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Revision of torrent mites (Parasitengona, Torrenticolidae, Torrenticola) of the United States and Canada: 90 descriptions, molecular phylogenetics, and a key to species
expand article infoJ. Ray Fisher, Danielle M. Fisher, Michael J. Skvarla§, Whitney A. Nelson, Ashley P.G. Dowling
‡ University of Arkansas, Fayetteville, United States of America
§ U.S. Department of Agriculture, Beltsville, United States of America
Open Access

Abstract

The descriptive biology of torrent mites (Parasitengona: Torrenticolidae: Torrenticola) of North America (north of Mexico) is investigated using integrative methods. Material examined includes approximately 2,300 specimens from nearly 500 localities across the United States and Canada, and a few collections in Mexico and Central America. Species hypotheses are derived from a phylogenetic analysis of the barcoding region of cytochrome c oxidase subunit 1 (COI) for 476 specimens and supported with morphology and biogeography. Relationships between species are examined with a combined analysis of COI and two expansion regions (D2–3) of the large ribosomal subunit (28S rDNA) for 57 specimens. All previously described species from the US and Canada are examined. Our results indicate the need to synonymize four species: T. mercedensis (Marshall, 1943) is a junior synonym of T. sierrensis (Marshall, 1943); T. rectiforma Habeeb, 1974 is a junior synonym of T. ellipsoidalis (Marshall, 1943); T. neoconnexa Habeeb, 1957 is a junior synonym of T. magnexa Habeeb, 1955; and T. esbelta Cramer, 1992 is a junior synonym of T. boettgeri KO Viets, 1977. We describe 66 new species and re-describe all previously described regional species. Our findings indicate that total diversity of Torrenticola in the United States and Canada comprises 90 species, 57 known from the east and 33 from the west. We organize these species into four species complexes that include 13 identification groups. An additional 13 species do not fit within an identification group. The southern Appalachians are suspected to contain the highest concentration of remaining undescribed diversity. A key is provided to all known species in the US and Canada.

Keywords

Acari, Acariformes, Hydrachnidiae, water mites, integrative taxonomy, turbo-taxonomy

Introduction

Torrenticolidae Piersig, 1902 are known as “torrent mites” due to the typical habitat of most species—fast-flowing, rocky or sandy-bottom streams (Goldschmidt 2007, Smith et al. 2010, Proctor et al. 2015). In fact, in many streams, torrenticolids are among the most abundant of all arthropods. Adults are active predators that crawl through sandy sediment with stout legs and sclerotized bodies and roam interstitial zones in search of micro-crustaceans (Goldschmidt 2007, Smith et al. 2010, Proctor et al. 2015). The dorsal cuticle of adults is often ornamented with colorful patterns that can be useful in differentiating species, especially locally (Figure 1). Larvae are ecto-parasites of chironomids, but cannot yet be identified to the species-level, so questions such as host specificity and loss of parasitism remain open areas of investigation. This difficulty is compounded by the fact that larval and post-larval stages are difficult to link due to disparate morphologies, although future efforts employing molecular techniques may resolve this issue (e.g., Stålstedt et al. 2016).

Figure 1. 

Diversity of Torrenticola in Ouachita National Forest, Arkansas. Species from top-left to bottom-right: T. irapalpa sp. n., T. interiorensis sp. n., T. biscutella sp. n. female, T. biscutella sp. n. male, T. pearsoni sp. n., T. trimaculata Fisher & Dowling 2016, T. unimaculata sp. n., T. larvata Cherri et al. 2016, T. solisorta sp. n. Note that, for a given locality, many characters necessary for identification can be seen under low magnification. For example, species in the bottom row can be readily differentiated using color pattern. In contrast, species in the top row have similar color patterns; however, even they can be readily identified: T. irapalpa have dorsal glandularia (Dgl-4) closer together than the other species and T. biscutella have fused anterio-lateral platelets and strong sexual dimorphism (neither character is present in T. interiorensis). Only the identification of T. interiorensis is tentative based on this image, because T. neoanomala (not pictured), which are also found in this locality, can only be differentiated from T. interiorensis by comparing precise measurements of the anterio-lateral platelets, necessitating examining slide-prepared material under higher magnification. Also, note that some species can appear differently than what is depicted in species descriptions. For example, although T. pearsoni sp. n. has a nearly colorless cuticle, it can appear dark due to gut contents, as shown here. Image is a composite of stacked images from a Samsung GS7 and edited in Adobe Photoshop.

Torrenticolidae comprises two subfamilies (Wiles 1997): Testudacarinae Cook, 1974 and Torrenticolinae Piersig, 1902. Testudacarinae comprises two genera (O’Neill et al. 2016)—Debsacarus Habeeb, 1961 (one species) and Testudacarus Walter, 1928 (20 species)—and Torrenticolinae comprises five genera: Monatractides Viets, 1926 (approximately 100 species); Neoatractides Lundblad, 1941 (11 species); Pseudotorrenticola Walter, 1906 (eight species); Stygotorrenticola Pešić & Gerecke, 2014 (one species); and Torrenticola Piersig, 1896 (approximately 250 species, not including species described herein).

Torrenticola—the focus of this paper—are found worldwide, excepting Antarctica. In North America, 79 species have been described, most of which are known from Central America due to the efforts of Goldschmidt (2007). North of Mexico, only 23 Torrenticola have been described (Marshall 1929, 1930, 1933, 1943; Habeeb 1955, 1957, 1961, 1973, 1974; Crowell 1960; Fisher et al. 2015; and Cherri et al. 2016) and no comprehensive review is available. Most species are known from only one or a few specimens and sampling has been greatly limited.

The present study is the fourth in a series of ongoing taxonomic investigations of North American Torrenticolidae. The first study offered a detailed description of a single species, diagnoses of higher ranks, and a historical review (Fisher et al. 2015); the second investigated the systematics of Testudacarinae (O’Neill etal. 2016); and the third allowed an undergraduate researcher to describe a species endemic to the Ouachita Mountains, USA (Cherri et al. 2016). Herein, we describe 66 new species and re-describe 24 species, totaling 90 species. The ability to efficiently test species boundaries of such a widespread and diverse genus is a feat made possible with molecular data and by the researchers who contributed to the immense holdings of the Canadian National Collection (CNC), especially Ian Smith, who collected fresh material for molecular analyses. These collections were supplemented with local collections in Arkansas and four expeditions (Rocky Mountains, Pacific Northwest, California, and Alaska).

Methods

Curation: Mites were collected and curated using the protocol detailed by Proctor et al. (2015, p. 649–655) and summarized by Fisher et al. (2015). Specimens collected for molecular analyses were preserved in 95% ethanol, whereas others were preserved in GAW (50% glycerol, 10% glacial acetic acid, and 40% water by volume; also referred to as Koenike’s solution). Slide preparations were created after extraction of genomic DNA (see Molecular phylogenetics below) using either glycerin jelly or Hoyer’s medium. Where available, holotypes and allotypes were designated with glycerin jelly slide preparations, rather than Hoyer’s slide preparations. Unlike Hoyer’s, glycerin jelly does not obliterate body coloration, which can be important in identifying many Torrenticola. However, Hoyer’s medium has superior optical properties, which enables investigations at higher magnification (i.e., greater than 400×). Therefore, when available, some paratypes were prepared with Hoyer’s medium.

Geographic coverage: North America, north of Mexico (Figure 2). Importantly, Alaska, which is not represented in Figure 2, was sampled for water mites, but no Torrenticolidae were found. Similarly, the Great Plains region was sampled, but few Torrenticola specimens were found. These regions likely represent areas where Torrenticola, which appear to prefer cool, clean, mountainous streams, are not present or at least not common.

Figure 2. 

Summary of all sampling localities. 327 red dots represent material examined; 14 blue diamonds represent published accounts where material was not available for direct examination. Each of the 341 localities represent a generalized locality, not an individual collection event, as multiple events often occured in close proximity. In total, this map represents nearly 500 collection events. Note the absence of material from the Great Plains (e.g., Oklahoma, Kansas, Nebraska, and the Dakotas) and Alaska; these are areas that were sampled for water mites, but Torrenticola were not found.

Terminology: We follow the terminology of Fisher et al. (2015), who modified Goldschmidt (2007) and Proctor et al. (2015), with the addition of a new term, “anterior venter”, used to describe the distance between the gnathosomal bay to the genital field (Figure 6C). Colorful species generally have dorsal shades that ranges from bluish to orangish, and a medial stripe that ranges from orangish to reddish. We use the following terminology regarding the darker shade: navy blue – bluish-purple – purple – reddish-purple – orange (Figure 3).

Figure 3. 

Terminology for coloration: blue – bluish-purple – purple-pink – reddish-purple – orange. Species depicted from top-left to bottom-right: T. gorti; T. erectirostra; T. biscutella; T. neoanomala; T. nigroalba; T. gorti; T. skvarlai; T. magnexa, and T. delicatexa.

Images: Color micrographs of slide preparations were taken with a Leica DFC 300× camera using Leica Application Suite software and montaged with Helicon Focus 6 to ensure certain structures (e.g., glandularia) were visible. Resulting image stacks were then edited in Adobe Photoshop CS6 to remove debris, repair damaged specimens, and remove structures that are unnecessary for identification. For the latter, legs were removed from the venter and most setae were removed from the pedipalp. In addition to these structures not being useful for identification, they are also not depicted accurately with photographs. Final images were compared with the specimen and edited accordingly to best approximate realistic coloration. In some cases, specimens prepared in glycerin jelly (i.e., coloration preserved) were not available and only the Hoyer’s slide preparations (i.e., coloration destroyed) were available. For these, micrographs were taken of the Hoyer’s slides and coloration was added in Photoshop (Figure 69, 71). Note that color micrographs depict the best representation of specific characters (venter, dorsum, pedipalp, subcapitulum), so a given figure may be a composite of multiple specimens (e.g., dorsum and subcapitulum of specimen A; venter of specimen B).

Descriptions: The descriptions contained herein are streamlined to include information that best diagnoses a given species, so we depart from the standard of previous descriptions in two important ways. First, we depict species with photomicrographs rather than line drawings, which greatly reduces the time necessary for a given description. Although this is not generally possible with mites, torrenticolids lend themselves to such representation because the characters needed to diagnose species, such as color pattern, are viewable from such images—a condition not common in most mite groups. Second, we depart from the suggestion of Goldschmidt (2007) and recent torrenticolid descriptions (e.g., Pešić 2014; Pešić and Gerecke 2014; Pešić and Smit 2014) in that we do not depict the ejaculatory complex or legs as these systems were not useful in differentiating species that were easily distinguished by other systems (e.g., color, various measurements and proportions). In fact, in those cases where two or more species were difficult to differentiate using our methods (e.g., Miniforma Group), we examined the ejaculatory complex and legs for additional characters. However, in no case could species that are difficult to identify be separated by ejaculatory complex or leg character systems. Therefore, although such systems are morphologically interesting, we decline to describe them here.

Species delimitation: An integrative approach to species delimitation was employed, which included a combination of morphological, molecular, and biogeographic characters. The development of species hypotheses was an iterative process. Initial hypotheses were created during the sorting stage with a stereomicroscope by grouping specimens into recognizable units of overall similarity (i.e., morphotypes). These hypotheses were tested with early iterations of phylogenetic analyses, which was the basis for identifying preliminary clades in need of further sampling. For example, distinctive morphotypes recovered as monophyletic, and with low genetic variability across their distribution (i.e., less than one percent difference in COI), were considered as putative species; additional specimens were not added unless found far outside the known geographic range. In contrast, when morphotypes were found to span multiple clades in COI trees, additional specimens from those lineages were included to generate more data in order to test hypotheses for those morphologically cryptic clades.

Measurements (Figure 46): We generally follow Fisher et al. (2015), who modified Goldschmidt (2007), but with the following modifications. Two measurements were added: the distance between Dgl-4 and the height of the rostrum. The following measurements were excluded as they were not found to be important in diagnosing species: distance from genital field to excretory pore; distance from genital field to cauda; dorsal plate length; gnathosomal bend depth; cheliceral height; width of palpomeres, except for the tibia; and all genital skeleton measurements. For each species, when available, a minimum of five individuals of each sex were measured. An effort was made to include members that spanned the range of variability (morphological and molecular) from across the geographic range.

Figure 4. 

Dorsal measurements: A dorsal length B dorsal width C distance between dorsal glandularia, Dgl-4 D anterio-lateral platelet length E anterio-lateral platelet width F anterio-medial platelet length G anterio-medial platelet width. T. multiforma Habeeb, 1974 depicted.

Figure 5. 

Gnathosomal measurements. Palpomeres – A trochanter length B femur length C genu length D tibia length E tibia width F tarsus length. Chelicerae – G cheliceral base length H fang length. Subcapitulum – I ventral length J dorsal length K rostrum length L rostrum width M height. T. multiforma Habeeb, 1974 depicted.

Figure 6. 

