Phylogeny and biogeography of the ancient spider family Filistatidae (Araneae) is consistent both with long-distance dispersal and vicariance following continental drift.

Filistatids, the crevice weavers, are an ancient family of cribellate spiders without extant close relatives. As one of the first lineages of araneomorph spiders, they present a complicated mixture of primitive and derived characters that make them a key taxon to elucidate the phylogeny of spiders, as well as the evolution of phenotypic characters in this group. Their moderate diversity (187 species in 19 genera) is distributed mainly in arid and semi-arid subtropical zones of all continents, except Antarctica. The objective of this paper is to generate a comprehensive phylogenetic hypothesis for this family to advance the understanding of its morphological evolution and biogeography, as well as lay the basis for a natural classification scheme. By studying the morphology using optical and electronic microscopy techniques, we produced a matrix of 302 morphological characters coded for a sample of 103 species of filistatids chosen to represent the phylogenetic diversity of the family. In addition, we included sequences of four molecular markers (COI, 16S, H3 and 28S; 3787 aligned positions) of 70 filistatid species. The analysis of the data (morphological, molecular, and combined) consistently indicates the separation of the Filistatidae into two subfamilies, Prithinae and Filistatinae, in addition to supporting several groups of genera: Filistata, Zaitunia and an undescribed genus from Madagascar; Sahastata and Kukulcania; all Prithinae except Filistatinella and Microfilistata; Antilloides and Filistatoides; a large Old World group including Pritha, Tricalamus, Afrofilistata, Labahitha, Yardiella, Wandella and putative new genera; and a South American group formed by Lihuelistata, Pikelinia and Misionella. Pholcoides is transferred to Filistatinae and Microfilistata is transferred to Prithinae, and each represents the sister group to the remaining genera of its own subfamily. Most genera are valid, although Pikelinia is paraphyletic with respect to Misionella, so we consider the two genera as synonyms and propose a few new generic combinations. Considering the new phylogenetic hypothesis, we discuss the evolution of some morphological character systems and the biogeography of the family. The ages of divergence between clades were estimated using a total-evidence tip-dating approach by including fossils of Filistatidae and early spider clades; this approach resulted in younger age estimates than those obtained with traditional node-dating. Filistatidae is an ancient family that started diversifying in the Mesozoic and most genera date to the Cretaceous. Clades displaying transcontinental distributions were most likely affected by continental drift, but at least one clade shows unequivocal signs of transoceanic long-distance dispersal.

Switching identities: a revision of the Afrotropical spider genus Carteronius Simon, 1897 (Araneae, Corinnidae), senior synonym of Mandaneta Strand, 1932, with a new genus of the Pronophaea group.

The genus Carteronius Simon, 1897 is transferred from Clubionidae to Corinnidae and recognized as the senior synonym of Mandaneta Strand, 1932, being the oldest available name for the pre-occupied Mandane Karsch, 1880. Upon comparing the respective type specimens, the type species of Carteronius and the type species of Mandaneta were found to represent the same species. Whence the type species Carteronius helluo Simon, 1896, is considered a junior synonym of the type species Mandaneta sudana (Karsch, 1880). Three other species of Carteronius are transferred to Donuea Strand, 1932 (Corinnidae): D. fuscus (Simon, 1896) comb. nov. from Mauritius, D. vittiger (Simon, 1896) comb. nov. and D. argenticomus (Keyserling, 1877) comb. nov., both from Madagascar. The type species, Carteronius sudanus comb. nov., from Côte d'Ivoire, Democratic Republic of the Congo, Ghana, Guinea, Sierra Leone, and C. gentilis (Simon, 1909) comb. nov., from Equatorial Guinea and Cameroon (the latter transferred from Procopius Thorell, 1899) are redescribed, and the female of C. gentilis is described for the first time. Six new species of Carteronius are described: C. ashanti Bonaldo & Silva-Junior sp. nov. from Ghana, C. myene Bonaldo & Labarque sp. nov., and C. simoni Bonaldo & Shimano sp. nov. from Gabon, C. lumumba Bonaldo & Ramírez sp. nov. from Cameroon, Gabon and the Democratic Republic of the Congo, and C. arboreus Bonaldo & Haddad sp. nov. and C. teke Bonaldo & Bosselaers sp. nov. from the Democratic Republic of the Congo. A key and distribution maps to all eight species in the genus are presented. The related Bunyoronius Bonaldo, Ramírez & Haddad gen. nov. is proposed to include B. femoralis Bonaldo, Ramírez & Haddad sp. nov. from the Central African Republic, Uganda, and Rwanda.

