these last two families. We did not recover monophyletic Austrochiloidea and Leptonetidae, but our data suggest that both groups are more closely related to the Cylindrical Gland Spigot clade rather than to Synspermiata. Palpimanoidea is not recovered by our analyses, but also not strongly contradicted. We find support for Entelegynae and Oecobioidea (Oecobiidae plus Hersiliidae), and ambiguous placement of cribellate orb-weavers, compatible with their non-monophyly. Nicodamoidea (Nicodamidae plus Megadictynidae) and Araneoidea composition and relationships are consistent with recent analyses. We did not obtain resolution for the titanoecoids (Titanoecidae and Phyxelididae), but the Retrolateral Tibial Apophysis clade is well supported. Penestomidae, and probably Homalonychidae, are part of Zodarioidea, although the latter family was set apart by recent transcriptomic analyses. Our data support a large group that we call the marronoid clade (including the families Amaurobiidae, Desidae, Dictynidae, Hahniidae, Stiphidiidae, Agelenidae and Toxopidae). The circumscription of most marronoid families is redefined here. Amaurobiidae include the Amaurobiinae and provisionally Macrobuninae. We transfer Malenellinae (Malenella, from Anyphaenidae), Chummidae (Chumma) (new syn.) and Tasmarubriinae (Tasmarubrius, Tasmabrochus and Teeatta, from Amphinectidae) to Macrobuninae. Cybaeidae are redefined to include Calymmaria, Cryphoeca, Ethobuella and Willisius (transferred from Hahniidae), and Blabomma and Yorima (transferred from Dictynidae). Cycloctenidae are redefined to include Orepukia (transferred from Agelenidae) and Pakeha and Paravoca (transferred from Amaurobiidae). Desidae are redefined to include five subfamilies: Amphinectinae, with Amphinecta, Mamoea, Maniho, Paramamoea and Rangitata (transferred from Amphinectidae); Ischaleinae, with Bakala and Manjala (transferred from Amaurobiidae) and Ischalea (transferred from Stiphidiidae); Metaltellinae, with Austmusia, Buyina, Calacadia, Cunnawarra, Jalkaraburra, Keera, Magua, Metaltella, Penaoola and Quemusia; Porteriinae (new rank), with Baiami, Cambridgea, Corasoides and Nanocambridgea (transferred from Stiphidiidae); and Desinae, with Desis, and provisionally Poaka (transferred from Amaurobiidae) and Barahna (transferred from Stiphidiidae). Argyroneta is transferred from Cybaeidae to Dictynidae. Cicurina is transferred from Dictynidae to Hahniidae. The genera Neoramia (from Agelenidae) and Aorangia, Marplesia and Neolana (from Amphinectidae) are transferred to Stiphidiidae. The family Toxopidae (restored status) includes two subfamilies: Myroinae, with Gasparia, Gohia, Hulua, Neomyro, Myro, Ommatauxesis and Otagoa (transferred from Desidae); and Toxopinae, with Midgee and Jamara, formerly Midgeeinae, new syn. (transferred from Amaurobiidae) and Hapona, Laestrygones, Lamina, Toxops and Toxopsoides (transferred from Desidae). We obtain a monophyletic Oval Calamistrum clade and Dionycha; Sparassidae, however, are not dionychans, but probably the sister group of those two clades. The composition of the Oval Calamistrum clade is confirmed (including Zoropsidae, Udubidae, Ctenidae, Oxyopidae, Senoculidae, Pisauridae, Trechaleidae, Lycosidae, Psechridae and Thomisidae), affirming previous findings on the uncertain relationships of the "ctenids" Ancylometes and Cupiennius, although a core group of Ctenidae are well supported. Our data were ambiguous as to the monophyly of Oxyopidae. In Dionycha, we found a first split of core Prodidomidae, excluding the Australian Molycriinae, which fall distantly from core prodidomids, among gnaphosoids. The rest of the dionychans form two main groups, Dionycha part A and part B. The former includes much of the Oblique Median Tapetum clade (Trochanteriidae, Gnaphosidae, Gallieniellidae, Phrurolithidae, Trachelidae, Gnaphosidae, Ammoxenidae, Lamponidae and the Molycriinae), and also Anyphaenidae and Clubionidae. Orthobula is transferred from Phrurolithidae to Trachelidae. Our data did not allow for complete resolution for the gnaphosoid families. Dionycha part B includes the families Salticidae, Eutichuridae, Miturgidae, Philodromidae, Viridasiidae, Selenopidae, Corinnidae and Xenoctenidae (new fam., including Xenoctenus, Paravulsor and Odo, transferred from Miturgidae, as well as Incasoctenus from Ctenidae). We confirm the inclusion of Zora (formerly Zoridae) within Miturgidae.