Venter measurements: A ventral length B ventral width C anterior venter D coxa-I total length E coxa-I medial length F medial suture length G coxa-III width H gnathosomal bay length I gnathosomal bay width J genital field length K genital field width. T. multiforma Habeeb, 1974 depicted.

To increase time-efficiency, Fisher et al. (2015) suggested digitally measuring compound light micrographs of specimens. However, we abandon that method herein, as many closely-related species of North American Torrenticola are identifiable only with precisely-measured ratios. Measurements were found to be highly inaccurate if taken from images that were not in crisp focus. High-quality, stacked images of each structure can negate this problem (e.g., Hamilton et al. 2016). However, creating such images of slide-prepared mites greatly increases the time invested per specimen and was beyond the scope of this study. Therefore, our measurements were taken with an objective micrometer on a Leica DM 2500 compound microscope.

Molecular phylogenetics: Samples from most sites were sorted to morphotype. Male and female representatives of each morphotype, per sample locality, were chosen for extraction. This resulted in clades containing members from across the geographic range, regardless of our initial speculations on species hypotheses. However, the high abundance of specimens in samples from the far west (California, Oregon, Washington) necessitated that each sample be scanned for morphotypes rather than sorting the entire sample. This was likely sufficient to find most species, but increased the likelihood of missing species present in low abundance. For example, despite obtaining abundant specimens of the Miniforma Group in nearly every sample, we were unable to re-collect T. miniforma Habeeb, 1974, even within the type locality.

To test species hypotheses and guide the description process, the barcoding region of COI was analyzed. We then used the resulting tree as a guide to increase sampling in certain geographic regions, or within certain morphotypes. The result is that many species are represented by specimens from across their distributions and spanning morphological variation. We also analyzed the D2-3 region of 28S rDNA in combination with COI to resolve relationships between species. This analysis included representatives of every species group. All sequences have been deposited in GenBank and accession numbers are located in Suppl. material 1. We follow the recommendation of Chakrabarty et al. (2013) in using GenSeq nomenclature.

Genomic DNA was extracted using Qiagen DNeasy Tissue Kit (Qiagen Inc., Valencia, Calif.). The target regions of COI (450 bp) and 28S (725 bp) were amplified with LCOI and HCOI (Folmer et al. 1994) and D23F and D6R (Park and Ó Foighill 2000), respectively, and purified with Qiagen QIAquick PCR Purification Kits. Test gels (1.5% agarose) confirmed PCR product quality. Purified PCR products were sequenced by Macrogen USA (Rockville, Maryland). Forward and reverse sequences were reconciled with DNASTAR© Lasergene SeqMan (Madison, Wisconsin). Resulting contigs were checked for contamination with BLASTn searches of NCBI’s nr database. Sequences were assessed for the presence of nuclear mitochondrial DNA segments (NUMT) by scanning for in-frame stop codons and indels (Song et al. 2008). COI sequences were aligned with Clustal X (Thompson et al. 1997) and 28S sequences were aligned with MAFFT Version 7 (Katoh and Standley 2013); both alignments were conservatively edited with BioEdit (Hall 1999). Bayesian analyses were performed with MrBayes (3.2.2) using the Extreme Science and Engineering Discovery Environment (XSEDE) infrastructure on the Cipres Portal (Miller et al. 2010), which submits jobs to the Gordon Compute Cluster, a network of 16 supercomputers sponsored by NSF XSEDE at the University of California, San Diego. Each analysis consisted of four simultaneous runs, each with four chains sampling every 1000 generations for 20 million generations, under a GTR+I+Γ model of molecular evolution; 50,000 trees were discarded as burn-in. The resulting majority-rule consensus trees were viewed with Dendroscope 3 (v. 3.5.7) (Huson and Scornavacca 2012); tree image files were then exported in PDF format and edited for final figures in Adobe Illustrator CS6.

Type designation: Earlier works have suggested that males should be used as holotypes due to the importance of characters of the genital skeleton (e.g., Goldschmidt 2007). At least with North American fauna, this is not the case, as females are often more distinctive than males, especially within difficult-to-identify lineages (e.g., Rusetria Complex). Therefore, we have chosen female specimens for most holotypes. Of all species in the region, only nine have male holotypes. Of these nine, four are only known from males (T. anoplopalpa Fisher & Dowling, sp. n.; T. bittikoferae Crowell, 1960; T. dolichodactyla Fisher & Dowling, sp. n.; and T. longitibia Fisher & Dowling, sp. n.) and four were previously described (T. delicatexa Habeeb, 1955; T. neoanomala Habeeb, 1957; T. rufoalba Habeeb, 1955; T. tricolor Habeeb, 1957). Only one species (T. interiorensis Fisher & Dowling, sp. n.) has been deliberately chosen herein to be represented by a male holotype, which we have done to reflect that this species can only be readily-differentiated from its sister species, T. neoanomala, with male characters.

When possible, allotypes (a paratypic member of the opposite sex as the holotype) that best embody the proposed species hypotheses have been selected from the paratypic series. Allotypes are connected to the holotype based upon sampling location, genetic variation, and character consistency. Allotypes are considered to be the best representatives of the paratypic series that are also connected to the holotype with greater confidence than other paratypes.

When possible, the entire type series is represented in our phylogenetic analyses. However, we have found that coloration is valuable for species identification, yet for some lineages a given sex is represented only by specimens preserved in Hoyer’s medium, which destroys coloration. To resolve this problem, we searched GAW-preserved material for representatives. These specimens lack molecular data, but in each instance, specimens were selected that adhere to our species hypothesis with high confidence. For example, specimens were preferred if from the same sample or region, and especially from collection events that also lacked similar species.

Deposition: Material examined for both newly-described and previously-described species (holotypes, allotypes, most paratypes, and most other material examined) were deposited in the Canadian National Collection of Insects, Arachnids, and Nematodes (CNC) in Ottawa, Canada. An exception is T. bittikoferae Crowell, 1960, for which we examined two paratypes deposited in the Ohio State University Acarology Laboratory (OSAL) collection. For all species, when additional specimens were available, representative paratypes and other material examined were also deposited in the Acari Collection of the University of Arkansas (ACUA) in Fayetteville, Arkansas.

Results and discussion

Summary of material: Approximately 2,300 specimens representing 90 species were examined from nearly 500 localities spanning the United States, Canada, and a few collections in Mexico and Central America (Figure 2). Specimens preserved in ethanol (for morphology and molecular data) and GAW (for morphology alone) were available for most locations across the US and Canada, but only GAW samples were available from Mexico and Central America.

Herein, we discuss 90 species of Torrenticola from North America. Previously, 23 species were known from the United States and Canada. Herein we propose the following four synonymies: T. mercedensis (Marshall, 1943) is a junior synonym of T. sierrensis (Marshall, 1943); T. rectiforma Habeeb, 1974 is a junior synonym of T. ellipsoidalis (Marshall, 1943); T. neoconnexa Habeeb, 1957 is a junior synonym of T. magnexa Habeeb, 1955; T. esbelta Cramer, 1992 is a junior synonym of T. boettgeri KO Viets, 1977. Five species previously known only from Mexico and Central America are reported from the southwestern U.S.: T. boettgeri KO Viets, 1977; T. keesdavidsi Cramer, 1992; T. kurtvietsi Cramer, 1992; T. lamellipalpis Viets, 1997; and T. rala Cook, 1980. This raises the number of previously described species known from North America north of Mexico to 24. We also describe 66 new species, which raises the total number of species known from the U.S. and Canada to 90. All previously described species are re-described with color images and updated information.

Summary of phylogenetic analyses: A combined dataset of 28S (725 bp) and COI (450 bp) of 42 species (Suppl. material 1) recovered four species complexes (described below) (Figure 78). These complexes were determined based upon a combination of monophyly and note-worthy characters (e.g., postero-lateral plate fusion in the Rusetria Complex; short, conical rostrum in the Tricolor Complex). The Rusetria and Raptor Complexes were recovered as closely related with strong support. However, we consider relationships among species complexes tentative, pending worldwide sampling. Within these complexes, 12 species groups are noted that are helpful for identification (an additional group, the Rala Group, does not fit within a complex), although many of these groups are not monophyletic.

Figure 7. 

Bayesian inferred cladogram for combined analysis (28S+COI) of North American Torrenticola: species complexes and identification groups. Four well-supported complexes are organized into 13 groups that are helpful for identification. Note that 1) not all identification groups are monophyletic (in quotation marks); 2) not all species are placed in a complex; and 3) some species are not placed within a group, reflecting their identification difficulty. Numbers are DNA identification numbers. Dots denote posterior probably >95%.

COI sequence data were obtained for 481 individuals from across the United States and Canada (Suppl. material 1), including all specimens from the combined analysis. These 481 individuals group into 57 clades that represent well-supported species hypotheses (Figures 918). Of these 57 species hypotheses, 13 clades are identifiable to previously-described species: T. delicatexa Habeeb, 1955; T. ellipsoidalis (Marshall, 1943); T. larvata Cherri, Fisher, & Dowling, 2016; T. multiforma Habeeb, 1974; T. neoanomala Habeeb, 1957; T. magnexa Habeeb, 1955; T. nigroalba Habeeb, 1955; T. projector Habeeb, 1961; T. rala Cook, 1980; T. sierrensis (Marshall, 1943); T. tahoei (Marshall, 1943), T. tricolor Habeeb, 1957, T. trimaculata Fisher, 2015; T. ventura Habeeb, 1973. Two clades include multiple previously-named species, necessitating the synonymization of three species. Ethanol-preserved specimens of seven previously described species, known from the United States and Canada (T. bittikoferae Crowell, 1960; T. indistincta (Marshall, 1929); T. kittatinniana Habeeb, 1955; T. miniforma Habeeb, 1974; T. occidentalis (Marshall, 1933); and T. rufoalba), were not available, but the type series for each of these were examined morphologically. Of the 57 species in our molecular analysis, 44 represent new species.

Species Complexes and Groups (Table 1): As considered herein, species complexes are monophyletic collections of closely-related species recovered by our combined (COI+28S) phylogenetic analyses (Figure 78). In addition, most species known only from morphology are placed within the tree in suspected complexes. Species complexes proposed herein and their relationships to each other should be considered a first step in need of future exploration that will greatly benefit from worldwide sampling. Our analyses recover four species complexes: Miniforma; Raptor; Rusetria; and Tricolor.

Distribution of the 90 species of Torrenticola from the US and Canada among Species Complexes and Groups. Number of species within a group are denoted parenthetically. Species for which molecular data was available are denoted with an asterisk. Species known only from one sex are denoted with sex symbols (♀ or ♂). Species Groups in quotation marks were not recovered as monophyletic in our molecular analyses.

Tricolor Complex (14 sp.) 1. “Tricolor Group” (13 sp.) a. T. bittikoferae ♂ b. T. cardia c. T. dimorpha* d. T. hoosieri* e. T. kringi f. T. larvata* g. T. mohawk h. T. olliei* i. T. pearsoni* j. T. sierrensis* k. T. tricolor* l. T. trimaculata* m. T. unimaculata* 2. Unplaced (1 sp.) a. T. projector* Miniforma Complex (13 sp.) 1. Ellipsoidalis Group (4 sp.) a. T. ellipsoidalis* b. T. leviathan c. T. multiforma* d. T. occidentalis ♀ 2. Miniforma Group (7 sp.) a. T. copipalpa* b. T. manni* c. T. miniforma d. T. oliveri e. T. pacificensis* f. T. pinocchio g. T. rockyensis* 3. Unplaced (2 sp.) a. T. regalis* b. T. tahoei* Raptor Complex (24 sp.) 1. Elongata Group (3 sp.) a. T. elongata* b. T. gorti* c. T. reduncarostra 2. Erectirostra Group (3 sp.) a. T. erectirostra* b. T. karambita* c. T. robisoni* 3. Neoanomala Group (2 sp.) a. T. interiorensis* b. T. neoanomala* 4. Nigroalba Group (4 sp.) a. T. dentirostra b. T. flangipalpa* c. T. nigroalba* d. T. solisorta* 5. “Raptor Group” (10 sp.) a. T. daemon b. T. danielleae c. T. elusiva* ♀ d. T. gnoma* e. T. irapalpa* f. T. ivyae g. T. longitibia* ♂ h. T. mjolniri* i. T. racupalpa* j. T. raptor* 6. Unplaced (2 sp.) a. T. bondi* b. T. skvarlai* Rusetria Complex (26 sp.) 1. “Eastern 2-plates” (12 sp.) a. T. bicutella* b. T. caerulea* c. T. delicatexa* d. T. feminellai e. T. indistincta f. T. malarkeyorum* g. T. microbiscutella* h. T. pendula* i. T. sellersorum* j. T. tysoni* k. T. ululata* l. T. whitneyae 2. “Eastern 4-plates” (6 sp.) a. T. dunni* b. T. glomerabilis* c. T. kittatinniana d. T. pollani* e. T. rufoalba f. T. shubini* 3. Partial 2-plates (4 sp.) a. T. folkertsae b. T. magnexa* c. T. priapus d. T. pulchra 4. Western 2-plates (4 sp.) a. T. mulleni* b. T. nortoni* c. T. walteri* d. T. welbourni* ♀
Unplaced (13 sp.) 1. Rala Group (7 sp.) a. T. anoplopalpa ♂ b. T. boettgeri c. T. dolichodactyla ♂ d. T. keesdavidsi e. T. kurtvietsi f. T. lamellipalpis b. T. rala* 2. Unplaced (6 sp.) a. T. arktonyx b. T. raptoroides* c. T. sharkeyi* d. T. ventura* e. T. wiedenmanni f. T. oregonensis

Four western species included in the molecular analysis do not fit into a species complex: T. ventura Habeeb, 1973; T. raptoroides Fisher & Dowling, sp. n.; T. sharkeyi Fisher & Dowling, sp. n.; and T. rala Cook, 1980. Additionally, fresh material for molecular analysis was not available for six other species that are suspected to be closely related to T. rala and therefore included in the Rala Group (below): T. boettgeri KO Viets, 1977; T. kurtvietsi Cramer, 1992; T. lamellipalpis KO Viets, 1977; T. keesdavidsi Cramer, 1992; T. dolichodactyla Fisher & Dowling, sp. n.; and T. anoplopalpa Fisher & Dowling, sp. n. Collectively these species (T. ventura, T. raptoroides, T. sharkeyi and the Rala Group), resemble fauna from south of the US. The phylogenetic affinity of these species can only be determined with the addition of worldwide fauna, especially from Mexico and Central America.