A taxonomic revision of the spider genus Meriola Banks (Araneae: Trachelidae).

The genus Meriola Banks currently includes 24 known species of spiders distributed across America, especially in southern South America. They have a nearly straight and narrower posterior eye row compared to other American genera of Trachelidae, and elongated and sharply tipped ventral leg cuspules. The study of specimens of Meriola available in collections revealed two undescribed species, M. avalosi sp. nov. and M. peras sp. nov., and the previously unknown females of M. balcarce Platnick Ewing and M. quilicura Platnick Ewing, all of which are described here. The female of M. lineolata (Mello-Leitão) comb. nov., transferred from Cetonana Strand, is described for the first time. Two further new combinations are proposed here: M. macrocephala (Nicolet) comb. nov., transferred from Trachelopachys Simon, and M. setosa (Simon) comb. nov., transferred from Cetonana; these two species are also considered senior synonyms of M. barrosi (Mello-Leitão) and M. hyltonae (Mello-Leitão), respectively. Additionally, a new sex matching of M. ramirezi Platnick Ewing and M. davidi Grismado is proposed, with a redescription of the female of M. ramirezi provided. New geographical records are provided for all of the species, with images and a revised diagnosis for the genus and the previously known species.

Revision and phylogeny of the genus Andocaeculus (Acari: Caeculidae) I: the A. weyrauchi species group.

Two new caeculid mite species, Andocaeculus beatrizrosso sp. nov. and Andocaeculus burmeisteri sp. nov., are described and A. weyrauchi (Franz, 1964) is redescribed based on material collected at the type locality. All post-larval stages are described for A. weyrauchi and Andocaeculus beatrizrosso sp. nov. and stochastic variation in the idiosomal and appendages chaetotaxy is considered. A clade of Andocaeculus containing the three species (the A. weyrauchi group) is established based on morphological characters, and confirmed with a Bayesian phylogenetic analysis of sequences from the CO1 marker. As result of the same analysis, the absence of the (st) pair of setae on leg II is proposed as a derived condition for the genus Andocaeculus, and the presence of the φ solenidion on leg IV is a derived condition for some Andocaeculus species of the A. weyrauchi species group.

Sequence Capture Phylogenomics of True Spiders Reveals Convergent Evolution of Respiratory Systems.

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 evolution is still poorly understood, and that might be related to the evolution of silk glands. Emphasizing a dense sampling of multiple araneomorph lineages where tracheal systems likely originated, we gathered genomic-scale data and reconstructed a phylogeny of true spiders. This robust phylogenomic framework was used to conduct maximum likelihood and Bayesian character evolution analyses for respiratory systems, silk glands, and aerial webs, based on a combination of original and published data. Our results indicate that in true spiders, posterior book lungs were transformed into morphologically similar tracheal systems six times independently, after the evolution of novel silk gland systems and the origin of aerial webs. From these comparative data, we put forth a novel hypothesis that early-diverging web-building spiders were faced with new energetic demands for spinning, which prompted the evolution of similar tracheal systems via convergence; we also propose tests of predictions derived from this hypothesis.[Book lungs; discrete character evolution; respiratory systems; silk; spider web evolution; ultraconserved elements.].

The fossil record of spiders revisited: implications for calibrating trees and evidence for a major faunal turnover since the Mesozoic.

