Petra Sierwald

Linking of digital images to phylogenetic data matrices using a morphological ontology.

Images are paramount in documentation of morphological data. Production and reproduction costs have traditionally limited how many illustrations taxonomy could afford to publish, and much comparative knowledge continues to be lost as generations turn over. Now digital images are cheaply produced and easily disseminated electronically but pose problems in maintenance, curation, sharing, and use, particularly in long-term data sets involving multiple collaborators and institutions. We propose an efficient linkage of images to phylogenetic data sets via an ontology of morphological terms; an underlying, fine-grained database of specimens, images, and associated metadata; fixation of the meaning of morphological terms (homolog names) by ostensive references to particular taxa; and formalization of images as standard views. The ontology provides the intellectual structure and fundamental design of the relationships and enables intelligent queries to populate phylogenetic data sets with images. The database itself documents primary morphological observations, their vouchers, and associated metadata, rather than the conventional data set cell, and thereby facilitates data maintenance despite character redefinition or specimen reidentification. It minimizes reexamination of specimens, loss of information or data quality, and echoes the data models of web-based repositories for images, specimens, and taxonomic names. Confusion and ambiguity in the meanings of technical morphological terms are reduced by ostensive definitions pointing to features in particular taxa, which may serve as reference for globally unique identifiers of characters. Finally, the concept of standard views (an image illustrating one or more homologs in a specific sex and life stage, in a specific orientation, using a specific device and preparation technique) enables efficient, dynamic linkage of images to the data set and automatic population of matrix cells with images independently of scoring decisions.

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 origins of the giant pill-millipedes from Madagascar (Diplopoda: Sphaerotheriida: Arthrosphaeridae)

Giant pill-millipedes (order Sphaerotheriida) are large-bodied millipedes without poison glands which can roll-up into a complete ball. Their disconnected area of distribution spanning South Africa, Madagascar, India, SE Asia, Australia and New Zealand makes them interesting model organisms for biogeographic studies. The here presented phylogeny is based on a molecular dataset covering all areas of distribution with a special focus on Madagascar, where some species of giant pill-millipedes show island gigantism, reaching the size of a baseball. For our study, two mitochondrial genes (partial 16S rRNA and COI) as well as the complete nuclear 18S rDNA were sequenced. While many recent vertebrate studies hint that the ancestors of the recent Malagasy fauna crossed the >350 km wide Mozambique Channel several times, no such crossing was discovered in the Sphaerotheriida. For the first time in a molecular phylogenetic study of soil arthropods, a Madagascar-India group, the family Arthrosphaeridae, is recovered, hinting to a Gondwanan origin of the Sphaerotheriida. The Malagasy-Indian family Arthrosphaeridae forms a monophyletic, statistically well-supported group in all obtained trees. The giant pill-millipedes from Madagascar are paraphyletic because the Malagasy genus Sphaeromimus is the sister-taxon of the Indian Arthrosphaera. In Sphaeromimus, an ecotone shift occurred only once: the spiny forest species Sphaeromimus musicus forms the sister-clade to the species collected in rainforests and littoral rainforests. The two species of the Malagasy genus Zoosphaerium which express island gigantism form a monophyletic group in some trees, but these trees lack good statistical support. Deeper nodes inside the Sphaerotheriida, like the position of the Australian genera Procyliosoma and Epicyliosoma, the Southeast Asian family Zephroniidae and the South African genus Sphaerotherium could not be resolved. This study is the first genetic study inside the order Sphaerotheriida and provides a proper basis for future molecular biogeographic studies in millipedes and soil organisms from Madagascar.

A worldwide catalog of the family Paradoxosomatidae Daday, 1889 (Diplopoda: Polydesmida)

A catalog of the family Paradoxosomatidae (order Polydesmida) is presented here listing a total of 198 genera with 975 valid species. Of these, 190 genera containing 960 valid species are arranged in three subfamilies and 22 tribes following Jeekel’s 1968 classification (with one addition by Jeekel in 1983). In addition, eight genera with 15 species are currently of “uncertain tribal position.” Eighty-five of the 198 paradoxosomatid genera are monotypic. Currently, 40 trinom- inals and one variety are accepted in the family. Furthermore, there are 83 species listed as “doubtful species”. The species Porcullosoma albipes , P. castaneum, P. connectens, P. jaujense, P. mamillatum and P. muticum are listed as new combinations, assigned to the genus Ergethus . For each tribe, genus and species bibliographic citations are given, including a citation of the original description and all significant subsequent citations in chronological order. Eighty-eight generic and 120 species- level synonymies are fully referenced, and presented in table form. Geographic information for each species is based mainly on the locality of the type specimen. An index to genus and species names is included. Electronic files of all valid genera, species, with author and year, as well as all generic and species-level synonyms, as well as subspecies are available from the authors.