As considered herein, “species groups” represent closely related species that are readily identifiable to the species group level, often even under low magnification (i.e., stereoscope). The function of these species groups is to aid identification, as often species of a group are easier to recognize at the group-level than the species-level. Occasionally, identifying a given specimen to species-level requires merely combining group-level identification with locality. Importantly, four species groups are not monophyletic: Tricolor Group, Raptor Group, Eastern 4-Plates, and Eastern 2-Plates. This is because these species groups serve to ease identification, not to inform relationship. However, despite the utility of learning these species groups, they are meant to merely augment the key. Accurate identification of most species still requires keying slide preparations under higher magnification (i.e., compound microscope) with precise measurements.

Thirteen species do not fit into species groups. Three of these species are among the most recognizable species in the genus, but are single-species and thus not “groups”: T. arktonyx Fisher & Dowling, sp. n.; T. projector Habeeb, 1961; and T. tahoei (Marshall, 1943). Nine of the 13 species are not placed within a species group because they are difficult to identify under lower magnification and have little in common with sister species. The reader is referred to the key and diagnoses for these species: T. oregonensis Fisher & Dowling, sp. n.; T. wiedenmanni Fisher & Dowling, sp. n.; T. bondi Fisher & Dowling, sp. n.; T. occidentalis (Marshall, 1933); T. raptoroides Fisher & Dowling, sp. n.; T. regalis Fisher & Dowling, sp. n.; T. sharkeyi Fisher & Dowling, sp. n.; T. skvarlai Fisher & Dowling, sp. n.; and T. ventura Habeeb, 1973.

The Rala Group (Figure 79, 11) is the only species group that does not fit within a species complex. This group comprises seven species: T. boettgeri; T. kurtvietsi; T. lamellipalpis; T. keesdavidsi; T. rala; T. dolichodactyla; and T. anoplopalpa. These species are colorless and have incomplete or indistinct hind coxal margins. They are southwestern (Arizona, New Mexico, and Texas), with a few extending into Mexico and Guatemala. However, we suspect all seven species extend well into Mexico and perhaps further south, with US populations representing the northern-most extents of their distributions.

Figure 8. 

Bayesian inferred phylogram for combined analysis (28S+COI) of North American Torrenticola: distributions of 57 species. Pink represents lineages distributed in eastern North America and blue represents lineages distributed in western North America (gray branches represent lineages distributed in both regions). Species marked with an asterisk are primarily eastern but have distributions extending into the west. Note that larger lineages contain species distributed in either the east or west, but not both. Numbers are DNA identification numbers. Scale bar indicates 0.1 substitutions per site. Dots denote posterior probably >95%.

Figure 9. 

Bayesian inferred circular phylograms of North American Torrenticola, COI only. Scale bars indicate 1.0 substitutions per site. A Overall tree depicting the four complexes recovered in combined analysis. Dots denote posterior probability >95% B Species distributions; branch colors correspond to regions of North America depicted on the right (eastern, pink; western teal). Asterisks denote eastern species that extend into the west. Note that lineages tend to contain species distributed in either the east or west, but not both. Detail of this figure is presented in Figures 1018.

Raptor Complex (Figure 711): Molecular analyses recovered 19 species in this complex. One of these species, T. racupalpa Fisher & Dowling, sp. n., could not be included in the combined dataset, but was included in the COI analysis, where it was recovered as sister to T. elusiva Fisher & Dowling, sp. n. Additionally, fresh material for molecular analysis was not available for five other species suspected as being within this complex based upon morphology: T. danielleae Fisher & Dowling, sp. n.; T. daemon Fisher & Dowling, sp. n.; T. ivyae Fisher & Dowling, sp. n.; T. reduncarostra Fisher & Dowling, sp. n.; and T. dentirostra Fisher & Dowling, sp. n. In total, there are 24 species within the Raptor Complex.

Figure 10. 

Raptor Complex (part I). These species are eastern, although T. irapalpa extends further west. Note that two species (T. skvarlai and T. bondi) do not fit into identification groups. A Species guide (overview of 28S+COI analysis from Figure 78) B Overview of Raptor Complex from Figure 9 (COI-only analysis from Figure 9) C Detail of Raptor Complex (in part) from B. Colored lineages in A and B correspond to Group names and brackets in C. Dots denote posterior probably of greater than 95%. Taxa are displayed by DNA number and state or territory abbreviation.

Figure 11. 

Raptor Complex (part II). Species of the Raptor Complex are eastern, but the species that do not fit into the complex (T. rala; T. sharkeyi; T. raptoroides; & T. skvarlai) are western. A Species guide (overview of 28S+COI analysis from Figure 78) B Overview of Raptor Complex from Figure 9 (COI-only analysis from Figure 9) C Detail of Raptor Complex (in part) from B. Colored lineages in A and B correspond to Group names and brackets in C. Dots denote posterior probably of greater than 95%. Taxa are displayed by DNA number and state or territory abbreviation.

All members of the Raptor Complex are eastern. A collection of western species (T. raptoroides Fisher & Dowling, sp. n.; T. sharkeyi Fisher & Dowling, sp. n.; T. ventura Habeeb, 1973; and the Rala Group) that do not form a monophyletic group, are often recovered as sister to the Raptor Complex. This suggests that the Raptor Complex may have originated from a Neotropical ancestor that dispersed into the eastern US and subsequently diversified. However, this speculation requires worldwide sampling for support.

Basal lineages within the Raptor Complex remain unsupported/unresolved, but a few relationships within the complex were recovered with high support. First, the Erectirostra Group was not recovered as monophyletic in the COI analysis, but is well-supported in the combined analysis. Second, the Neoanomala and Erectirostra Groups are closest relatives; indeed, apart from the apomorphic subcapitulum of the Erectirostra Group, the two groups are otherwise similar. Third, a sister relationship between the Erectirostra + Neoanomala lineage and the Nigroalba Group was recovered with high support. Finally, both analyses support a sister relationship between T. bondi Fisher & Dowling, sp. n. and the Elongata Group.

22 of the 24 species of the Raptor Complex are organized into five species groups: Raptor Group, Elongata Group, Nigroalba Group, Erectirostra Group, and Neoanomala Group. Two species do not fit within an identification group: T. bondi Fisher & Dowling, sp. n. and T. skvarlai Fisher & Dowling, sp. n.

The Raptor Group (Figure 10) comprises ten eastern species: T. danielleae Fisher & Dowling, sp. n.; T. daemon Fisher & Dowling, sp. n.; T. ivyae Fisher & Dowling, sp. n.; T. gnoma Fisher & Dowling, sp. n.; T. irapalpa Fisher & Dowling, sp. n.; T. longitibia Fisher & Dowling, sp. n.; T. mjolniri Fisher & Dowling, sp. n.; T. elusiva Fisher & Dowling, sp. n.; T. racupalpa Fisher & Dowling, sp. n.; and T. raptor Fisher & Dowling, sp. n. These closely related species are readily identifiable with the following combination of characters: round bodies; Dgl-4 close to muscles scars; long thin subcapitular rostra; and long, thin pedipalp tibiae. This group was not recovered as monophyletic in our analyses; however, relationships within this group are also not well-supported, so neither monophyly nor non-monophyly can be rejected.

The Elongata Group (Figure 10) comprises three eastern species: T. elongata Fisher & Dowling, sp. n.; T. reduncarostra Fisher & Dowling, sp. n.; and T. gorti Fisher & Dowling, sp. n. Members of this distinctive group are readily identifiable by the combination of characters: small body size; bold, distinctive coloration; and elongate, ovoid body shape. No other species share this combination of characters. Furthermore, no other Torrenticola have similar coloration to the dark morph of T. gorti and no other Torrenticola are as elongate as T. elongata.

The Nigroalba Group (Figure 11) comprises four eastern species: T. flangipalpa Fisher & Dowling, sp. n.; T. dentirostra Fisher & Dowling, sp. n.; T. nigroalba Habeeb, 1955; and T. solisorta Fisher & Dowling, sp. n. These closely related species are readily identifiable with the following combination of characters: small body size; long, thin subcapitular rostra and pedipalp tibiae; dorsal purple coloration restricted to posterior half; somewhat elongate bodies; and hind coxal margins that are distinctly present, but the margins of are poorly defined. Several western species can superficially resemble members of the Nigroalba Group under low magnification (e.g., T. tahoei and T. regalis).

The Erectirostra Group (Figure 11) comprises three eastern species: T. erectirostra Fisher & Dowling, sp. n.; T. karambita Fisher & Dowling, sp. n.; and T. robisoni Fisher & Dowling, sp. n. These closely related species are readily identifiable with the following combination of characters: thick subcapitular rostra (when viewed ventrally) that is strongly upturned and dentate and Cxgl-4 nearly halfway down gnathosomal bay. No other Torrenticola in North America have similar subcapitular rostra and members of the Erectirostra Group can be readily sorted under low magnification. In terms of body shape and coloration, they superficially resemble Neoanomala Group, to which they are closely related.

The Neoanomala Group (Figure 11) comprises two eastern species: T. interiorensis Fisher & Dowling, sp. n. and T. neoanomala Habeeb, 1957. These closely related species are readily identifiable with the following combination of characters: dorsal coloration often purplish, separated into anterior and posterior portions; hind coxal apodemes distinct; and other characters relatively unmodified. Members of the Neoanomala Group are readily differentiated from the superficially similar Rusetria Group by having distinct hind coxal apodemes, whereas all members of the Rusetria Group have indistinct hind coxal apodemes. Another species—T. bondi Fisher & Dowling, sp. n.—resembles members of this group, but is not closely related. T. bondi have a shorter medial suture (10–15 µm in female T. bondi, 22–40 µm in female Neoanomala Group; 55–70 µm in male T. bondi, 75–110 µm in male Neoanomala Group). Also, T. bondi is currently known only from Haywood County (North Carolina), which implies that this species is either rare, or has a restricted distribution.

Miniforma Complex (Figure 79, 1213): Molecular analyses recovered eight western species in this complex. Additionally, fresh material for molecular analysis was not available for five other species (T. occidentalis (Marshall, 1933); T. miniforma Habeeb, 1974; T. oliveri Fisher & Dowling, sp. n.; T. leviathan Fisher & Dowling, sp. n.; and T. pinocchio Fisher & Dowling, sp. n.) that are suspected to be within this complex based upon morphology. Another species, T. oregonensis Fisher & Dowling, sp. n., superficially resembles members of this complex, but is divergent enough that we avoid placing this species pending molecular support. Therefore, in total, we propose 13 species in this complex.

Figure 12. 

Miniforma Complex (part I). All species are western. Note that the phylogenetic structure in the Miniforma Group corresponds to biographic regions (Rockies, Pacific Ranges, southwest). A Species guide (overview of 28S+COI analysis from Figure 78) B Overview of Miniforma Complex from Figure 9 (COI-only analysis from Figure 9) C Detail of Miniforma Complex (in part) from B C Detail of Miniforma Complex (in part) from Figure 9. Colored lineages in A and B correspond to Group names and brackets in C. Dots denote posterior probably of greater than 95%. Taxa are displayed by DNA number and state (U.S.) or territory (Canada) abbreviation.

Figure 13. 