Studies in evolutionary biology and biogeography increasingly rely on the estimation of dated phylogenetic trees using molecular clocks. In turn, the calibration of such clocks is critically dependent on external evidence (i.e. fossils) anchoring the ages of particular nodes to known absolute ages. In recent years, a plethora of new fossil spiders, especially from the Mesozoic, have been described, while the number of studies presenting dated spider phylogenies based on fossil calibrations increased sharply. We critically evaluate 44 of these studies, which collectively employed 67 unique fossils in 180 calibrations. Approximately 54% of these calibrations are problematic, particularly regarding unsupported assignment of fossils to extant clades (44%) and crown (rather than stem) dating (9%). Most of these cases result from an assumed equivalence between taxonomic placement of fossils and their phylogenetic position. To overcome this limitation, we extensively review the literature on fossil spiders, with a special focus on putative synapomorphies and the phylogenetic placement of fossil species with regard to their importance for calibrating higher taxa (families and above) in the spider tree of life. We provide a curated list including 41 key fossils intended to be a basis for future estimations of dated spider phylogenies. In a second step, we use a revised set of 23 calibrations to estimate a new dated spider tree of life based on transcriptomic data. The revised placement of key fossils and the new calibrated tree are used to resolve a long-standing debate in spider evolution - we tested whether there has been a major turnover in the spider fauna between the Mesozoic and Cenozoic. At least 17 (out of 117) extant families have been recorded from the Cretaceous, implying that at least 41 spider lineages in the family level or above crossed the Cretaeous-Paleogene (K-Pg) boundary. The putative phylogenetic affinities of families known only from the Mesozoic suggest that at least seven Cretaceous families appear to have no close living relatives and might represent extinct lineages. There is no unambiguous fossil evidence of the retrolateral tibial apophysis clade (RTA-clade) in the Mesozoic, although molecular clock analyses estimated the major lineages within this clade to be at least ∼100 million years old. Our review of the fossil record supports a major turnover showing that the spider faunas in the Mesozoic and the Cenozoic are very distinct at high taxonomic levels, with the Mesozoic dominated by Palpimanoidea and Synspermiata, while the Cenozoic is dominated by Araneoidea and RTA-clade spiders.

Copulatory mechanics in the wolf spider Agalenocosa pirity reveals a hidden diversity of locking systems in Lycosidae (Araneae).

Genital traits are among the fastest to evolve, and the processes that drive their evolution are intensively studied. Spiders are characterized by complex genitalia, but the functional role of the different structures during genital coupling is largely unknown. Members of one of the largest spider groups, the retrolateral tibial apophysis (RTA) clade, are characterized by a RTA on the male palp, which is thought to play a crucial role during genital coupling. However, the RTA was lost in several families including the species-rich wolf spiders (Lycosidae) leading to the hypothesis that the genital coupling is achieved by alternative mechanisms. Here, we investigate the genital interactions during copulation in the wolf spider Agalenocosa pirity (Zoicinae) on cryofixed mating pairs using electron, optical and X-ray microscopy and compare our findings with other lycosids and entelegyne spiders. We found an unprecedented coupling mechanism for lycosid spiders involving the palea and a membranous cuticle folding adjacent to the epigynal plate. Additionally, we show an uncommon coupling between the median apophysis and the contralateral genital opening, and confirmed that the terminal apophysis acts as functional conductor, as previously hypothesized for males of Zoicinae. Phylogenetic mapping of RTA indicated that the basal tibial process found in Agalenocosa is a secondary acquisition rather than a modified RTA.

The Grass was Greener: Repeated Evolution of Specialized Morphologies and Habitat Shifts in Ghost Spiders Following Grassland Expansion in South America.

While grasslands, one of Earth's major biomes, are known for their close evolutionary ties with ungulate grazers, these habitats are also paramount to the origins and diversification of other animals. Within the primarily South American spider subfamily Amaurobioidinae (Anyphaenidae), several species are found living in the continent's grasslands, with some displaying putative morphological adaptations to dwelling unnoticed in the grass blades. Herein, a dated molecular phylogeny provides the backbone for analyses revealing the ecological and morphological processes behind these spiders' grassland adaptations. The multiple switches from Patagonian forests to open habitats coincide with the expansion of South America's grasslands during the Miocene, while the specialized morphology of several grass-dwelling spiders originated at least three independent times and is best described as the result of different selective regimes operating on macroevolutionary timescales. Although grass-adapted lineages evolved towards different peaks in adaptive landscape, they all share one characteristic: an anterior narrowing of the prosoma allowing spiders to extend the first two pairs of legs, thus maintaining a slender resting posture in the grass blade. By combining phylogenetic, morphological, and biogeographic perspectives we disentangle multiple factors determining the evolution of a clade of terrestrial invertebrate predators alongside their biomes.

