Charles Griswold

• PhD, University of California, Berkeley
• Professor Emeritus at California Academy of Sciences

Two new genera, Tabiboka gen. nov. and Zorascar gen. nov., are described to accommodate five new Udubidae species from Madagascar: Tabiboka milleri gen. et sp. nov. (type species; ♂♀), T. milloti gen. et sp. nov. (♂♀), T. polotowae gen. et sp. nov. (♂♀), Zorascar pasunepipe gen. et sp. nov. (type species; ♂♀) and Z. pasunepomme gen. et sp. nov. (♀)...

We revise the Chilean genus Porteria, including the type species, Porteria albopunctata, and 11 new species: Porteria ajimayo sp. nov., Porteria alopobre sp. nov., Porteria ariasbohartae sp. nov., Porteria bunnyana sp. nov., Porteria contulmo sp. nov., Porteria correcaminos sp. nov., Porteria eddardstarki sp. nov., Porteria faberi sp. nov., Porteri...

Spiders are among the most diverse animals, which developed different morphological and behavioral traits for capturing prey. We studied the anatomy and functionality of the rare and apomorphic raptorial spider feet using 3D reconstruction modeling, among other imaging techniques. The evolutionary reconstruction of the raptorial feet (tarsus plus p...

The Indo-Pacific Phyxelididae, all members of the genus Vytfutia Deeleman-Rein-hold, 1986, are revised. Vytfutia bedel Deeleman-Reinhold, 1986 from Sumatra and Vytfutia pallens Deeleman-Reinhold 1989 from Borneo are redescribed, and V. labalaba, n. sp from Borneo and V. halandrefana, n. sp. from Madagascar are described. Vytfutiini remain in Phyxel...

Leptonetidae are rarely encountered spiders, usually associated with caves and mesic habitats, and are disjunctly distributed across the Holarctic. Data from ultraconserved elements (UCEs) were used in concatenated and coalescent-based analyses to estimate the phylogenetic history of the family. Our taxon sample included close outgroups, and 90% of...

The common ancestor of spiders likely used silk to line burrows or make simple webs, with specialized spinning organs and aerial webs originating with the evolution of the megadiverse "true spiders" (Araneomorphae). The base of the araneomorph tree also concentrates the greatest number of changes in respiratory structures, a character system whose...

The eight‐eyed haplogyne spider family Plectreuridae Simon, 1893 is one of the oldest of spider families, currently comprising two genera (Kibramoa and Plectreurys), 30 extant species and one subspecies. Plectreuridae have not been rigorously examined since 1958, with only three new species added to Plectreurys. This study revisited a subset of tax...

Aim As a continental island, much of the biota of New Zealand was initially thought to have been shaped by vicariance. Recent studies, however, have highlighted the role of dispersal, with some even suggesting that the entire biota is the product of dispersal events following emergence of the islands. This study focuses on the interplay between dis...

The new zoropsid spider genus Chinja Polotow & Griswold is diagnosed and described and the following two new species are described: C. chinja sp. nov. and C. scharffi sp. nov. The species were collected in the Eastern Arc Mountains of Tanzania and appear to be endemic to this region. The genus belongs to the oval-calamistrum clade (OC Clade), which...

Silk is the most recognizable trait of spiders, and silk use has changed throughout spider evolutionary history. While morphology of the adult silk spigot has been a useful character for systematics, few studies have examined the ontogeny of the spinning apparatus, and none of these included cribellate spiders. Here, we report the first published f...

Spiders may be good models for studying the evolution of genitalia given their peculiar copulatory mechanism. Gnaphosidae, in particular, have remarkable diversity in copulatory apparatus. The family contains species with bipartite and tripartite palps. Previous studies suggest intermediate palp complexity as the ancestral condition, with an evolut...

Spiders are well known for their silk and its varying use across taxa. Very few studies have examined the silk spigot ontogeny of the entire spinning field of a spider. Historically the spider phylogeny was based on morphological data and behavioral data associated with silk. Recent phylogenomics studies have shifted major paradigms in our understa...