Miniforma Complex (part II). All species are western. One species (T. regalis) does not fit within a species group. Note that T. ellipsoidalis and T. multiforma are widespread in the west (C). A Species guide (overview of 28S+COI analysis from Figure 78) B Overview of Miniforma Complex from Figure 9 (COI-only analysis from Figure 9) C Detail of Miniforma Complex (in part) from B C Detail of Miniforma Complex (in part) from Figure 9. Colored lineages in A and B correspond to Group names and brackets in C. Dots denote posterior probably of greater than 95%. Taxa are displayed by DNA number and state (U.S.) or territory (Canada) abbreviation.

These species are organized into two monophyletic species groups: Ellipsoidalis Group and Miniforma Group. The two members of this complex that are not placed within a species group include T. tahoei (Marshall, 1943), which is among the most recognizable species of the genus, and T. regalis Fisher & Dowling, sp. n., which can only be confidently identified by keying slide-mounted specimens. Torrenticola tahoei is consistently recovered as sister to the rest of the species in this complex and T. regalis is consistently recovered sister to Ellipsoidalis Group.

The Miniforma Group (Figure 12) comprises seven western species: T. manni Fisher & Dowling, sp. n.; T. oliveri Fisher & Dowling, sp. n.; T. pinocchio Fisher & Dowling, sp. n.; T. miniforma Habeeb, 1974; T. pacificensis Fisher & Dowling, sp. n.; T. rockyensis Fisher & Dowling, sp. n.; and T. copipalpa Fisher & Dowling, sp. n. Members of this group are often readily identified by their smaller body size than other Torrenticola within their range except for Wester 2-Plates (T. mulleni Fisher & Dowling, sp. n.; T. nortoni Fisher & Dowling, sp. n.; T. walteri Fisher & Dowling, sp. n.; and T. welbourni Fisher & Dowling, sp. n.). Members of the Miniforma Group are easily distinguished from Western 2-Plates, which have dorso-lateral platelets fused to the dorsal plate. The Miniforma Group can be further differentiated from all other Torrenticola by their distinctive pedipalp genual extensions. Females of this group have a relatively large genital field, which we suspect coincides with loss of larval parasitism.

The Ellipsoidalis Group (Figure 13) comprises four western species: T. occidentalis (Marshall, 1933); T. ellipsoidalis (Marshall, 1943); T. leviathan Fisher & Dowling, sp. n.; and T. multiforma Habeeb, 1974. Members of this group are readily identified from co-occurring Torrenticola via their larger body size than other species in a given sample. Torrenticola occidentalis may fit within this species group based upon overall similarity and by having a short, conical rostrum, but we were not able to obtain fresh material to confirm this hypothesis.

Rusetria Complex (Figure 79, 1416): Molecular analyses recovered 17 species in this complex. Additionally, fresh material for molecular analysis was not available for nine other species that are included here based upon morphology: T. rufoalba Habeeb, 1955; T. kittatinniana Habeeb, 1955; T. indistincta (Marshall, 1929); T. whitneyae Fisher & Dowling, sp. n.; T. microbiscutella Fisher & Dowling, sp. n.; T. feminellai Fisher & Dowling, sp. n.; T. priapus Fisher & Dowling, sp. n.; T. folkertsae Fisher & Dowling, sp. n.; and T. pulchra Fisher & Dowling, sp. n. In total, there are 26 species in this complex.

Figure 14. 

Rusetria Complex (part I). All species are eastern, but note the distribution of T. sellersorum extends into New Mexico and Saskatchewan. A Species guide (overview of 28S+COI analysis from Figure 78) B Overview of Rusetria Complex from Figure 9 (COI-only analysis from Figure 9) C Detail of Rusetria Complex (in part) from B C Detail of Rusetria Complex (in part) from Figure 9. Colored lineages in A and B correspond to Group names and brackets in C. Dots denote posterior probably of greater than 95%. Taxa are displayed by DNA number and state (U.S.) or territory (Canada) abbreviation.

Figure 15. 

Rusetria Complex (part II). All species are eastern. Relationships among species groups remain unresolved, except for three species (T. dunni; T. pollani; and T. shubini), which form a well-supported clade in all analyses. A Species guide (overview of 28S+COI analysis from Figure 78) B Overview of Rusetria Complex from Figure 9 (COI-only analysis from Figure 9) C Detail of Rusetria Complex (in part) from B C Detail of Rusetria Complex (in part) from Figure 9. Colored lineages in A and B correspond to Group names and brackets in C. Dots denote posterior probably of greater than 95%. Taxa are displayed by DNA number and state (U.S.) or territory (Canada) abbreviation.

Figure 16. 

Rusetria Complex (part III). All species are western. A Species guide (overview of 28S+COI analysis from Figure 78) B Overview of Rusetria Complex from Figure 9 (COI-only analysis from Figure 9) C Detail of Rusetria Complex (in part) from B. D Detail of Rusetria Complex (in part) from Figure 9. Colored lineages in A and B correspond to Group names and brackets in C. Dots denote posterior probably of greater than 95%. Taxa are displayed by DNA number and state (U.S.) or territory (Canada) abbreviation.

Unlike other species complexes, the Rusetria Complex corresponds to a previously recognized subgenus—“Rusetria”—that occurs worldwide and is identified by the fusion of the lateral platelets. Goldschmidt (2007) called species with fused lateral platelets “Rusetria-like species” and we continue that terminology here. However, our analyses indicate that a reversal to free lateral platelets occurred twice: once in a clade sister to the Eastern 2-Plates and once within the Eastern 2-Plates (T. glomerabilis). Members of the Rusetria Complex can also be differentiated from superficially similar species by having indistinct hind coxae (distinct in most similar species, such as the Neoanomala and Raptor Groups). Only a few other species have indistinct hind coxal margins (T. dolichodactyla, T. skvarlai, and occasionally T. flangipalpa) or partial hind coxal margins (T. regalis, T. sharkeyi, and membes of the Nigroalba and Rala Groups,). This character is particularly useful when differentiating Eastern 4-Plates from similar species.

Two interesting evolutionary stories are suggested by the combined molecular analysis. The first involves the fusion of the lateral platelets, which seems to have a complex evolutionary history. For example, if this character is synapomorphic for the Rusetria Complex, then there have been multiple reversals. Furthermore, the position of the Partial 2-Plates suggests that the partial fusion of their lateral platelets represents a transitional step to complete fusion. But this story is even more complicated by T. glomerabilis, which has regained free lateral platelets separately from the rest of the Eastern 4-Plates. Ultimately, understanding the evolution of lateral platelet fusion, even within the eastern U.S., will depend on a much greater sampling of this worldwide species complex. Second, significant sexual size dimorphism (i.e., males 20–30% smaller than females), which is found in most members of this complex worldwide, likely arose once within the complex basally, but was subsequently and independently lost (i.e., males 5–15% smaller than females) three times: twice within the Eastern 4-Plates (once with the sister species T. dunni Fisher & Dowling, sp. n. and T. pollani Fisher & Dowling, sp. n., and once with T. glomerabilis Fisher & Dowling, sp. n.) and once with T. ululata Fisher & Dowling, sp. n.

The 23 species of the Rusetria Complex are organized into four species groups: Eastern 4-Plates, Eastern 2-Plates, Partial 2-Plates, and Western 2-Plates.

The Eastern 2-Plate Group (Figure 1415) comprises 12 eastern species: T. biscutella Fisher & Dowling, sp. n.; T. whitneyae Fisher & Dowling, sp. n.; T. microbiscutella Fisher & Dowling, sp. n.; T. feminellai Fisher & Dowling, sp. n.; T. caerulea Fisher & Dowling, sp. n.; T. delicatexa Habeeb, 1955; T. indistincta (Marshall, 1929); T. malarkeyorum Fisher & Dowling, sp. n.; T. pendula Fisher & Dowling, sp. n.; T. sellersorum Fisher & Dowling, sp. n.; T. tysoni Fisher & Dowling, sp. n.; and T. ululata Fisher & Dowling, sp. n. This group is not monophyletic because of the exclusion of T. glomerabilis Fisher & Dowling, sp. n., which is considered an Eastern 4-Plate based on morphology. Members of this group are readily identifiable by having dorso-lateral platelets fused with the dorsal plate, thus giving the appearance of having only two plates, the anterio-medial platelets. Most members of this group are colorful (except for T. indistincta, which resembles Western 2-Plates) and nearly all have similar dorsal patterns that are separated into anterior and posterior portions, which are occasionally connected medially (except T. ululata, Figures 262263). Members of the Eastern 2-Plate group are similar to the western counterpart (below) in having fused lateral platelets. However, the majority of eastern species are distinctly more colorful than most western species, which are either colorless or only faintly colored, and the groups exhibit non-overlapping ranges.

The Western 2-Plate Group (Figure 16) comprises four western species: T. mulleni Fisher & Dowling, sp. n.; T. nortoni Fisher & Dowling, sp. n.; T. walteri Fisher & Dowling, sp. n.; and T. welbourni Fisher & Dowling, sp. n. Members of this group are readily identifiable by having dorso-lateral platelets fused with the dorsal plate, thus giving the appearance of having only two plates. Although resembling Eastern 2-Plates, ranges are non-overlapping and western species are immediately identifiable by being colorless or nearly so, whereas most eastern species (except T. indistincta) are colorful. Besides range information, Torrenticola indistincta can be differentiated from Western 2-Plates by having coxal apodemes I-II not meeting posteriorly (meeting in western-two plates, usually with an accompanying medial suture).

The Partial 2-Plate Group (Figure 15) comprises four eastern species: T. priapus Fisher & Dowling, sp. n.; T. folkertsae Fisher & Dowling, sp. n.; T. pulchra Fisher & Dowling, sp. n.; and T. magnexa Habeeb, 1955. These species are readily distinguished from all other Torrenticola by having dorso-lateral platelets only partially fused to the dorsal plate. In practice, when sorting under low magnification (i.e., stereoscope), this group appears to be Eastern 2-Plates, but closer inspection will show that the lateral platelet borders are distinct, even under low magnification. This group will likely not be confused with members of the Rusetria 4-Plates (below), as the latter do not appear to have lateral plate fusion, even under low magnification.

The Eastern 4-Plate Group (Figure 15) comprises six eastern species: T. dunni Fisher & Dowling, sp. n.; T. glomerabilis Fisher & Dowling, sp. n.; T. kittatinniana Habeeb,1955; T. pollani Fisher & Dowling, sp. n.; T. rufoalba Habeeb, 1955; and T. shubini Fisher & Dowling, sp. n. This group is not monophyletic because T. glomerabilis is more closely related to Eastern 2-Plates. Members of this group can be readily differentiated from all other species of the Rusetria Complex by having anterio-lateral platelets completely free from the dorsal plate, hence “4-Plates”. All members of this group are colorful and have similar dorsal patterns that are separated into anterior and posterior portions and occasionally connected medially. Members of this group resemble the Neoanomala Group in overall appearance and coloration; however, they can be readily differentiated by having indistinct hind coxal margins (distinct in Neoanomala Group). Members of this group also resemble T. skvarlai in terms of overall appearance and coloration, and in having indistinct hind coxal margins; however, they can be readily differentiated by having conical, tuberculate pedipalp femoral extensions (broadly tuberculate in T. skvarlai).

Tricolor Complex (Figure 79, 1718): Molecular analyses recovered 10 species in this complex. Fresh material for molecular analysis was not available for four other species (T. bittikoferae Crowell, 1960; T. cardia Fisher & Dowling, sp. n.; T. kringi Fisher & Dowling, sp. n.; and T. mohawk Fisher & Dowling, sp. n.) that are grouped within this complex based upon morphology. In total, there are 14 species within the Tricolor Complex.

Figure 17. 

Tricolor Complex (part I). All species are eastern. These are some of the most recognizable of all Torrenticola, namely the elongate T. projector and a clade of species with diagnostic color patterns (T. trimaculata, T. unimaculata, and T. tricolor). Note that three species (T. pearsoni, T. hoosieri, and T. projector) are repeated in Figure 18. A Species guide (overview of 28S+COI analysis from Figure 78) B Overview of Tricolor Complex from Figure 9 (COI-only analysis from Figure 9) C Detail of Tricolor Complex (in part) from B C Detail of Tricolor Complex (in part) from Figure 9. Colored lineages in A and B correspond to Group names and brackets in C. Dots denote posterior probably of greater than 95%. Taxa are displayed by DNA number and state (U.S.) or territory (Canada) abbreviation.

Figure 18. 

Tricolor Complex (part II). Most species of the Tricolor Complex are eastern, but one lineage is western: T. sierrensis, widespread; and T. olliei, north Pacific coast. The most apomorphic Torrrenticola (T. dimorpha) is recovered here in all analyses. Note that three species (T. pearsoni, T. hoosieri, and T. projector) are repeated in Figure 17. A Species guide (overview of 28S+COI analysis from Figure 78) B Overview of Tricolor Complex from Figure 9 (COI-only analysis from Figure 9) C Detail of Tricolor Complex (in part) from B C Detail of Tricolor Complex (in part) from Figure 9. Colored lineages in A and B correspond to Group names and brackets in C. Dots denote posterior probably of greater than 95%. Taxa are displayed by DNA number and state (U.S.) or territory (Canada) abbreviation.