Millipede Taxonomy after 250 Years: Classification and Taxonomic Practices in a Mega-Diverse yet Understudied Arthropod Group

Background The arthropod class Diplopoda is a mega-diverse group comprising >12,000 described millipede species. The history of taxonomic research within the group is tumultuous and, consequently, has yielded a questionable higher-level classification. Few higher-taxa are defined using synapomorphies, and the practice of single taxon descriptions lacking a revisionary framework has produced many monotypic taxa. Additionally, taxonomic and geographic biases render global species diversity estimations unreliable. We test whether the ordinal taxa of the Diplopoda are consistent with regards to underlying taxonomic diversity, attempt to provide estimates for global species diversity, and examine millipede taxonomic effort at a global geographic scale. Methodology/Principal Findings A taxonomic distinctness metric was employed to assess uniformity of millipede ordinal taxa. We found that ordinal-level taxa are not uniform and are likely overinflated with higher-taxa when compared to related groups. Several methods of estimating global species richness were employed (Bayesian, variation in taxonomic productivity, extrapolation from nearly fully described taxa). Two of the three methods provided estimates ranging from 13,413–16,760 species. Variations in geographic diversity show biases to North America and Europe and a paucity of works on tropical taxa. Conclusions/Significance Before taxa can be used in an extensible way, they must be definable with respect to the diversity they contain and the diagnostic characters used to delineate them. The higher classification for millipedes is shown to be problematic from a number of perspectives. Namely, the ordinal taxa are not uniform in their underlying diversity, and millipedes appear to have a disproportionate number of higher-taxa. Species diversity estimates are unreliable due to inconsistent taxonomic effort at temporal, geographic, and phylogenetic scales. Lack of knowledge concerning many millipede groups compounds these issues. Diplopods are likely not unique in this regard as these issues may persist in many other diverse yet poorly studied groups.

Common House Spiders Are Not Likely Vectors of Community-Acquired Methicillin-Resistant Staphylococcus aureus Infections

Abstract There is an increasing incidence in the number of community-acquired methicillin-resistant Staphylococcus aureus (MRSA) infections in the United States. Skin and soft tissue infections caused by MRSA are often perceived as being preceded by a spider bite. Several possibilities exist to explain this phenomenon, including 1) spiders introduce MRSA into the bite wound and thereby serve as a potential vehicle or vector for MRSA; 2) MRSA colonization is an event secondary to the spider bite; and 3) the spider bite is a misguided way for patients or their physicians to explain the initial lesion of their skin or soft tissue infection. We hypothesized that if spiders were able to serve as vehicles or vectors for MRSA infections, they would be colonized with this pathogen. To test this hypothesis, we captured common household spiders and determined the patterns of normal microbial flora isolated from them. Spiders were collected from several homes by their occupants, photographed for identification, and cultured for external and internal microbial flora. Of >100 spiders collected, none was found to carry Staphylococcus aureus or MRSA. Relatively low numbers of microbial flora were isolated, and only a single isolate with pathogenic potential in humans (Aeromonas spp.) was isolated. Common house spiders are unlikely to be a source of MRSA.