Full ontogeny dataset of twenty-two species comprising thirteen spider families This includes average numbers of each spigot for each instar, as well as categorical data of foraging strategies, silk used, maximum number of instars and a score of the diversity of silk spigots each species possesses. 1, Wąsowska, 1977; 2, Hajer, 1991; 3, RE Alfaro, 2...

Model selection results for modes of evolution AICc and ΔAICc values for Model Selection for modes of evolution for PGLS Model, using Instar as the predictor variable, as Model Selection found Instar to be most important or significantly most important.

Model selection AICc and ΔAICc results for all independent variables AICc and ΔAICc values for all model iterations for each independent variable, by response variable.

PGLS results for Female data Full results of the PGLS and ANOVA analyses of adult female spigot numbers.

Scaled likelihood values for the nodes on each ACE phylogram The scaled likelihoods of each possible character state at each node, starting from the root towards to the tips. These correspond to the pie charts at each node in Fig. 3. Scaled likelihood at the nodes for each of the ancestral character state estimation analyses for the four independen...

Females dataset Raw data for adult Female spigot numbers, as well as the categorical codes for the independent variables: Foraging, Specific, Instar, Type, and Silk.

Second Instars dataset Raw data for second instar spiderling numbers, as well as the categorical codes for the independent variables: Foraging, Specific, Instar, Type, and Silk.

PGLS results for Second Instar data Full results of the PGLS and ANOVA analyses of second instar spigot numbers.

Substitution rate matrices for ACE of independent variables The substitution rate matrices for the four independent variables: Foraging (foraging strategy), Specific (specific foraging strategy), Silk (main type of silk used), and Type (variety of spigot types possessed) used as the models for the ancestral character estimation analyses. The rows a...

Final R codes All R codes for tests for phylogenetic signal, model selection, PGLS analyses, ACE analyses and creating and labelling the resulting phylograms.

Gnaphosidae Pocock are a very diverse spider family with remarkable spinning organ morphology. Although the family has received intense taxonomic attention in recent years, its intergeneric relationships remain obscure. A phylogenetic analysis of Gnaphosidae genera was performed to untangle the evolutionary history of the family. A matrix of 324 mo...

Rapa Nui is one of the most remote islands in the world. As a young island, its biota is a consequence of both natural dispersals over the last ~1 million years and recent human introductions. It therefore provides an opportunity to study a unique community assemblage. Here, we extract DNA from museum-preserved and newly field-collected spiders fro...

The family Drymusidae includes 16 cryptic spiders that build irregular webs in dark places. The family is distributed in South Africa, the Neotropical and Andean regions. Here, we use a molecular approach to infer the relationships of Drymusidae using three mitochondrial (COI, 16S, 12S) and three nuclear (H3, 28S, 18S) markers. Our preferred analys...

The fauna of Udubidae and Zoropsidae of Sri Lanka is reviewed, the families are diagnosed and a distribution map is provided. The udubid genus Campostichomma Karsch, 1892 is diagnosed and redescribed, C. manicatum Karsch, 1892 is redescribed, and we propose three new species, based on females: C. harasbedda sp. nov., C. mudduk sp. nov. and C. alawa...

The tip of the legs concentrates the interactions that a spider has with the substrate where it lives. We review the morphology and evolution of spider feet, discussing the functional anatomy of their articulations and proposing a coherent terminology. All spiders consistently have two tendons to operate their feet and show a stereotyped sequence o...

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 American gnaphosid genus Apopyllus Platnick & Shadab is found from southern Mexico to southern Argentina. It can be diagnosed by the complex shape of the RTA, by the membranous tegular extension, the long coiled embolus, the retrolateral incision on the cymbium, the long convoluted copulatory duct extending anteriorly to the copulatory openings...

We test the limits of the spider superfamily Araneoidea and reconstruct its interfamilial relationships using standard molecular markers. The taxon sample (363 terminals) comprises for the first time representatives of all araneoid families, including the first molecular data of the family Synaphridae. We use the resulting phylogenetic framework to...

Small animals possess intriguing morphological and behavioral traits that allow them to capture prey, including innovative structural mechanisms that produce ballistic movements by amplifying power [1-6]. Power amplification occurs when an organism produces a relatively high power output by releasing slowly stored energy almost instantaneously, res...