This complex is divided into eastern and western lineages. The eastern lineage contains species that are among the most distinctive of all Torrenticola (e.g., T. trimaculata Fisher, 2015; T. dimorpha Fisher & Dowling, sp. n.; and T. projector Habeeb, 1961). This complex contains one species group, the Tricolor Group, which is paraphyletic with respect to T. projector. This species has an elongate morphology that is unique among species of the Tricolor Complex.

The Tricolor Group (Figure 1718) comprises 13 species: T. bittikoferae Crowell, 1960; T. cardia Fisher & Dowling, sp. n.; T. kringi Fisher & Dowling, sp. n.; T. dimorpha Fisher & Dowling, sp. n.; T. mohawk Fisher & Dowling, sp. n.; T. hoosieri Fisher & Dowling, sp. n.; T. larvata Cherri, Fisher, & Dowling, 2016; T. olliei Fisher & Dowling, sp. n.; T. pearsoni Fisher & Dowling, sp. n.; T. projector Habeeb, 1961; T. sierrensis (Marshall, 1943); T. tricolor Habeeb, 1957; T. trimaculata Fisher, 2015; and T. unimaculata Fisher & Dowling, sp. n. Most species are distributed in eastern North America, except for T. sierrensis and T. olliei, which are western. The most distinguishing characteristic of this group is a short, conical rostrum, which is noticeably downturned in males. The only exception is T. kringi, which has a conical rostrum that is longer than other members of the group. Five members of the Tricolor Group form a monophyletic subgroup and can be readily recognized by having dorsal coloration composed of one or more spots (T. kringi, T. mohawk, T. tricolor, T. trimaculata, T. unimaculata). The Tricolor Group also contains a morphologically enigmatic species that has strong sexual dimorphism: T. dimorpha. Finally, T. bittikoferae likely fits within this complex, but fresh material was not available to test this hypothesis.

Biogeography

Of the 90 species discussed herein, 57 are primarily eastern (east of the 100th Meridian) and 33 are western (west of the 100th Meridian). Only a few species are distributed on either side of this barrier: T. irapalpa is widespread in the east and has populations in Saskatchewan; T. rala is widespread from Costa Rica to Arizona and is also found in south-central Texas; and T. sellersorum has a widespread, but sporadic, distribution spanning Arizona to Manitoba and Ohio to Pennsylvania. Aside from these few species, the Great Plains acts as a biogeographic barrier for most species and Torrenticola are not common within this region.

Much of the discrepancy between the eastern and western diversity is due to increased diversity in the Appalachians, particularly due to increased speciation within the Raptor and Rusetria Complexes, which have 17 (14 endemic) and 15 (12 endemic) species found in the Appalachians, respectively. The Southern Appalachian region is particularly diverse, with 14 endemics (T. arktonyx, T. bondi, T. karambita, T. longitibia, T. danielleae, T. daemon, T. dentirostra, T. racupalpa, T. dunni, T. glomerabilis, T. whitneyae, T. microbiscutella, T. feminellai, and T. pollani). The Northern Appalachian region contains six species known only from that area (T. elusiva, T. kittatinniana, T. folkertsae, T. pendula, T. rufoalba, and T. mohawk); however, two of these are known only from their type localities in northern New Jersey (T. kittatinniana and T. rufoalba). Eight species are restricted to the Appalachians, but range throughout both northern and southern regions (T. erectirostra, T. gorti, T. reduncarostra, T. skvarlai, T. delicatexa, T. shubini, T. cardia, T. projector). Three species appear to be widespread in the northeast (i.e., extending into the Great Lakes region in Ottawa), but also extend southward throughout the Appalachians (T. nigroalba, T. raptor, and T. tricolor). The remaining eastern diversity includes seven widespread species (T. malarkeyorum, T. trimaculata, T. irapalpa, T. neoanomala, T. magnexa, T. priapus, and T. sellersorum); seven species endemic to the Interior Highlands (T. biscutella, T. interiorensis, T. robisoni, T. solisorta, T. larvata, T. pearsoni, and T. unimaculata); six species from the non-mountainous southeast (T. caerulea, T. elongata, T. flangipalpa, T. gnoma, T. tysoni, and T. ululata); four species known only from the Midwest (T. bittikoferae, T. indistincta, T. hoosieri, and T. pulchra); one species from Florida (T. ivyae); and one species from eastern Texas (T. kringi).

Our results point to two eastern regions that have high diversity and an increased proportion of endemic species: the southern Appalachians (including southern Pennsylvania) and the Interior Highlands (Ozarks and Ouachitas of Missouri, Arkansas, and Oklahoma). Of the 31 species known from the southern Appalachians, 14 are endemic. Of the 13 species known from the Interior Highlands, seven are endemic. Each of these regions is well-known for its diversity and endemism of aquatic taxa, due to a complex biogeographic history where they have acted as refugia (Robison & Allen 1995; Radwell et al. 2011; Skvarla et al. 2015).

By contrast, western Torrenticola are less diverse, with 32 species known from the region. Four species are widespread (T. ellipsoidalis, T. multiforma, T. sierrensis, and T. tahoei), although only one of these, T. multiforma, is also found in the southwest. Two species are found throughout much of California (T. nortoni and T. ventura) with only T. ventura extending northward into southwestern Oregon. One species is known from south-central Texas (T. dimorpha). The remaining western diversity is distributed into patterns that roughly correspond to three major ecoregions: Rocky Mountains, Pacific Ranges, and the arid southwest. The Rockies contain three endemics (T. occidentalis, T. rockyensis, and T. mulleni). The Pacific Ranges of British Columbia, Washington, Oregon, and northern California contain nine endemics (T. miniforma, T. oregonensis, T. oliveri, T. leviathan, T. pinocchio, T. pacificensis, T. regalis, T. welbourni, and T. olliei), and two species that also range into the Sierra Nevada (T. copipalpa and T. walteri), although the latter (T. walteri) is also known from one locality in the Rockies of southern British Columbia. The arid southwest (southern California, Arizona, and New Mexico) contains eleven species restricted to that region in the US (T. boettgeri, T. keesdavidsi, T. kurtvietsi, T. lamellipalpis, T. dolichodactyla, T. anoplopalpa, T. raptoroides, T. sharkeyi, T. wiedenmanni, and T. rala), although five of these extend southward into Mexico and Central America (T. boettgeri, T. keesdavidsi, T. kurtvietsi, T. lamellipalpis, and T. rala) and one is also found in southern Texas (T. rala).

Our results suggest the diversity of Torrenticola in eastern North America can be explained by as few as three dispersal events, followed by subsequent radiations (Figure 8). Western diversity is more complex and was likely influenced from southern species extending northward and also by northern species that crossed Beringia and extended southward. However, even these results are speculative, pending worldwide sampling.

The distributions described above show the variable dispersal capabilities of different species within the genus. For instance, some species are wide-ranging, with distributions that span multiple topographic barriers (e.g., T. irapalpa, T. multiforma, and T. ellipsoidalis), whereas other groups are endemic to specific geographic areas and so are not able to cross such barriers (e.g., T. pacificensis, T. rockyensis, T. arktonyx, and T. solisorta). To explain this variation, adequate knowledge of the dispersal capabilities of both the adult and larval stages is required.

It seems well-understood that water mite larvae utilize the dispersal capabilities of their winged insect hosts. Indeed, it is normally inferred that dispersal is the primary function of the parasitic larvae, as is exemplified by the following passage from Proctor et al. (2015, pg. 639):

“The dominant strategic role of the larval instar in the life history of most Hydryphantoidea, Lebertioidea, Hygrobatoidea, and Arrenuroidea appears to be dispersal rather than growth. Species that have the parasitic larval stage suppressed illustrate that development of large eggs can obviate the need for larval feeding. However, the relative rarity of this phenomenon attests to the crucial role of larval dispersal in more derived water mite species.”

Consistent with this view, many lentic water mites utilize far-flying hosts such as dragonflies (e.g., Arrenurus), various true bugs (e.g., Hydrachna), and beetles (e.g., Eylais) (Proctor et al. 2015). However, if Torrenticola were utilizing such hosts, we would expect fewer instances of endemism, as well as more dispersal events across the east-west divide. Instead, all eastern Torrenticola diversity could be explained by as little as three dispersal events. We speculate that this pattern can be explained by Torrenticola larval ecology. Torrenticola, like most lotic water mites, parasitize nematocerous flies, especially Chironomidae. Torrenticola are reported from the thoraces (rarely abdomen) of three chironomid subfamilies and ten genera: Tanypodinae (1 genus), Orthocladiinae (6 genera), and Chironominae (3 genera) (Smith & Oliver 1976, 1986). In the proper habitat (clean fast-flowing streams and riffles), Torrenticola adults are typically far more abundant than other water mites and larvae are easily identified by possessing fused coxal plates (unlike all other Lebertioidea [Smith 1982]), which should increase both the likelihood of larvae being sampled on a host and being identified by researchers. However, larval Torrenticola are rarely reported. Unfortunately, Torrenticola larvae are not currently correlated with adults, so there is no way to link species to hosts and we can only conclude that at least some unidentified species parasitize chironomids.

This lack of knowledge has important ramifications because chironomids appear to have different inter- and intra-species natal fidelity. Although only a few studies have addressed adult chironomid dispersal (e.g., Delettre and Morvan 2000, Krosch et al. 2011), they suggest that the propensity for adult chironomids to disperse decreases when the natal stream is bordered by dense vegetation, which appears to confine them to the stream from which they emerged. This trend is not restricted to chironomids as other aquatic insects that inhabit flowing water are also unable to disperse through dense riparian vegetation (Titmus 1980, Jackson and Resh 1989, Peterson et al. 1999, Delettre and Morvan 2000). Conversely, in areas with low vegetation, chironomids have been shown to disperse further (Delettre and Morvan 2000), and under such conditions can even cross into nearby catchments (Krosch et al. 2011).

We speculate that broadly distributed water mite species (e.g., T. ellipsoidalis, T. multiforma, T. sierrensis, T. tahoei, T. malarkeyorum, T. trimaculata, T. irapalpa, T. neoanomala, T. magnexa, T. priapus, and T. sellersorum) disperse primarily during the larval stage by utilizing chironomids with low natal fidelity, whereas species that are confined to a smaller geographic region (e.g., T. solisorta, T. larvata, T. pacificensis, T. rockyensis, T. arktonyx) either utilize chironomids that have high natal fidelity, or they have lost parasitism altogether. As discussed previously, loss of parasitism is especially likely with members of the Miniforma Group, which in addition to high endemism, also possess characters that have been proposed for species that have lost parasitism, such as smaller body size and a larger female genital opening (Smith 1998). In either case (host natal fidelity or loss of parasitism), the result is that species confined to a smaller geographic region may be more dependent on the adult stage for dispersal. A sound understanding of Torrenticola larval ecology, including host associations, is critical to understanding the biology of the genus. Unfortunately, this area of inquiry is inhibited by our limited understanding, not only of water mites, but also chironomids, which are also understudied in North America, leaving most aspects of the biology of nearly all species involved a mystery.

To fully understand the evolution of Torrenticola within North America, analyses are needed that include worldwide taxon sampling and robust analyses of multiple genes, as well as a comprehensive understanding of larval ecology. To this end, we recommend the following areas of study for future investigation in North America: 1) surveys that correlate larval and adult chironomids; 2) dispersal studies on lotic chironomids; 3) surveys that correlate larval and adult water mites; and 4) surveys that investigate host specificity (i.e., identify both water mite larvae on hosts as well as the species of the hosts).

Taxonomy

Torrenticolidae Piersig, 1902

Familial diagnosis. See Fisher et al. (2015).

Torrenticolinae Piersig, 1902

Subfamilial diagnosis. See Fisher et al. (2015)

Torrenticola Piersig, 1896

Type species. T. anomala (Koch, 1837), originally Atractides anomalus

Generic diagnosis. See Fisher et al. (2015)

Descriptions

Torrenticola anoplopalpa Fisher & Dowling, sp. n.

Material examined

HOLOTYPE (♂): USA, New Mexico, Catron County, Glenwood; Whitewater Picnic Area, 8 km east of Rt. 180, (33°22'22"N, 108°50'50"W), 12 July 1987, by IM Smith, IMS870084.

PARATYPES (0 ♀; 0 ♂):

Type deposition

Holotype (♂) deposited in the CNC.