Revisiting the spider genus Eutichurus Simon, 1897 (Araneae, Eutichuridae): new species and complementary descriptions.

Six new species of the genus Eutichurus Simon, 1897 are described: E. murgai new species (based on male and female) and E. paredesi new species (male) from Peru; E. cumbia new species (female) and E. tequendama new species (male) from Colombia; E. yungas new species (male and female) from Bolivia, and E. nancyae new species (male and female) from Brazil. The males of E. marquesae Bonaldo, 1994, E. madre Bonaldo, 1994 and E. zarate Bonaldo, 1994 are described for the first time. Eutichurus brescoviti Bonaldo, 1994, described on males, is synonymized with E. tropicus (L. Koch, 1866), previously known only from females. New records for E. ibiuna Bonaldo, 1994, E. ravidus Simon, 1897, E. putus O. Pickard-Cambridge, 1898, E. silvae Bonaldo, 1994, E. lizeri Mello-Leitão, 1938, E. saylapampa Bonaldo, 1994, E. tropicus and E. manu Bonaldo, 1994 are presented. A key to all species of Eutichurus is provided, variation in the epigynal morphology of E. ibiuna is recorded and the fine morphology of E. marquesae is documented.

Multiple approaches to understanding the taxonomic status of an enigmatic new scorpion species of the genus Tityus (Buthidae) from the biogeographic island of Paraje Tres Cerros (Argentina).

Tityus curupi n. sp., belonging to the bolivianus complex, is described from the biogeographically distinct area of Paraje Tres Cerros in north-eastern Argentina. We also present a molecular species delimitation analysis between Tityus curupi n. sp. and its sister species Tityus uruguayensis Borelli 1901 to confirm species integrity. Furthermore, a cytogenetic analysis is presented for these two species which contain different multivalent associations in meiosis, as a consequence of chromosome rearrangements, and the highest chromosome numbers in the genus.

Dated phylogenetic studies of the southernmost American buthids (Scorpiones; Buthidae).

A dated molecular phylogeny of the southernmost American species of the family Buthidae, based on two nuclear and two mitochondrial genes, is presented. Based on this study, analyzed species of the subgenus Tityus (Archaeotityus) are neither sister to the remaining species of the genus Tityus, nor are they closely related to the New World microbuthids with decreasing neobothriotaxy. Analyzed species of the subgenus Tityus do not form a monophyletic group. Based on ancestral area estimation analyses, known geoclimatic events of the region and comparisons to the diversification processes of other epigean groups from the area, a generalized hypothesis about the patterns of historical colonization processes of the family Buthidae in southern South America is presented. Furthermore, for the first time, a Paleogene-African ingression route for the colonization of America by the family Buthidae is proposed as a plausible hypothesis.

New species and phylogenetic relationships of the spider genus Coptoprepes using morphological and sequence data (Araneae: Anyphaenidae).

We present evidence from the standard cytochrome c oxidase subunit I (COI) barcoding marker and from new collections, showing that the males and females of C. ecotono Werenkraut & Ramírez were mismatched, and describe the female of that species for the first time. An undescribed male from Chile is assigned to the new species Coptoprepes laudani, together with the female that was previously thought as C. ecotono. The matching of sexes is justified after a dual cladistics analysis of morphological and sequence data in combination. New locality data and barcoding sequences are provided for other species of Coptoprepes, all endemic of the temperate forests of Chile and adjacent Argentina. Although morphology and sequences are not conclusive on the relationships of Coptoprepes species, the sequence data suggests that the species without a retrolateral tibial apophysis may belong to an independent lineage.

Around the World in Eight Million Years: Historical Biogeography and Evolution of the Spray Zone Spider Amaurobioides (Araneae: Anyphaenidae).

