Gustavo Hormiga

Phylogenetic placement of the Tasmanian spider Acrobleps hygrophilus (Araneae, Anapidae) with comments on the evolution of the capture web in Araneoidea

This paper studies the family‐level phylogenetic placement of the conflicting Tasmanian spider genus Acrobleps using both morphological and behavioral data. We also provide a formal taxonomic revision of Acrobleps, including information on its web architecture and natural history, as well as detailed morphological information for A. hygrophilus, its only species. Acrobleps hygrophilus lacks the typical mysmenid features. Furthermore A. hygrophilus does have all typical and diagnostic characteristics of Anapidae, except for the labral spur. We also discuss two noteworthy morphological features of Acrobleps: the pore bearing depressions of the carapace and the granulated cuticle of the spinnerets. Variation in the latter feature might provide a useful phylogenetic character. Based on the results of cladistic analyses we propose the transfer of Acrobleps from the Mysmenidae to its original placement within the Anapidae. We also propose a new lineage, informally labeled as the “clawless female clade”, which includes synaphrids, cyatholipids and “symphytognathoids.” The secondary absence of the female palpal claw provides support for the “clawless female clade.” We discuss the evolution of the orb web within anapids and other symphytognathoids based on the results of our cladistic analyses. The identical bi‐dimensional webs of the anapid Elanapis and of symphytognathids have evolved independently. Finally, we comment on the implications of one of our analyses regarding araneoid web evolution. We conclude that the taxon sample included in the previous orbicularian data matrix (modified and used in this study) is adequate to test the phylogenetic placement of Acrobleps in Anapidae but insufficient to significantly assess web evolution within Araneoidea.

Undersampling bias: the null hypothesis for singleton species in tropical arthropod surveys

1. Frequency of singletons - species represented by single individuals - is anomalously high in most large tropical arthropod surveys (average, 32%). 2. We sampled 5965 adult spiders of 352 species (29% singletons) from 1 ha of lowland tropical moist forest in Guyana. 3. Four common hypotheses (small body size, male-biased sex ratio, cryptic habits, clumped distributions) failed to explain singleton frequency. Singletons are larger than other species, not gender-biased, share no particular lifestyle, and are not clumped at 0.25-1 ha scales. 4. Monte Carlo simulation of the best-fit lognormal community shows that the observed data fit a random sample from a community of approximately 700 species and 1-2 million individuals, implying approximately 4% true singleton frequency. 5. Undersampling causes systematic negative bias of species richness, and should be the default null hypothesis for singleton frequencies. 6. Drastically greater sampling intensity in tropical arthropod inventory studies is required to yield realistic species richness estimates. 7. The lognormal distribution deserves greater consideration as a richness estimator when undersampling bias is severe.

Phylogeny of Extant Nephilid Orb-Weaving Spiders (Araneae, Nephilidae): Testing Morphological and Ethological Homologies

The Pantropical spider clade Nephilidae is famous for its extreme sexual size dimorphism, for constructing the largest orb‐webs known, and for unusual sexual behaviors, which include emasculation and extreme polygamy. We synthesize the available data for the genera Nephila, Nephilengys, Herennia and Clitaetra to produce the first species level phylogeny of the family. We score 231 characters (197 morphological, 34 behavioral) for 61 taxa: 32 of the 37 known nephilid species plus two Phonognatha and one Deliochus species, 10 tetragnathid outgroups, nine araneids, and one genus each of Nesticidae, Theridiidae, Theridiosomatidae, Linyphiidae, Pimoidae, Uloboridae and Deinopidae. Four most parsimonious trees resulted, among which successive weighting preferred one ingroup topology. Neither an analysis of an alternative data set based on different morphological interpretations, nor separate analyses of morphology and behavior are superior to the total evidence analysis, which we therefore propose as the working hypothesis of nephilid relationships, and the basis for classification. Ingroup generic relationships are (Clitaetra (Herennia (Nephila, Nephilengys))). Deliochus and Phonognatha group with Araneidae rather than Nephilidae. Nephilidae is sister to all other araneoids (contra most recent literature). Ethological data, although difficult to obtain and thus frequently missing for rare taxa, are phylogenetically informative. We explore the evolution of selected morphological and behavioral characters, discuss and redefine the homology of palpal sclerites, disprove semientelegyny in spiders, trace the newly interpreted evolution of the orb web, and show that nephilid genital morphologies coevolve with sexual behaviors and extreme sexual size dimorphism. Phylogenetic interpretations of behavior suggest new insights into spider biology and avenues for future research.