The spider tree of life: phylogeny of Araneae based on target-gene analyses from an extensive taxon sampling

We present a phylogenetic analysis of spiders using a dataset of 932 spider species, representing 115 families (only the family Synaphridae is unrepresented), 700 known genera, and additional representatives of 26 unidentified or undescribed genera. Eleven genera of the orders Amblypygi, Palpigradi, Schizomida and Uropygi are included as outgroups. The dataset includes six markers from the mitochondrial (12S, 16S, COI) and nuclear (histone H3, 18S, 28S) genomes, and was analysed by multiple methods, including constrained analyses using a highly supported backbone tree from transcriptomic data. We recover most of the higher-level structure of the spider tree with good support, including Mesothelae, Opisthothelae, Mygalomorphae and Araneomorphae. Several of our analyses recover Hypochilidae and Filistatidae as sister groups, as suggested by previous transcriptomic analyses. The Synspermiata are robustly supported, and the families Trogloraptoridae and Caponiidae are found as sister to the Dysderoidea. Our results support the Lost Tracheae clade, including Pholcidae, Tetrablemmidae, Diguetidae, Plectreuridae and the family Pacullidae (restored status) separate from Tetrablemmidae. The Scytodoidea include Ochyroceratidae along with Sicariidae, Scytodidae, Drymusidae and Periegopidae; our results are inconclusive about the separation of these last two families. We did not recover monophyletic Austrochiloidea and Leptonetidae, but our data suggest that both groups are more closely related to the Cylindrical Gland Spigot clade rather than to Synspermiata. Palpimanoidea is not recovered by our analyses, but also not strongly contradicted. We find support for Entelegynae and Oecobioidea (Oecobiidae plus Hersiliidae), and ambiguous placement of cribellate orb-weavers, compatible with their non-monophyly. Nicodamoidea (Nicodamidae plus Megadictynidae) and Araneoidea composition and relationships are consistent with recent analyses. We did not obtain resolution for the titanoecoids (Titanoecidae and Phyxelididae), but the Retrolateral Tibial Apophysis clade is well supported. Penestomidae, and probably Homalonychidae, are part of Zodarioidea, although the latter family was set apart by recent transcriptomic analyses. Our data support a large group that we call the marronoid clade (including the families Amaurobiidae, Desidae, Dictynidae, Hahniidae, Stiphidiidae, Agelenidae and Toxopidae). The circumscription of most marronoid families is redefined here. Amaurobiidae include the Amaurobiinae and provisionally Macrobuninae. We transfer Malenellinae (Malenella, from Anyphaenidae), Chummidae (Chumma) (new syn.) and Tasmarubriinae (Tasmarubrius, Tasmabrochus and Teeatta, from Amphinectidae) to Macrobuninae. Cybaeidae are redefined to include Calymmaria, Cryphoeca, Ethobuella and Willisius (transferred from Hahniidae), and Blabomma and Yorima (transferred from Dictynidae). Cycloctenidae are redefined to include Orepukia (transferred from Agelenidae) and Pakeha and Paravoca (transferred from Amaurobiidae). Desidae are redefined to include five subfamilies: Amphinectinae, with Amphinecta, Mamoea, Maniho, Paramamoea and Rangitata (transferred from Amphinectidae); Ischaleinae, with Bakala and Manjala (transferred from Amaurobiidae) and Ischalea (transferred from Stiphidiidae); Metaltellinae, with Austmusia, Buyina, Calacadia, Cunnawarra, Jalkaraburra, Keera, Magua, Metaltella, Penaoola and Quemusia; Porteriinae (new rank), with Baiami, Cambridgea, Corasoides and Nanocambridgea (transferred from Stiphidiidae); and Desinae, with Desis, and provisionally Poaka (transferred from Amaurobiidae) and Barahna (transferred from Stiphidiidae). Argyroneta is transferred from Cybaeidae to Dictynidae. Cicurina is transferred from Dictynidae to Hahniidae. The genera Neoramia (from Agelenidae) and Aorangia, Marplesia and Neolana (from Amphinectidae) are transferred to Stiphidiidae. The family Toxopidae (restored status) includes two subfamilies: Myroinae, with Gasparia, Gohia, Hulua, Neomyro, Myro, Ommatauxesis and Otagoa (transferred from Desidae); and Toxopinae, with Midgee and Jamara, formerly Midgeeinae, new syn. (transferred from Amaurobiidae) and Hapona, Laestrygones, Lamina, Toxops and Toxopsoides (transferred from Desidae). We obtain a monophyletic Oval Calamistrum clade and Dionycha; Sparassidae, however, are not dionychans, but probably the sister group of those two clades. The composition of the Oval Calamistrum clade is confirmed (including Zoropsidae, Udubidae, Ctenidae, Oxyopidae, Senoculidae, Pisauridae, Trechaleidae, Lycosidae, Psechridae and Thomisidae), affirming previous findings on the uncertain relationships of the "ctenids" Ancylometes and Cupiennius, although a core group of Ctenidae are well supported. Our data were ambiguous as to the monophyly of Oxyopidae. In Dionycha, we found a first split of core Prodidomidae, excluding the Australian Molycriinae, which fall distantly from core prodidomids, among gnaphosoids. The rest of the dionychans form two main groups, Dionycha part A and part B. The former includes much of the Oblique Median Tapetum clade (Trochanteriidae, Gnaphosidae, Gallieniellidae, Phrurolithidae, Trachelidae, Gnaphosidae, Ammoxenidae, Lamponidae and the Molycriinae), and also Anyphaenidae and Clubionidae. Orthobula is transferred from Phrurolithidae to Trachelidae. Our data did not allow for complete resolution for the gnaphosoid families. Dionycha part B includes the families Salticidae, Eutichuridae, Miturgidae, Philodromidae, Viridasiidae, Selenopidae, Corinnidae and Xenoctenidae (new fam., including Xenoctenus, Paravulsor and Odo, transferred from Miturgidae, as well as Incasoctenus from Ctenidae). We confirm the inclusion of Zora (formerly Zoridae) within Miturgidae.