Closely related organisms with transoceanic distributions have long been the focus of historical biogeography, prompting the question of whether long-distance dispersal, or tectonic-driven vicariance shaped their current distribution. Regarding the Southern Hemisphere continents, this question deals with the break-up of the Gondwanan landmass, which has also affected global wind and oceanic current patterns since the Miocene. With the advent of phylogenetic node age estimation and parametric bioinformatic advances, researchers have been able to disentangle historical evolutionary processes of taxa with greater accuracy. In this study, we used the coastal spider genus Amaurobioides to investigate the historical biogeographical and evolutionary processes that shaped the modern-day distribution of species of this exceptional genus of spiders. As the only genus of the subfamily Amaurobioidinae found on three Southern Hemisphere continents, its distribution is well-suited to study in the context of Gondwanic vicariance versus long-distance, transoceanic dispersal. Ancestral species of the genus Amaurobioides appear to have undergone several long-distance dispersal events followed by successful establishments and speciation, starting from the mid-Miocene through to the Pleistocene. The most recent common ancestor of all present-day Amaurobioides species is estimated to have originated in Africa after arriving from South America during the Miocene. From Africa the subsequent dispersals are likely to have taken place predominantly in an eastward direction. The long-distance dispersal events by Amaurobioides mostly involved transoceanic crossings, which we propose occurred by rafting, aided by the Antarctic Circumpolar Current and the West Wind Drift.

Phylogeny, species delimitation and convergence in the South American bothriurid scorpion genus Brachistosternus Pocock 1893: Integrating morphology, nuclear and mitochondrial DNA.

A phylogenetic analysis of the scorpion genus Brachistosternus Pocock, 1893 (Bothriuridae Simon, 1880) is presented, based on a dataset including 41 of the 43 described species and five outgroups, 116 morphological characters and more than 4150 base-pairs of DNA sequence from the nuclear 18S rDNA and 28S rDNA gene loci, and the mitochondrial 12S rDNA, 16S rDNA, and Cytochrome c Oxidase Subunit I gene loci. Analyses conducted using parsimony, Maximum Likelihood and Bayesian Inference were largely congruent with high support for most clades. The results confirmed the monophyly of Brachistosternus, the nominal subgenus, and subgenus Ministernus Francke, 1985, as in previous analyses based only on morphology, but differed in several other respects. Species from the plains of the Atacama Desert diverged basally whereas the high altitude Andean species radiated from a more derived ancestor, presumably as a consequence of Andean uplift and associated changes in climate. Species limits were assessed among species that contain intraspecific variation (e.g., different morphs), are difficult to separate morphologically, and/or exhibit widespread or disjunct distributions. The extent of convergence in morphological adaptation to life on sandy substrata (psammophily) and the complexity of the male genitalia, or hemispermatophores, was investigated. Psammophily evolved on at least four independent occasions. The lobe regions of the hemispermatophore increased in complexity on three independent occasions, and decreased in complexity on another three independent occasions.

First phylogenetic analysis of the family Neriidae (Diptera), with a study on the issue of scaling continuous characters.

Neriidae are a small family of acalyptratae flies, mostly distributed in the tropics. Very little is known about their biology, and the evolutionary relationships among species have never been evaluated. We perform the first comprehensive phylogenetic analysis of the family, including 48 species from all biogeographic regions inhabited, as well as five species of Micropezidae and one Cypselosomatidae as outgroups. We build a morphological data matrix of 194 characters, including 72 continuous characters. We first explore ways to deal with the issue of scaling continuous characters, including rescaling ranges to unity and using implied weighting. We find that both strategies result in very different phylogenetic hypotheses, and that implied weighting reduces the problem of scaling, but only partially. Furthermore, using implied weighting after rescaling characters improves the congruence between partitions and results in higher values of group support. With respect to the Neriidae, we confirm the monophyly of the family and of most its genera, although we do not obtain any of the currently accepted suprageneric groups. We propose to restrict the Eoneria and Nerius groups exclusively to the Neotropical fauna, and synonymize Glyphidops subgenus Oncopsia Enderlein with Glyphidops subgenus Glyphidops Enderlein, eliminating the subgeneric divisions. This revised phylogeny presents a striking biogeographic consistency, and shows that previous main divisions of the family were based on events of convergence.

First Harvestman Record for the Juan Fernández Archipelago, Chile, with Morphological Notes on Acropsopilio chilensis (Opiliones: Caddidae: Acroposopilioninae).