A Protocol For Digesting Internal Soft Tissues And Mounting Spiders For Scanning Electron Microscopy

Abstract We describe a simple protocol for digesting the internal soft tissues of spiders using an enzyme complex known as pancreatin. This technique is preferred over digestions with caustic agents because it better preserves the cuticle surface, allowing its study by means of scanning electron or transmitted light microscopy. In addition, we describe a technique for mounting spider body parts for scanning electron microscopy using an acryloid polymer.

Phylogenomic interrogation of Arachnida reveals systemic conflicts in phylogenetic signal

Chelicerata represents one of the oldest groups of arthropods, with a fossil record extending to the Cambrian, and is sister group to the remaining extant arthropods, the mandibulates. Attempts to resolve the internal phylogeny of chelicerates have achieved little consensus, due to marked discord in both morphological and molecular hypotheses of chelicerate phylogeny. The monophyly of Arachnida, the terrestrial chelicerates, is generally accepted, but has garnered little support from molecular data, which have been limited either in breadth of taxonomic sampling or in depth of sequencing. To address the internal phylogeny of this group, we employed a phylogenomic approach, generating transcriptomic data for 17 species in combination with existing data, including two complete genomes. We analyzed multiple data sets containing up to 1,235,912 sites across 3,644 loci, using alternative approaches to optimization of matrix composition. Here, we show that phylogenetic signal for the monophyly of Arachnida is restricted to the 500 slowest-evolving genes in the data set. Accelerated evolutionary rates in Acariformes, Pseudoscorpiones, and Parasitiformes potentially engender long-branch attraction artifacts, yielding nonmonophyly of Arachnida with increasing support upon incrementing the number of concatenated genes. Mutually exclusive hypotheses are supported by locus groups of variable evolutionary rate, revealing significant conflicts in phylogenetic signal. Analyses of gene-tree discordance indicate marked incongruence in relationships among chelicerate orders, whereas derived relationships are demonstrably robust. Consistently recovered and supported relationships include the monophyly of Chelicerata, Euchelicerata, Tetrapulmonata, and all orders represented by multiple terminals. Relationships supported by subsets of slow-evolving genes include Ricinulei + Solifugae; a clade comprised of Ricinulei, Opiliones, and Solifugae; and a clade comprised of Tetrapulmonata, Scorpiones, and Pseudoscorpiones. We demonstrate that outgroup selection without regard for branch length distribution exacerbates long-branch attraction artifacts and does not mitigate gene-tree discordance, regardless of high gene representation for outgroups that are model organisms. Arachnopulmonata (new name) is proposed for the clade comprising Scorpiones + Tetrapulmonata (previously named Pulmonata).

Speciation on a Conveyor Belt: Sequential Colonization of the Hawaiian Islands by Orsonwelles Spiders (Araneae, Linyphiidae)

Spiders of the recently described linyphiid genus Orsonwelles (Araneae, Linyphiidae) are one of the most conspicuous groups of terrestrial arthropods of Hawaiian native forests. There are 13 known Orsonwelles species, and all are single- island endemics. This radiation provides an excellent example of insular gigantism. We reconstructed the cladistic relationships of Orsonwelles species using a combination of morphological and molecular characters (both mitochondrial and nuclear sequences) within a parsimony framework. We explored and quantified the contribution of different character partitions and their sensitivity to changes in the traditional parameters (gap, transition, and transversion costs). The character data show a strong phylogenetic signal, robust to parameter changes. The monophyly of the genus Orsonwelles is strongly supported. The parsimony analysis of all character evidence combined recovered a clade with of all the non-Kauai Orsonwelles species; the species from Kauai form a paraphyletic assemblage with respect to the latter former clade. The biogeographic pattern of the Hawaiian Orsonwelles species is consistent with colonization by island progression, but alternative explanations for our data exist. Although the geographic origin of the radiation remains unknown, it appears that the ancestral colonizing species arrived first on Kauai (or an older island). The ambiguity in the area cladogram (i.e., post-Oahu colonization) is not derived from conflicting or unresolved phylogenetic signal among Orsonwelles species but rather from the number of taxa on the youngest islands. Speciation in Orsonwelles occurred more often within islands (8 of the 12 cladogenic events) than between islands. A molecular clock was rejected for the sequence data. Divergence times were estimated by using the nonparametric rate smoothing method of Sanderson (1997, Mol. Biol. Evol. 14:1218-1231) and the available geological data for calibration. The results suggest that the oldest divergences of Orsonwelles spiders (on Kauai) go back about 4 million years.