We evaluate the phylogenetic and biogeographical relationships of the members of the family Pettalidae (Opiliones, Cyphophthalmi), a textbook example of an ancient temperate Gondwanan taxon, by means of DNA sequence data from four markers. Taxon sampling is optimized to cover more than 70% of the described species in the family, with 117 ingroup sp...

Spiders (Order Araneae) are massively abundant generalist arthropod predators that are found in nearly every ecosystem on the planet and have persisted for over 380 million years. Spiders have long served as evolutionary models for studying complex mating and web spinning behaviors, key innovation and adaptive radiation hypotheses, and have been in...

Clustering/heatmap analysis depicting degree of shared data between each species pair Missing data reduction (matrix 3, Table 2). Degree of positive data overlap indicated by a color-coded heatmap (yellow = low, red = high); species order from right to left in the same order as listed from top to bottom on right side of figure. Lack of phylogenetic...

Clustering/heatmap analysis depicting degree of shared data between each species pair Missing data reduction (matrix 2, Table 2). Degree of positive data overlap indicated by a color-coded heatmap (yellow = low, red = high); species order from right to left in the same order as listed from top to bottom on right side of figure. Lack of phylogenetic...

Clustering/heatmap analysis depicting degree of shared data between each species pair BaCoCa reduced matrix (matrix 4, Table 2). Degree of positive data overlap indicated by a color-coded heatmap (yellow = low, red = high); species order from right to left in the same order as listed from top to bottom on right side of figure. Lack of phylogenetic...

Clustering/heatmap analysis depicting degree of shared data between each species pair Full spider ortholog matrix (matrix 1, Table 2). Degree of positive data overlap indicated by a color-coded heatmap (yellow = low, red = high); species order from right to left in the same order as listed from top to bottom on right side of figure. Lack of phyloge...

Clustering/heatmap analysis depicting degree of shared data between each species pair MARE matrix (matrix 7, Table 2). Degree of positive data overlap indicated by a color-coded heatmap (yellow = low, red = high); species order from right to left in the same order as listed from top to bottom on right side of figure. Lack of phylogenetic clustering...

Gene occupancy of matrix 3 (see Table 2) Colored squares represent partitions present in matrix for each OTU (y-axis, in descending order of OTU representation from bottom to top) and each partition or gene (x-axis, in descending order of partition representation from left to right).

Gene occupancy of matrix 7 (see Table 2) Colored squares represent partitions present in matrix for each OTU (y-axis, in descending order of OTU representation from bottom to top) and each partition or gene (x-axis, in descending order of partition representation from left to right).

Gene occupancy of matrix 5 (see Table 2) Colored squares represent partitions present in matrix for each OTU (y-axis, in descending order of OTU representation from bottom to top) and each partition or gene (x-axis, in descending order of partition representation from left to right).

Graph Level 2 Pie Chart of Molecular Function Gene Ontology molecular functions, levels 2 for OGs shared by Arthropod and Spider Core sets. Figures generated by Blast2GO analysis.

Latex source for supplemental material—including figure legends and Tables 1 and 2

Supplemental Table 3 Spider Core Ortholog Annotations. BLASTP (top hit, evalue 1E–10) of Spider Core OGs found in reference taxon (Acanthoscurria geniculata) against either (1) custom BLAST database of Stegodyphus mimosarum proteins downloaded from GenBank (BioProject Accession PRJNA222714; 27,135 protein sequences) or (2) nr, the non-redundant pro...

Clustering/heatmap analysis depicting degree of shared data between each species pair Arthropod core ortholog matrix (matrix 5, Table 2). Degree of positive data overlap indicated by a color-coded heatmap (yellow = low, red = high); species order from right to left in the same order as listed from top to bottom on right side of figure. Lack of phyl...

Gene occupancy of matrix 1 (see Table 2) Colored squares represent partitions present in matrix for each OTU (x-axis, in descending order of OTU representation from left to right) and each partition or gene (y-axis, in ascending order of partition representation).