IS FARRIS OPTIMIZATION PERFECT?: THREE-TAXON STATEMENTS AND MULTIPLE BRANCHING.

- The three-taxon approach to phylogenetic analysis separates the universe of cladograms into a larger number of classes of solutions showing decreasing degrees of fit to data than does conventional Farris optimization. The three-taxon approach applies to character analysis Nelson and Platnick's interpretation 2 of multiple branching in cladograms.

AN EMPIRICAL COMPARISON OF MICROCOMPUTER PARSIMONY PROGRAMS, II.

Abstract- The effectiveness and efficiency of J. S. Farris' new microcomputer parsimony program (Hennig86, version 1.5) are evaluated with reference to 60 data sets, including those used to benchmark earlier mainframe and microcomputer packages. By overcoming the arbitrary resolution and consequent redundancy problems that have plagued previously available microcomputer programs, as well as their limitations on data set size, cladogram storage space, and execution speed, Hennig86 advances enormously the accuracy and ease with which cladistic analyses can be conducted. Hennig86 has such an impressive edge in both effectiveness and efficiency that earlier parsimony programs (including those by Farris) have essentially been rendered obsolete. For exact analyses, both exhaustive and minimal options arc provided; of the options available for approximate analyses, the branch breaker (bb) used in conjunction with the mhennig* and tread commands performed best.

On the Information Content of Classifications.

Abstract- The best fit estimate of the retrospective information content of a classification is argued to be the proportion of the maximal sum of informative subgroups to which each included taxon is assigned. The best fit estimate of the prospective information content of a classification is argued to be the proportion of fully resolved dadograms prohibited by that classification. Those (wo proportions are multiplied to provide a general information index that is sensitive to both the degree of resolution and the contents of individual subgroups. That index and the best fit considerations supporting it have implications for congruence studies, consensus indices, and the choice between Adams and Nelson consensus trees.

AN EMPIRICAL COMPARISON OF MICROCOMPUTER PARSIMONY PROGRAMS.

Abstract- The effectiveness and efficiency of two microcomputer parsimony programs (Felsenstein's PHYLIP and Swofford's PAUP), and a prototype version of a third (SHEN, to be incorporated in Farris' HENNIG-86), are evaluated with reference to 35 data sets, including those previously used by Luckow and Pimentel (1985) to benchmark mainframe programs. Both PHYLIP and PAUP can be used effectively; with careful selection of options, their accuracy can equal or surpass that of older mainframe programs. PHYLIP is relatively inefficient (in use of computer time); its usefulness is also limited by the inability of its heuristic algorithms to detect multiple equally parsimonious solutions in a single run. PHYLIP's exact algorithm (branch-and-bound) and PAUP's heuristic and exact algorithms do not share the latter drawback but require users to spend unreasonable amounts of time coping with redundant output. The remarkable effectiveness of these programs offers hope that (particularly with the advent of multitasking microcomputers) exact and cost-effective solutions will be obtainable in many, if not all, cladistic analyses.