Diagnosis

Torrenticola anoplopalpa are similar to other members of the Rala Group (T. boettgeri, T. dolichodactyla, T. kurtvietsi, T. keesdavidsi, T. lamellipalpis, and T. rala) by being colorless, having incomplete hind coxal margins and being distributed in the southwest. Torrenticola anoplopalpa can be differentiated from all other Rala Group by having a more elongate subcapitulum (ventral length/width ♂ = 4.16 in T. anoplopalpa, 2.04–3.56 in others). Additionally, T. anoplopalpa can be differentiated from all other Rala Group by femur/genu (♂ 1.94 in T. anoplopalpa, 0.98–1.86 in others), except T. keesdavidsi (1.84–1.96).

Description

Female unknown.

Male (Figure 20) (n = 1) (holotype only) with characters of the genus with following specifications.

Dorsum — (640 long; 440 wide) ellipsoid and colorless. Anterio-medial platelets (152.5 long; 60 wide). Anterio-lateral platelets (205 long; 72.5 wide) free from dorsal plate. Dgl-4 much closer to the edge of the dorsum than to the muscle scars (distance between Dgl-4 375). Dorsal plate proportions: dorsum length/width 1.45; dorsal width/distance between Dgl-4 1.17; anterio-medial platelet length/width 2.54; anterio-lateral platelet length/width 2.83; anterio-lateral/anterio-medial length 1.34.

Gnathosoma — Subcapitulum (322.5 long (ventral); 235 long (dorsal); 77.5 tall) elongate and colorless. Rostrum (110 long; 32.5 wide). Chelicerae (275 long) with curved fangs (40 long). Subcapitular proportions: ventral length/height 4.16; rostrum length/width 3.38. Pedipalps short and stocky (especially tibiae) without extensions on femora and genua. Palpomeres: trochanter (35 long); femur (77.5 long); genu (40 long); tibia (47.5 long; 17.5 wide); tarsus (12.5 long). Palpomere proportions: femur/genu 1.94; tibia/femur 0.61; tibia length/width 2.71.

Venter — (775 long; 510 wide) colorless. Gnathosomal bay (127.5 long; 70 wide). Cxgl-4 subapical. Medial suture (70 long). Genital plates (160 long; 120 wide). Additional measurements: Cx-1 (310 long (total); 180 long (medial)); Cx-3 (330 wide); anterior venter (270 long). Ventral proportions: gnathosomal bay length/width 1.82; anterior venter/genital field length 1.69; anterior venter length/genital field width 2.25; anterior venter/medial suture 3.86.

Immatures unknown.

Etymology

Specific epithet (anoplopalpa) refers to the pedipalps, which lack tubercles on the femora and genua, an uncommon condition in Torrenticola, which usually have tuberculate ventral extensions (anoplos, G. unarmed; palpus, L. hand, feeler).

Distribution

Southwest. New Mexico (probably also Arizona) (Figure 19).

Figure 19. 

Torrenticola anoplopalpa sp. n. distribution.

Figure 20. 

Torrenticola anoplopalpa sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Remarks

Unfortunately, we were unable to acquire fresh material of Torrenticola anoplopalpa and so this species is not included in our phylogenetic analyses. However, we were able to examine material preserved in GAW for morphology. The overall appearance, incomplete hind coxal margins, lack of coloration, and distribution, are consistent with placing this species in the Rala Identification Group.

Torrenticola arktonyx Fisher & Dowling, sp. n.

Material examined

HOLOTYPE (♀): USA, North Carolina, Macon County, Rainbow Springs; beside Forest Route 67, 2.0 km south of road to Standing Indian Campground, (35°3'3"N, 83°31'31"W), 1 July 1990, by IM Smith, IMS900072

PARATYPES (4 ♀; 5 ♂): Georgia, USA: 1 ♀ from White County, Helen; beside Road to Anna Ruby Falls just north of Unicoi State Park, (34°44'44"N, 83°43'43"W), 24 September 1992, by IM Smith, IMS920051 • North Carolina, USA: 1 ♀ from Haywood County, Great Smokey Mountains National Park, Big Creek downstream of the bridge at picnic area, (35°45'45"N, 83°6'6"W), 15 September 2009, by AJ Radwell, AJR090008A • 1 ♀ and 2 ♂ from Haywood County, Great Smoky Mountains National Park; Cataloochee; beside Mt. Sterling Rd. near bridge 1.7 km n. of road to Campground, (35°38'38"N, 83°4'4"W), 6 September 2009, by IM Smith, IMS090099 • 2 ♂ from Macon County, Rainbow Springs; beside Forest Route 67, 2.0 km south of road to Standing Indian Campground, (35°3'3"N, 83°31'31"W), 20 September 1991, by IM Smith, IMS910054 • 1 ♂ (ALLOTYPE) from Macon County, Rainbow Springs; beside Forest Route 67, 2.0 km south of road to Standing Indian Campground, (35°3'3"N, 83°31'31"W), 1 July 1990, by IM Smith, IMS900072 • Tennessee, USA: 1 ♀ from Blount County, Middle Prong of the Little River at Tremont, (35°38'38"N, 83°41'41"W), 16 September 2009, by AJ Radwell, AJR090009

Type deposition

Holotype (♀), allotype (♂), and some paratypes (2 ♀; 2 ♂) deposited in the CNC; other paratypes (2 ♀; 2 ♂) deposited in the ACUA.

Diagnosis

Torrenticola arktonyx are similar to species with similar dorsal patterning, such as the Rusetria “4-Plate” group (T. dunni, T. glomerabilis, T. kittatinniana, T. pollani, T. rufoalba, and T. shubini), Elongata Group (T. elongata, T. gorti, and T. reduncarostra), Neoanomala Group (T. interiorensis and T. neoanomala), and T. erectirostra, T. robisoni, T. irapalpa, T. racupalpa, T. skvarlai, and T. bondi. Torrenticola arktonyx can be differentiated from all other Torrenticola by having distinctive longitudinal dark markings on the anterior portion of the dorsal plate that fade posteriorly.

Description

Female (Figure 22) (n = 4) (holotype measurements in parentheses when available) with characters of the genus with following specifications.

Dorsum — (645–680 (670) long; 480–520 (510) wide) ovoid with purple coloration separated into anterior and posterior portions with faint orange medially, also with distinctive longitudinal dark markings on the anterior portion of the dorsal plate that fade posteriorly. Anterio-medial platelets (137.5–145 (137.5) long; 65–70 (65) wide). Anterio-lateral platelets (200–207.5 (200) long; 80–90 (80) wide) free from dorsal plate. Dgl-4 much closer to the edge of the dorsum than to the muscle scars (distance between Dgl-4 335–375 (375)). Dorsal plate proportions: dorsum length/width 1.29–1.34 (1.31); dorsal width/distance between Dgl-4 1.36–1.43 (1.36); anterio-medial platelet length/width 2.07–2.23 (2.12); anterio-lateral platelet length/width 2.31–2.56 (2.50); anterio-lateral/anterio-medial length 1.38–1.48 (1.45).

Gnathosoma — Subcapitulum (345–362.5 (355) long (ventral); 260–270 (265) long (dorsal); 117.5–120 (117.5) tall) with purple coloration. Rostrum (135–135 (135) long; 45–47.5 (45) wide). Chelicerae (335–350 (350) long) with curved fangs (45–55 (45) long). Subcapitular proportions: ventral length/height 2.88–3.09 (3.02); rostrum length/width 2.84–3.00 (3.00). Pedipalps with dentate, flanged ventral extensions on femora and genua. Palpomeres: trochanter (40–42.5 (41.25) long); femur (107.5–111.25 (111.25) long); genu (75–80 (77.5) long); tibia (92.5–95 (95) long; 25–27.5 (25) wide); tarsus (25–25 (25) long). Palpomere proportions: femur/genu 1.38–1.44 (1.44); tibia/femur 0.85–0.86 (0.85); tibia length/width 3.36–3.80 (3.80).

Venter — (815–840 (815) long; 510–600 (600) wide) with purple coloration. Gnathosomal bay (137.5–155 (137.5) long; 82.5–92.5 (92.5) wide). Cxgl-4 subapical. Medial suture (50–60 (50) long). Genital plates (180–187.5 (180) long; 155–162.5 (155) wide). Additional measurements: Cx-1 (300–310 (300) long (total); 150–165 (165) long (medial)); Cx-3 (350–385 (385) wide); anterior venter (230–232.5 (230) long). Ventral proportions: gnathosomal bay length/width 1.49–1.82 (1.49); anterior venter/genital field length 1.24–1.28 (1.28); anterior venter length/genital field width 1.42–1.48 (1.48); anterior venter/medial suture 3.83–4.60 (4.60).

Male (Figure 23) (n = 5) (allotypic measurements in parentheses when available) with characters of the genus with following specifications.

Dorsum — (500–570 (570) long; 400–450 (450) wide) ovoid with purple coloration separated into anterior and posterior portions with faint orange medially, also with distinctive longitudinal dark markings on the anterior portion of the dorsal plate that fade posteriorly. Anterio-medial platelets (110–125 (125) long; 55–62.5 (62.5) wide). Anterio-lateral platelets (155–180 (180) long; 65–75 (75) wide) free from dorsal plate. Dgl-4 much closer to the edge of the dorsum than to the muscle scars (distance between Dgl-4 305–345 (345)). Dorsal plate proportions: dorsum length/width 1.23–1.35 (1.27); dorsal width/distance between Dgl-4 1.30–1.35 (1.30); anterio-medial platelet length/width 2.00–2.18 (2.00); anterio-lateral platelet length/width 2.21–2.69 (2.40); anterio-lateral/anterio-medial length 1.41–1.59 (1.44).

Gnathosoma — Subcapitulum (275–300 (300) long (ventral); 210–235 (235) long (dorsal); 92.5–102.5 (102.5) tall) with purple coloration. Rostrum (110–125 (125) long; 35–40 (40) wide). Chelicerae (270–305 (305) long) with curved fangs (40–45 (40) long). Subcapitular proportions: ventral length/height 2.75–3.08 (2.93); rostrum length/width 3.00–3.20 (3.13). Pedipalps with dentate, flanged ventral extensions on femora and genua. Palpomeres: trochanter (32.5–37.5 (35) long); femur (85–97.5 (97.5) long); genu (60–68.75 (65) long); tibia (75–85 (85) long; 22.5–26.25 (25) wide); tarsus (21.25–25 (22.5) long). Palpomere proportions: femur/genu 1.42–1.50 (1.50); tibia/femur 0.81–0.91 (0.87); tibia length/width 3.10–3.40 (3.40).

Venter — (650–725 (725) long; 430–490 (490) wide) with purple coloration. Gnathosomal bay (125–135 (135) long; 65–72.5 (72.5) wide). Cxgl-4 subapical. Medial suture (75–90 (80) long). Genital plates (160–172.5 (172.5) long; 117.5–130 (130) wide). Additional measurements: Cx-1 (250–290 (290) long (total); 130–150 (145) long (medial)); Cx-3 (295–350 (350) wide); anterior venter (220–260 (260) long). Ventral proportions: gnathosomal bay length/width 1.85–2.08 (1.86); anterior venter/genital field length 1.38–1.55 (1.51); anterior venter length/genital field width 1.87–2.17 (2.00); anterior venter/medial suture 2.72–3.25 (3.25).

Immatures unknown.

Etymology

Specific epithet (arktonyx) refers to the distinctive longitudinal markings on the anterior dorsal plate, which resemble claw marks from a bear (árktos, G. bear; ónyx, G. claw).

Distribution

Southern Appalachians (Figure 21).

Figure 21. 

Torrenticola arktonyx sp. n. distribution.

Figure 22. 

Torrenticola arktonyx sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 23. 

Torrenticola arktonyx sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Remarks

Unfortunately, we were unable to acquire fresh material of Torrenticola arktonyx and therefore this species is not included in our phylogenetic analyses. We were able to examine material preserved in GAW for morphology, but due its unique characteristics, we are unable to place this species into either a species complex or identification group. However, based upon coloration, distribution, and gnathosomal shape, we speculate that future analyses will place this species in the Raptor Complex.

Torrenticola biscutella Fisher & Dowling, sp. n.

Material examined

HOLOTYPE (♀): from USA, Arkansas, Montgomery County, South Fork Ouachita River, access off County Road 17 at Forest Road 903, 29 Jul 2011, by IM Smith, IMS110040, DNA 1263.

PARATYPES (4 ♀; 3 ♂): Arkansas, USA: 1 ♂ (ALLOTYPE) from Montgomery County, Ouachita River (34°34'53.20"N, 93°53'0.16"W), 5 Oct 2007, by AJ Radwell, & HW Robison, AJR070300A • 2 ♂ Montgomery County, Ouachita River (34°34'53.20"N, 93°53'0.16"W), 5 Oct 2007, by AJ Radwell, & HW Robison, AJR070300A • 3 ♀ from Montgomery County, South Fork Ouachita River, access off County Road 17 at Forest Road 903, 29 Jul 2011, by IM Smith, IMS110040 • Missouri, USA: 1 ♀ from Crawford County, Huzzah Creek, Red Bluff campground, off Road V east of Davisville, 23 Jul 2011, by IM Smith, IMS110029.