Phylogenomic Analysis of Spiders Reveals Nonmonophyly of Orb Weavers

Spiders constitute one of the most successful clades of terrestrial predators. Their extraordinary diversity, paralleled only by some insects and mites, is often attributed to the use of silk, and, in one of the largest lineages, to stereotyped behaviors for building foraging webs of remarkable biomechanical properties. However, our understanding of higher-level spider relationships is poor and is largely based on morphology. Prior molecular efforts have focused on a handful of genes but have provided little resolution to key questions such as the origin of the orb weavers. We apply a next-generation sequencing approach to resolve spider phylogeny, examining the relationships among its major lineages. We further explore possible pitfalls in phylogenomic reconstruction, including missing data, unequal rates of evolution, and others. Analyses of multiple data sets all agree on the basic structure of the spider tree and all reject the long-accepted monophyly of Orbiculariae, by placing the cribellate orb weavers (Deinopoidea) with other groups and not with the ecribellate orb weavers (Araneoidea). These results imply independent origins for the two types of orb webs (cribellate and ecribellate) or a much more ancestral origin of the orb web with subsequent loss in the so-called RTA clade. Either alternative demands a major reevaluation of our current understanding of the spider evolutionary chronicle.

Tangled in a sparse spider web: single origin of orb weavers and their spinning work unravelled by denser taxonomic sampling

In order to study the tempo and the mode of spider orb web evolution and diversification, we conducted a phylogenetic analysis using six genetic markers along with a comprehensive taxon sample. The present analyses are the first to recover the monophyly of orb-weaving spiders based solely on DNA sequence data and an extensive taxon sample. We present the first dated orb weaver phylogeny. Our results suggest that orb weavers appeared by the Middle Triassic and underwent a rapid diversification during the end of the Triassic and Early Jurassic. By the second half of the Jurassic, most of the extant orb-weaving families and web designs were already present. The processes that may have given origin to this diversification of lineages and web architectures are discussed. A combination of biotic factors, such as key innovations in web design and silk composition, as well as abiotic environmental changes, may have played important roles in the diversification of orb weavers. Our analyses also show that increased taxon sampling density in both ingroups and outgroups greatly improves phylogenetic accuracy even when extensive data are missing. This effect is particularly important when addition of character data improves gene overlap.

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.

Clade stability and the addition of data: A case study from erigonine spiders (Araneae: Linyphiidae, Erigoninae)

This study presents a new phylogeny of erigonine spiders with emphasis on genera from the Neotropics. Thirty‐nine exemplar taxa representing mostly Neotropical genera were added to a global sample of 31 erigonine and 12 outgroup exemplar taxa analyzed in a previous study. These 82 taxa were coded for 176 (172 informative) mostly morphological characters. Eighty‐one characters were identical to or modified from the 73 (67 informative) characters included in a previous study; the remaining 95 characters are new. The complete data set includes 70 erigonine exemplars representing 65 genera, seven nonerigonine linyphiid exemplars, and five exemplars representing four araneoid families in the outgroup. Cladistic analysis resulted in a single most parsimonious tree (L =904, CI = 0.23, RI = 0.58; uninformative characters excluded: L = 900, CI = 0.23). This paper explores the implications of the new topology for the evolution of several characters of interest in erigonine evolution. The phylogeny implies that the desmitracheate condition is a synapomorphy of erigonines, with a reversal to the haplotracheate condition in one large clade within Erigoninae. We infer that the loss of the paracymbium in Neotropical erigonines occurred twice and may have progressed by different evolutionary pathways. Our phylogeny differs markedly from the previous cladistic hypothesis of erigonine relationships. We investigate how the addition of characters and taxa (alone and together) have altered the earlier hypothesis of erigonine phylogeny. We conclude that topological changes from the previous study to the current one are largely the result of adding and modifying characters, not adding taxa. Continuous Jackknife Function (CJF) analysis predicts that the inclusion of additional character data will continue to imply changes in the relationships among taxa in our analysis.