Phylogeny of the millipede order Spirobolida (Arthropoda: Diplopoda: Helminthomorpha)

This study examines relationships within the millipede order Spirobolida using an exemplar approach, sampling within families to maximize geographical and morphological diversity; due to lack of available material, Allopocockiidae and Hoffmanobolidae were not included in analyses. The focus of this study was to test monophyly of the order, the suborders, and the families of Spirobolida and to propose interfamilial relationships using morphological and molecular data in a total‐evidence approach. Both maximum‐parsimony analyses and Bayesian inference were employed to analyse two datasets consisting of combined morphological and molecular data, one aligned using progressive alignment methods and the second aligned by secondary structure models. Rhinocricidae was recovered sister to all remaining spirobolidan millipedes and is elevated to suborder status as suborder Rhinocricidea. Trigoniulidea was recovered as monophyletic as was Spirobolidea excluding Rhinocricidae; Spirobolidea is redefined to reflect this change. All previously recognized families were recovered, with the exception of Spirobolidae; in all instances, this family was paraphyletic or part of a polytomy that lacked sufficient resolution to assess its monophyly. The results reaffirm much of the existing taxonomic foundation within Spirobolida. This study provides the first phylogenetic test of higher‐level relationships within Spirobolida and will serve as a foundation for future work in this group at finer levels.

Morphological and systematic study of the tribe Australiosomatini (Diplopoda : Polydesmida : Paradoxosomatidea : Paradoxosomatidae) and a revision of the genus Australiosoma Brölemann

The collection of several paradoxosomatid species in the context of ecological studies prompted an investigation into the morphology and species-level characteristics of Australian millipedes in the tribe Australiosomatini Brolemann, 1916 (Polydesmida : Paradoxosomatidae). Three new species are described: Akamptogonus caragoon, sp. nov., Australiosoma fulbrighti, sp. nov. and Australiosoma combei, sp. nov. Notes or re-descriptions are provided for nine additional species belonging to the tribe. Scanning electron microscopy was utilised to examine details of the antennal sensory fields, the fifth sternite lamella and associated pores. The presence of the fifth sternite lamella in adult males is considered a synapomorphy for the family Paradoxosomatidae, whereas the prominent tubercle on the first femur in males (adenostyle) represents a synapomorphy of the subfamily Australiosomatinae. With the description of two new species in the genus Australiosoma Brolemann, 1913 a revision of the genus was undertaken with the purpose of constructing a species-level phylogeny. The most commonly described and utilised species-specific characteristics were scored in a data matrix and analysed using PAUP. The analysis resulted in a single, fully resolved tree of the following structure: Hoplatria clavigera ((A. clavigerum, A. inusitatum) (((A. rainbowi, A. nodulosum) A. michelseni) (A. laminatum (A. combei, A. fulbrighti))).

A review of subspecies recognition in polydesmidan millipedes (Diplopoda) with a revision of the subspecies of Euryuridae (Xystodesmoidea)

Jorgensen, M. C., Sierwald, P. & Mason‐Gamer, R. J. (2012). A review of subspecies recognition in polydesmidan millipedes (Diplopoda) with a revision of the subspecies of Euryuridae (Xystodesmoidea). —Zoologica Scripta, 42, 317–326.

Revision of the endemic giant fire millipedes of Madagascar, genus Aphistogoniulus (Diplopoda: Spirobolida: Pachybolidae)

The Malagasy fire millipede genus Aphistogoniulus (Silvestri, 1897) is revised. All previously described species, A. cowani (Butler, 1882), A. erythrocephalus (Pocock, 1893), A. hova (de Saussure & Zehntner, 1897), A. corallipes (de Saussure & Zehntner, 1902), A. sakalava (de Saussure & Zehntner, 1897), are redescribed. Four new synonyms are confirmed: Aphistogoniulus sanguinemaculatus (Silvestri, 1897) new synonym of A. cowani, A. quadridentatus (Attems, 1910) new synonym of A. erythrocephalus, A. polleni Jeekel, 1971 and A. brolemanni Jeekel, 1971 new synonyms of A. hova. Scanning electron microscopy is utilized to investigate the sexual differences on the antenna, mandible and gnathochilarium in A. cowani. The intraspecific variation of the taxonomic characters within and between different populations of A. erythrocephalus (Pocock, 1893) is examined. Five new species (A. sanguineus Wesener, n. sp., A. infernalis Wesener, n. sp., A. diabolicus Wesener, n. sp., A. aridus Wesener, n. sp., A. vampyrus Wesener, n. sp.) are described, including the first records of the genus from the Malagasy spiny forest and azonal Western deciduous forest ecosystem. A key to Aphistogoniulus species is provided.