Type deposition

Holotype (♀), allotype (♂), and some paratypes (2 ♀; 1 ♂) deposited in the CNC; other paratypes (2 ♀; 1 ♂) deposited in the ACUA.

Diagnosis

Torrenticola biscutella are similar to other members of the Rusetria “Eastern 2-Plates” group (T. caerulea, T. delicatexa, T. feminellai, T. indistincta, T. malarkeyorum, T. microbiscutella, T. pendula, T. sellersorum, T. tysoni, T. ululata, and T. whitneyae) in having anterio-lateral platelets fused to the dorsal plate, having dorsal coloration separated into anterior and posterior portions (except T. indistincta and T. ululata), and being distributed in the east. It is one of only four Eastern 2-Plates that have dark, bold, bluish-purple coloration (also T. pendula, T. sellersorum, and T. tysoni). Torrenticola biscutella can be differentiated from T. caerulea, T. ululata, T. indistincta, and T. feminellai by dorsal coloration and pattern. T. biscutella can be differentiated from T. tysoni by having a stockier rostrum (length/width = 2.55–2.83 in T. biscutella, 3.06–3.50 in T. tysoni). Female T. biscutella can be differentiated from female T. malarkeyorum by having a shorter subcapitulum (ventral length = 290–315 in T. biscutella, 317.5–335 in T. malarkeyorum). Male T. biscutella can be differentiated from male T. malarkeyorum by having a slightly rounder dorsum (length/width 1.37–1.42 in T. biscutella, 1.42–1.56 in T. malarkeyorum). Additionally, although T. biscutella and T. malarkeyorum have the same dorsal coloration and pattern, often the coloration is bold in T. biscutella and faint in T. malarkeyorum. Female T. biscutella can be differentiated from female T. delicatexa by having a shorter genital field (152.5–167.5 in T. biscutella, 175–198 in T. delicatexa) and male T. biscutella can be differentiated from male T. delicatexa by having a slightly rounder dorsum (length/width = 1.37–1.42 in T. biscutella, 1.44–1.56 in T. delicatexa). Female T. biscutella can be differentiated from female T. sellersorum by anterior venter/genital field length (0.82–0.88 in T. biscutella, 0.69–0.77 in T. sellersorum). Male T. biscutella can be differentiated from male T. sellersorum by having slightly stockier anterio-lateral platelets (length/width = 2.58–2.74 in T. biscutella, 2.76–3.00 in T. sellersorum). T. biscutella can be differentiated from T. pendula by having a stockier gnathosomal bay (1.55–1.85 in T. biscutella, 2.42–2.9 in T. pendula); more elongate tibiae (3.11–3.45 in T. biscutella, 2.78–3.05 in T. biscutella); and by dorsal pattern. T. biscutella can be differentiated from T. microbiscutella by having a more ovoid dorsum (length/width = 1.33–1.42 in T. biscutella, 1.63–1.75 in T. microbiscutella) and by anterior venter/genital field width (♀ = 0.84–0.91 in T. biscutella, 1.25–1.33 in T. microbiscutella; ♂ = 1.68–1.80 in T. biscutella and 1.95–2.29 in T. microbiscutella). T. biscutella can be differentiated from T. whitneyae by having more elongate pedipalpal tibiae (3.11–3.45 in T. biscutella, 2.42–2.95 in T. whitneyae) and by anterior venter/genital field length (♀ = 0.82–0.88 in T. biscutella, 0.59–0.75 in T. whitneyae; ♂ = 1.55–1.76 in T. biscutella and 1.37–1.43 in T. whitneyae).

Description

Female (Figure 25) (n = 4) (holotype measurements in parentheses when available) with characters of the genus with following specifications.

Dorsum — (560–630 (560) long; 420–455 (420) wide) ovoid with bluish-purple coloration separated into anterior and posterior portions, and bold or faint orange medially. Anterio-medial platelets (122.5–135 (122.5) long; 40–45 (40) wide). Anterio-lateral platelets (140–170 (140) long; 62.5–75 (62.5) wide) fused to dorsal plate. Dgl-4 much closer to the edge of the dorsum than to the muscle scars (distance between Dgl-4 300–330 (300)). Dorsal plate proportions: dorsum length/width 1.33–1.38 (1.33); dorsal width/distance between Dgl-4 1.38–1.40 (1.40); anterio-medial platelet length/width 3.00–3.31 (3.06); anterio-lateral platelet length/width 2.24–2.48 (2.24); anterio-lateral/anterio-medial length 1.14–1.28 (1.14).

Gnathosoma — Subcapitulum (290–315 (290) long (ventral); 207–240 (208) long (dorsal); 137.5–155 (137.5) tall) colorless. Rostrum (110–125 (110) long; 42.5–47.5 (42.5) wide). Chelicerae (286–335 (286) long) with curved fangs (55–70 (56) long). Subcapitular proportions: ventral length/height 2.02–2.11 (2.11); rostrum length/width 2.56–2.67 (2.59). Pedipalps with tuberculate ventral extensions on femora and genua. Palpomeres: trochanter (43.75–50 (43.75) long); femur (107.5–122.5 (107.5) long); genu (65–72.5 (65) long); tibia (80–86.25 (80) long; 23.75–25 (23.75) wide); tarsus (20–20 (20) long). Palpomere proportions: femur/genu 1.59–1.74 (1.65); tibia/femur 0.69–0.74 (0.74); tibia length/width 3.35–3.45 (3.37).

Venter — (660–740 (660) long; 488–544 (489) wide) with faint bluish-purple coloration. Gnathosomal bay (151.25–172.5 (151.25) long; 97.5–100 (100) wide). Cxgl-4 subapical. Medial suture absent. Genital plates (152.5–167.5 (152.5) long; 142.5–160 (142.5) wide). Additional measurements: Cx-1 (274–309 (275) long (total); 118–135 (121) long (medial)); Cx-3 (319–392 (319) wide); anterior venter (130–147.5 (130) long). Ventral proportions: gnathosomal bay length/width 1.55–1.73 (1.55); anterior venter/genital field length 0.82–0.88 (0.85); anterior venter length/genital field width 0.84–0.91 (0.91).

Male (Figure 26) (n = 3) (allotypic measurements in parentheses when available) with characters of the genus with following specifications.

Dorsum — (430–445 (440) long; 310–315 (310) wide) ovoid with bluish-purple coloration separated into anterior and posterior portions, and bold or faint orange medially. Anterio-medial platelets (97.5–97.5 (97.5) long; 33.75–36.25 (35) wide). Anterio-lateral platelets (122.5–130 (130) long; 45–50 (47.5) wide) fused to dorsal plate. Dgl-4 much closer to the edge of the dorsum than to the muscle scars (distance between Dgl-4 230–242.5 (230)). Dorsal plate proportions: dorsum length/width 1.37–1.42 (1.42); dorsal width/distance between Dgl-4 1.30–1.35 (1.35); anterio-medial platelet length/width 2.69–2.89 (2.79); anterio-lateral platelet length/width 2.58–2.74 (2.74); anterio-lateral/anterio-medial length 1.26–1.33 (1.33).

Gnathosoma — Subcapitulum (230–235 (235) long (ventral); 175–177.5 (177) long (dorsal); 20–20 (20) tall) colorless. Rostrum (85–92.5 (92.5) long; 30–36.25 (36.25) wide). Chelicerae (225–241 (241) long) with curved fangs (45–50 (46) long). Subcapitular proportions: ventral length/height 2.29–2.47 (2.29); rostrum length/width 2.55–2.83 (2.55). Pedipalps with tuberculate ventral extensions on femora and genua. Palpomeres: trochanter (23.75–37.5 (37.5) long); femur (85–90 (90) long); genu (52.5–55 (55) long); tibia (68.75–72.5 (72.5) long; 21.25–22.5 (22.5) wide); tarsus (17.5–20 (20) long). Palpomere proportions: femur/genu 1.62–1.64 (1.64); tibia/femur 0.81–0.82 (0.81); tibia length/width 3.11–3.24 (3.22).

Venter — (510–525 (525) long; 335–380 (336) wide) with faint bluish-purple coloration. Gnathosomal bay (115–122.5 (122.5) long; 65–67.5 (67.5) wide). Cxgl-4 subapical. Medial suture (60–65 (65) long). Genital plates (102.5–110 (102.5) long; 100–100 (100) wide). Additional measurements: Cx-1 (215–226 (226) long (total); 99–110 (100) long (medial)); Cx-3 (252–275 (252) wide); anterior venter (167.5–180 (180) long). Ventral proportions: gnathosomal bay length/width 1.70–1.85 (1.81); anterior venter/genital field length 1.55–1.76 (1.76); anterior venter length/genital field width 1.68–1.80 (1.80); anterior venter/medial suture 2.68–2.83 (2.77).

Immatures unknown.

Etymology

Specific epithet (biscutella) refers to the appearance of only two anterio-dorsal platelets due to the fusion of lateral platelets with the dorsal shield (bi-, L. two; scutella, L. little plate).

Distribution

Interior Highlands (both Ozarks and Ouachitas), likely endemic (Figure 24).

Figure 24. 

Torrenticola biscutella sp. n. distribution.

Figure 25. 

Torrenticola biscutella sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 26. 

Torrenticola biscutella sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Remarks

Torrenticola biscutella groups with other members of the Rusetria Complex with high support and specimens of this species were less than 2% different in COI sequence from each other. In all analyses, T. biscutella groups with two other morphologically similar species: T. malarkeyorum and T. caerulea. These three species are 3–5% different from each other in COI sequence. The three of these species are morphologically similar to the more distantly-related T. delicatexa. Of these four species, the range of T. biscutella only overlaps with T. malarkeyorum in the Ozark Mountains and these species are easily differentiated by color. T. biscutella is the only one of these four species known from the Ouachita Mountains, and it is not known from east of the Mississippi River, where the other three species are distributed (only T. malarkeyorum is known from west of the Mississippi River).

Based upon overall similarity, dorso-lateral platelet fusion, and distribution, we were able to place this species within the Eastern 2-Plate Identification Group.

This species hypothesis is supported by biogeography, low COI divergence within the species (0–2%) and high divergence between species (3–15%), and by morphological characters outlined in the diagnoses.

Torrenticola bittikoferae Crowell, 1960

T. bittikoferae: Crowell 1960: 36; 1961: 330 • Johnston 1965: 44 • Modlin and Gannon 1973: 219, 221 • Viets 1987: 756.

Material examined

PARATYPES (0 ♀; 2 ♂): Ohio, USA: 2 ♂ from Ottawa County, Middle Bass Island, rubble beach, 29 June 1954, by R Crowell.

Type deposition

Holotype (♀) and some paratypes (unspecified number) deposited in the Chicago Natural History Museum (unexamined; types not located); other paratypes (2 ♂) deposited in the OSUAC.

Diagnosis

Torrenticola bittikoferae are similar to other members of the Tricolor Group (T. cardia, T. dimorpha, T. hoosieri, T. kringi, T. larvata, T. mohawk, T. olliei, T. pearsoni, T. sierrensis, T. tricolor, T. trimaculata, and T. unimaculata) in having a short, conical rostrum. Torrenticola bittikoferae can be differentiated from most other Tricolor Complex (except T. hoosieri, T. pearsoni, and T. dimorpha) by being colorless, whereas most other members have bold patterning. T. bittikoferae can be differentiated from T. hoosieri by having ventral extensions on the pedipalp femora and genua (lacking in T. hoosieri) and having stockier pedipalp tibiae (length/width = 2.7–2.8 in T. bittikoferae, 3.6–4.4 in T. hoosieri). T. bittikoferae can be differentiated from T. pearsoni by having Dgl-4 further from the dorsal edge (dorsal width/distance between Dgl-4 = 1.6–1.7 in T. bittikoferae, 1.2–1.3 in T. pearsoni); stockier pedipalp tibiae (length/width = 2.7–2.8 in T. bittikoferae, 3.0–3.3 in T. pearsoni); and a more elongate rostrum (length/width = 1.8–1.9 in T. bittikoferae, 2.0–2.4 in T. pearsoni). Torrenticola bittikoferae can be differentiated from T. dimorpha by having an unmodified dorsal plate (T. dimorpha has a dorsal plate medial extension covering nearly half the length of the anterio-medial platelets) and by males having unmodified pedipalps (male T. dimorpha have large, highly modified pedipalps which are expanded vertically and laterally).

Re-description

Male (Figure 28) (n = 2) with characters of the genus with following specifications.

Dorsum — (620–670 long; 500–530 wide) circular and colorless. Anterio-medial platelets (132.5–137.5 long; 70–70 wide). Anterio-lateral platelets (192.5–202.5 long; 90–92.5 wide) free from dorsal plate. Dgl-4 approaching midway between muscle scars and dorsum edge (distance between Dgl-4 305–330). Dorsal plate proportions: dorsum length/width 1.24–1.26; dorsal width/distance between Dgl-4 1.61–1.64; anterio-medial platelet length/width 1.89–1.96; anterio-lateral platelet length/width 2.14–2.19; anterio-lateral/anterio-medial length 1.40–1.53.