Acropsopilio chilensis Silvestri, 1904 (Eupnoi: Caddidae: Acropsopilioninae), is recorded for Robinson Crusoe Island, Chile. This is the first harvestman species recorded for the Juan Fernández Archipelago and also the first extra-continental record for this species. During the comparison with continental co-specific specimens, some previously unknown, remarkable morphological characteristics were discovered, among them: the absence of ovipositor seminal receptacles and tracheal system, small and probably imperforate spiracles and the presence of a subdistal spiny structure, maybe a stylus, in the major branch of the penis. 

Calculating structural complexity in phylogenies using ancestral ontologies.

Complexity is an important aspect of evolutionary biology, but there are many reasonable concepts of complexity, and its objective measurement is an elusive matter. Here we develop a simple measure of complexity based on counts of elements, incorporating the hierarchical information as represented in anatomical ontologies. Neomorphic and transformational characters are used to identify novelties and individuated morphological regions, respectively. By linking the characters to terms in an anatomical ontology a node-driven approach is implemented, where a node ontology and a complexity score are inferred from the optimization of individual characters on each ancestral or terminal node. From the atomized vector of character scorings, the anatomical ontology is used to integrate the hierarchical structure of morphology in terminals and ancestors. These node ontologies are used to calculate a measure of complexity that can be traced on phylogenetic trees and is harmonious with usual phylogenetic operations. This strategy is compared with a terminal-driven approach, in which the complexity scores are calculated only for terminals, and optimized as a continuous character on the internal nodes. These ideas are applied to a real dataset of 166 araneomorph spider species scored for 393 characters, using Spider Ontology (SPD, https://bioportal.bioontology.org/ontologies/SPD); complexity scores and transitions are calculated for each node and branch, respectively. This result in a distribution of transitions skewed towards simplification; the transitions in complexity have no apparent correlation with character branch lengths. The node-driven and terminal-driven estimations are generally correlated in the complexity scores, but have higher divergence in the transition values. The structure of the ontology is used to provide complexity scores for organ systems and body parts of the focal groups.

New species of the spider genera Aysenia and Aysenoides from Chile and Argentina: description and phylogenetic relationships (Araneae: Anyphaenidae, Amaurobioidinae).

New spider species of the genera Aysenia Tullgren and Aysenoides Ramírez are described and their phylogenetic relationships discussed. The new species Aysenia paposo, from the coastal desert in northern Chile is sister to Aysenia araucana Ramírez. The diagnosis of Aysenia araucana is updated and new somatic variability is reported for the species. We present new records for other species of Aysenia and Aysenoides. The new species Aysenoides simoi, from temperate forests in Chile and adjacent Argentina is sister to Aysenoides nahuel. The phylogenetic analysis confirmed the monophyly of both genera. The support values of the genera are relatively high, but some internal branches show low support values. The genus Aysenia is supported by three synapomorphies, two of these from leg spination and one from the male genitalia. Aysenoides is supported by three synapomorphies from male and female genitalia.

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.

Resampling measures of group support: a reply to Grant and Kluge.

Grant and Kluge (2003) associated resampling measures of group support with the aim of evaluating statistical stability, confidence, or the probability of recovering a true phylogenetic group. This interpretation is not necessary to methods such as jackknifing or bootstrapping, which are better interpreted as measures of support from the current dataset. Grant and Kluge only accepted the absolute Bremer value as a measure of group support, and considered resampling methods as irrelevant to phylogenetic inference. It is shown that under simple circumstances resampling indices better reflect the degree of support than Bremer values. Grant and Kluge associated the resampling methods (and the use of measures of group support in general) with what they call a "verificationist agenda", where strongly supported groups are first detected, and then protected against additional testing. They propose that identifying weakly supported groups, and then concentrating additional tests on them, will better serve science. Both programs are actually equivalent, and inert as to the selection of methods to estimate group support. The ranking of groups under a range of resampling strength is proposed as an additional criterion to evaluate resampling methods. A reexamination of the slope of symmetric resampling frequency as a function of resampling strength suggest that slopes can be problematic as well as a measure of group support.