Gene occupancy of matrix 2 (see Table 2) Colored squares represent partitions present in matrix for each OTU (x-axis, in descending order of OTU representation from left to right) and each partition or gene (y-axis, in ascending order of partition representation). Figure

Gene occupancy of matrix 4 (see Table 2) Colored squares represent partitions present in matrix for each OTU (y-axis, in descending order of OTU representation from bottom to top) and each partition or gene (x-axis, in descending order of partition representation from left to right).

Graph Level 3 Pie Chart of Molecular Function Gene Ontology molecular functions, level 3 for OGs shared by Arthropod and Spider Core sets. Figures generated by Blast2GO analysis.

Partition file for data matrix 1

Supplemental figures with legends (Table 2 omitted due to length)

Supplemental Table 2—232 pages

Spiders (Order Araneae) are massively abundant generalist arthropod predators that are found in nearly every ecosystem on the planet and have persisted for over 380 million years. Spiders have long served as evolutionary models for studying complex mating and web spinning behaviors, key innovation and adaptive radiation hypotheses, and have been in...

Spider-hunting wasps Trypoxylon (Trypargilum) lactitarse Saussure, 1867 and T. (Trypargilum) menkeanum Coville, 1982 were studied at the Organization for Tropical Studies fi eld station at La Selva, Costa Rica. The wasps nested in trap-nests with tube of 6.4, 8.0, or 9.5 mm diameter. Nest structure, cocoon morphology, and differences between male a...

Four new species of oonopid spiders are described from Madagascar, divided between the genera Silhouettella and Noideattella: S. perismontes, sp. nov., S. perisalma, sp. nov., N. omby, sp. nov., and N. sylvnata, sp. nov. A new diagnosis for Silhouettella is provided, its type species, S. curieusei, from the Seychelles is redescribed, and its morpho...

The genus Emmenomma is revised and now includes three species from Southern Chile, Argentina and Falkland Islands (Islas Malvinas). The type species Emmenomma oculatum is redescribed and considered a senior synonym of E. beauchenicum. Emmenomma oculatum obscurum is removed from synonymy with E. oculatum, raised to the species level and redescribed;...

A new genus of goblin spider from Madagascar, Volborattella Saucedo and Ubick, is proposed and its five included species newly described and illustrated: V. teresae, the type species, V. guenevera, V. nasario, V. paulyi, and V. toliara. These species differ from other oonopids in several unusual characters, especially the variously modified setae:...

Phylogenetic relationships within the superfamily Lycosoidea are investigated through the coding and analysis of character data derived from morphology, behaviour and DNA sequences. In total, 61 terminal taxa were studied, representing most of the major groups of the RTA-clade (i.e. spiders that have a retrolateral tibial apophysis on the male palp...

Although Madagascar is an ancient fragment of Gondwana, the majority of taxa studied thus far appear to have reached the island through dispersal from Cenozoic times. Ancient lineages may have experienced a different history compared to more recent Cenozoic arrivals, as such lineages would have encountered geoclimatic shifts over an extended time p...

We revise the relationships of the spider genus Cryptothele after reexamination of the morphology of the spinnerets, leg tarsal claws and maxillae with scanning electron microscopy. Cryptothele species have a particular conformation of the spinning field of the anterior lateral spigots that is typical of zodariids and close relatives: the field of...

Ilodesmus margaritiferus (Gervais), 1836 is redescribed based on several male and female specimens from Mount Makiling, Laguna Province, Luzon Island, Philippines. Detailed illustrations of the head, antennae, gnathochilarium, mandibles, rings, spiracles, legs, gonopods and eggs are provided. Platyrhacus philippinorum Chamberlin, 1921 and Polydesmu...

A new genus of Theridiosomatidae from the Philippines is described based on male and female characters: Tagalogonia gen. nov. Two new species are described for the genus: Tagalogonia banahaw sp. nov. and Tagalogonia isarog sp. nov. A new species of Coddingtonia Miller, Griswold & Yin, 2009 is described based on males and females from Malaysia: C. a...

The orb-weaving spiders (Orbiculariae) comprise more than 25% of the approximately 44,000 known living spider species and produce a remarkable variety of webs. The wheel-shaped orb web is primitive to this clade, but most Orbiculariae make webs hardly recognizable as orbs. Orb-weavers date at least to the Jurassic. With no evidence for convergence...

The genus Dolomedes Latreille, 1804 present a very wide range of distribution on every continent except Antarctica (Platnick 2013) However, only one species has been reported from Madagascar, Dolomedes saccalavus Strand, 1907, which is unfortunately based on a juvenile specimen and was destroyed in wartime. We consider this a nomen dubium.

Spiders, like other Madagascan organisms, exhibit remarkable richness and endemism (Griswold 2003). Whereas there are only just over 550 species known from Madagascar (Platnick 2013), the rate of new species discovery suggests that the total may be much higher. In a comprehensive study of the Madagascar spider fauna, Griswold (2003) found that 29%...

Phylogenetic relationships among archaeid spider lineages, as well as the placement of archaeids within the Araneomorphae, present a problem in the systematics of spiders. We investigate these relationships by broadly sampling taxa from the Araneomorphae and superfamily Palpimanoidea, as well as from extant and fossil archaeid lineages. Using parsi...

Incorporation of fossils into biogeographic studies can have a profound effect on the conclusions that result, particularly when fossil ranges are non-overlapping with extant ranges. This is the case in archaeid spiders, where there are known fossils from the Northern Hemisphere, yet all living members are restricted to the Southern Hemisphere. To...

The forest biota of Africa is a biological diversity hotspot reflecting a heterogeneous landscape, suggesting centers of endemism and strong selective gradients in species richness. The existence of a disjunct, cool-temperate and tropical montane region has been described as ‘archipelago-like’, which has led to high endemic diversity across many or...

The new spider genus and species Trogloraptor marchingtoni Griswold, Audisio & Ledford is described as the type of the new family Trogloraptoridae. The oblique membranous division of the basal segment of the anterior lateral spinnerets of Trogloraptor suggests that this haplogyne family is the sister group of the other Dysderoidea (Dysderidae, Oono...

The family Eresidae C. L. Koch, 1850 is reviewed at the genus level. The family comprises nine genera including one new genus. They are: Adonea Simon, 1873, Dorceus C. L. Koch, 1846, Dresserus Simon, 1876, Eresus Walckenaer, 1805, Gandanameno Lehtinen, 1967, Loureedia gen. n., ParadoneaLawrence, 1968, Seothyra Purcell, 1903, and Stegodyphus Simon,...

Two new genera of goblin spiders from Madagascar, Noideattella, new genus, and Tolegnaro, new genus, are described with 11 and two species respectively. Noideattella includes N. assumptia, new combination (transferred from Silhouettella), and 10 new species: N. amboa, N. famafa, N. fantara, N. farihy, N. gamela, N. lakana, N. mamba, N. saka, N. tan...

We briefly review the potential history of Madagascar as either a Darwinian or a Wallacean island, summarize the phylogenetic evidence regarding the biogeography of Madagascar spiders, examine the dispersal history of the Madagascar Phyxelididae, and monograph the family in Madagascar. Molecular phylogenetic analyses for 32 Malagasy phyxelidid exem...

Based on a survey of a wide variety of oonopid genera and outgroups, we hypothesize new synapomorphies uniting the Oonopidae (minus the South African genus Calculus Purcell, which is transferred to the Orsolobidae). The groundplan of the tarsal organ in Oonopidae is hypothe sized to be an exposed organ with a distinctive, longitudinal ridge origina...

The spider genus Tayshaneta is revised based on results from a three gene phylogenetic analysis (Ledford et al. 2011) and a comprehensive morphological survey using scanning electron (SEM) and compound light microscopy. The morphology and relationships within Tayshaneta are discussed and five species-groups are supported by phylogenetic analyses: t...

A new oonopid genus endemic to Madagascar, Molotra, is proposed and its six included species are newly described and illustrated: M. molotra, the type species, M. ninae, M. katarinae, M. suzannae, M. tsingy, and M. milloti. Molotra is defined on the basis of its unusual genitalia. The male has the palpal bulb terminating in broad, liplike lobes, un...