Gnathosoma — Subcapitulum (265 long (ventral); 202.5 long (dorsal); 125 tall) colorless. Rostrum (95–100 long; 52.5–52.5 wide). Chelicerae (260 long) with curved fangs (50 long) short and conical. Subcapitular proportions: ventral length/height 2.12; rostrum length/width 1.81–1.90. Pedipalps with tuberculate ventral extensions with dentate tip on femora and tuberculate ventral extensions on genua. Palpomeres: trochanter (42.5–42.5 long); femur (101.25–107.5 long); genu (72.5–75 long); tibia (87.5–90 long; 32.5–32.5 wide); tarsus (25–35 long). Palpomere proportions: femur/genu 1.40–1.43; tibia/femur 0.81–0.89; tibia length/width 2.69–2.77.

Venter — (790–800 long; 610–680 wide) colorless. Gnathosomal bay (122.5–125 long; 87.5–100 wide). Medial suture (102.5–117.5 long). Genital plates (137.5–142.5 long; 115–115 wide). Additional measurements: Cx-1 (270–280 long (total); 152.5–152.5 long (medial)); Cx-3 (405–410 wide); anterior venter (270–287.5 long). Ventral proportions: gnathosomal bay length/width 1.23–1.43; anterior venter/genital field length 1.96–2.02; anterior venter length/genital field width 2.35–2.50; anterior venter/medial suture 102.5–117.5.

Female type specimens unavailable for present study.

Immatures unknown.

Etymology

Robert Crowell (1960) named the specific epithet (bittikoferae) after Lelia Bittikofer, his high school biology teacher.

Distribution

Known only from type locality: Lake Erie, Ohio (Figure 27).

Figure 27. 

Torrenticola bittikoferae distribution.

Figure 28. 

Torrenticola bittikoferae male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Remarks

Unfortunately, we were unable to acquire fresh specimens of Torrenticola bittikoferae and therefore this species was not included in our phylogenetic analyses. We were able to examine paratypes of two males, neither of which were dissected during slide preparation, making precise examination difficult. These specimens were remounted, but proper dissection risked fully damaging the specimen and was therefore avoided. The images in Figure 28A–B that appear to display a properly dissected specimen, were created by photographing each section (dorsum and venter) of the fully intact specimen and digitally editing the photographs so that the dorsum and venter could be easily compared with other species.

The overall appearance, short conical rostrum, and distribution of this species allows us to places it within the Tricolor Complex and Tricolor Identification Group.

Torrenticola boettgeri K.O. Viets, 1977

Torrenticola boettgeri K.O. Viets, 1977a: 89.

Torrenticola esbelta Cramer, 1992: 22.

Material examined

(1 ♀; 4 ♂) . New Mexico, USA: 1 ♀ and 1 ♂ from Catron County, Glenwood; Whitewater Picnic Area 8 km east of Rt. 180, (33°22'22"N, 108°50'50"W), 12 July 1987, by IM Smith, IMS870084 • 3 ♂ from Catron County, beside Rt. 15, 65 km north of Rt. 180, Silver City, (33°12'12"N, 108°13'13"W), 10 July 1987, by IM Smith, IMS870081A

Type deposition

Holotype (♀), prep. no. 6381 SMF, Viets collection (not examined).

Diagnosis

Torrenticola boettgeri are similar to other members of the Rala Group (T. anoplopalpa, T. dolichodactyla, T. keesdavidsi, T. kurtvietsi, T. lamellipalpis, and T. rala) by being colorless, having incomplete hind coxal margins and being distributed in the southwest. T. boettgeri can be differentiated from all other Rala Group by having a more elongate dorsum (length/width ♀ = 1.74–1.82 in T. boettgeri, 1.21–1.60 in others) and a stockier subcapitulum (ventral length/width = 1.96 in T. boettgeri, 2.06–3.52 in others; ♂ = 2.04–2.07 in T. boettgeri, 2.14–4.16 in others).

Re-description

Female (Figure 30) (n = 3: one specimen examined from New Mexico; measurements from two additional specimens are included based upon those listed in Goldschmidt (2007) for K.O. Viets’s (1977a) specimen from Guatemala and Cramer’s (1992) specimen from Mexico) with characters of the genus with following specifications. Note: measurements below are from the above three combined sources; those in parentheses are from the Guatemalan holotype (Viets 1977a) as listed in Goldschmidt (2007).

Dorsum — (668–800 (675) long; 367–440 (440) wide) ovoid, elongate, and colorless. Anterio-medial platelets (103–135 (126) long; 38–52.5 wide). Anterio-lateral platelets (179–200 (199) long; 47–70 wide) free from dorsal plate. Dgl-4 much closer to the edge of the dorsum than to the muscle scars (distance between Dgl-4 370). Dorsal plate proportions: dorsum length/width 1.53–1.82 (1.82); dorsal width/distance between Dgl-4 1.19; anterio-medial platelet length/width 2.57–2.71; anterio-lateral platelet length/width 2.86–3.81; anterio-lateral/anterio-medial length 1.48–1.74 (1.58).

Gnathosoma — Subcapitulum (245–280 (245) long (ventral); 185 long (dorsal); 142.5 tall) colorless. Rostrum (70–87.5 (70) long; 40 wide). Chelicerae with curved fangs (65 long). Subcapitular proportions: ventral length/height 1.96; rostrum length/width 2.19. Pedipalps short and stocky (especially tibiae) without extensions on femora and genua. Palpomeres: trochanter (32.5 long); femur (76.25 long); genu (65 long); tibia (32.5 long; 17.5 wide); tarsus (15 long). Palpomere proportions: femur/genu 1.17; tibia/femur 0.43; tibia length/width 1.86.

Venter — (668–955 (845) long; 452–570 (500) wide) colorless. Gnathosomal bay (164–200 (164) long; 45–50 (47) wide). Cxgl-4 apical. Medial suture (65–73 (73) long). Genital plates (170–188 (170) long; 148–160 (148) wide). Additional measurements: Cx-1 (340 long (total); 133–135 (133) long (medial)); Cx-3 (350 wide); anterior venter (212.5 long). Ventral proportions: gnathosomal bay length/width 3.49–4.44 (3.49); anterior venter/genital field length 1.15; anterior venter length/genital field width 1.33; anterior venter/medial suture 3.27.

Male (Figure 31) (n = 4: new specimens from New Mexico)

Dorsum — (710–780 long; 400–430 wide) ovoid, elongate, and colorless. Anterio-medial platelets (122.5–140 long; 45–50 wide). Anterio-lateral platelets (175–192.5 long; 55–62.5 wide) free from dorsal plate. Dgl-4 much closer to the edge of the dorsum than to the muscle scars (distance between Dgl-4 335–365). Dorsal plate proportions: dorsum length/width 1.74–1.81; dorsal width/distance between Dgl-4 1.16–1.19; anterio-medial platelet length/width 2.68–2.95; anterio-lateral platelet length/width 2.96–3.27 (3.08); anterio-lateral/anterio-medial length 1.25–1.47.

Gnathosoma — Subcapitulum (255–285 long (ventral); 165–187.5 long (dorsal); 125–137.5 tall) colorless. Rostrum (80–92.5 long; 35–40 wide). Chelicerae (285–310 long) with curved fangs (62.5–65 long). Subcapitular proportions: ventral length/height 2.04–2.07; rostrum length/width 2.13–2.36. Pedipalps short and stocky (especially tibiae) without extensions on femora and genua. Palpomeres: trochanter (30–31.25 long); femur (58.75–72.5 long); genu (60–62.5 long); tibia (32.5–37.5 long; 17.5–17.5 wide); tarsus (15–15 long). Palpomere proportions: femur/genu 0.98–1.16; tibia/femur 0.48–0.55; tibia length/width 1.86–2.14.

Venter — (800–960 long; 485–510 wide) colorless. Gnathosomal bay (192.5–205 long; 40–50 wide). Cxgl-4 apical. Medial suture (105–125 long). Genital plates (160–177.5 long; 120–125 wide). Additional measurements: Cx-1 (305–330 long (total); 115–130 long (medial)); Cx-3 (320–325 wide); anterior venter (235–262.5 long). Ventral proportions: gnathosomal bay length/width 3.85–5.00; anterior venter/genital field length 1.38–1.59; anterior venter length/genital field width 1.96–2.13; anterior venter/medial suture 2.00–2.24.

Immatures unknown.

Etymology

Viets (1977a) named the specific epithet (boettgeri) in honor of Klaus Böttger of the University of Kiel, Germany, who collected the type specimen in Rio Chilax near Cobán, Guatemala.

Distribution

New Mexico (probably also Arizona) and extending southward into Mexico and Guatemala (Figure 29).

Figure 29. 

Torrenticola boettgeri distribution. Blue star represents type locality (Viets 1977a); blue diamond represents additional published record (Cramer 1992, as T. esbelta comb. n.); and red circles represent new records and material examined.

Figure 30. 

Torrenticola boettgeri female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 31. 

Torrenticola boettgeri male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Remarks

Unfortunately, we were unable to acquire fresh material of Torrenticola boettgeri and therefore this species is not included in our phylogenetic analyses. We were also unable to examine types, but were able to examine new material from New Mexico. The overall appearance, incomplete hind coxal margins, distribution, and lack of coloration are consistent with placing this species in the Rala Identification Group.

Viets (1977a) described T. boettgeri from a single female collected in Guatemala. Cramer (1992) described T. esbelta from three females collected in one stream (Peña Blanca) in San Francisco Oxtotilpan, State of Mexico, Mexico. Cramer (1992) differentiated T. esbelta from T. boettgeri by pedipalpal tibia length: 36 µm in T. esbelta; 26-27 µm (right, left, respectively). Given our experience with the variability of tibial length across species, which often range well over 10 µm, and especially considering the very few number of specimens examined, we do not consider slight variations in pedipalp tibial length to be good evidence for separate species. Furthermore, our material from New Mexico includes specimens with tibiae in between the previously recorded specimens (32.5 µm in our single female specimen; 32.5–37.7 µm in males). Therefore, we consider T. esbelta as a junior synonym of T. boettgeri.

Torrenticola bondi Fisher & Dowling, sp. n.

Material examined

HOLOTYPE (♀): from USA, North Carolina, Haywood County, Great Smokey Mountains National Park, Cataloochee (35°37'31"N, 83°6'46"W), 20 Sep 2010, by IM Smith, IMS100148, DNA 1431.

PARATYPES (4 ♀; 5 ♂): North Carolina, USA: 1 ♂ (ALLOTYPE) from Haywood County, Great Smoky Mountains National Park; Cataloochee; beside road to Nellie 0.2 km west of Pretty Hollow Gap Trailhead, (35°37'37"N, 83°6'6"W), 16 June 2006, by IM Smith, IMS060012 • 1 ♀ from Haywood County, Great Smoky Mountains National Park; Cataloochee; beside road to Nellie 0.2 km west of Pretty Hollow Gap Trailhead, (35°37'37"N, 83°6'6"W), 16 June 2006, by IM Smith, IMS060012 • 1 ♀ and 1 ♂ from Haywood County, Great Smoky Mountains National Park; Cataloochee; beside road to Nellie 0.8 km west of Pretty Hollow Gap Trailhead, (35°38'38"N, 83°4'4"W), 20 June 2006, by IM Smith, IMS060022 • 1 ♂ from Haywood County, Great Smoky Mountains National Park; Cataloochee; beside road to Nellie 0.4 km west of Pretty Hollow Gap Trailhead, (35°37'37"N, 83°6'6"W), 3 October 2007, by IM Smith, IMS0701002 • ♀ and 2 ♂ from Haywood County, Great Smoky Mountains National Park; Waterville; Big Creek Picnic Area, (35°45'45"N, 83°6'6"W), 26 September 2007, by IM Smith, IMS070089

Type deposition

Holotype (♀), allotype (♂), and some paratypes (2 ♀; 2 ♂) deposited in the CNC; other paratypes (2 ♀; 2 ♂) deposited in the ACUA.

Diagnosis

Torrenticola bondi are similar to species with similar dorsal patterning, such as the Rusetria “4-Plate” group (T. dunni, T. glomerabilis, T. kittatinniana, T. pollani, T. rufoalba, and T. shubini), Elongata Group (T. elongata, T. gorti, and T. reduncarostra), Neoanomala Group (T. interiorensis and T. neoanomala), and T. erectirostra, T. robisoni, T. irapalpa, T. racupalpa, T. skvarlai, and T. arktonyx. They can be differentiated from Rusetria 4-Plates and T. skvarlai by having distinct hind coxal margins. T. bondi can be differentiated from T. erectirostra and T. robisoni by having a straight, anteriorly-directed rostrum (upturned in others). T. bondi can be differentiated from T. arktonyx by having an unmodified dorsal plate (T. arktonyx has distinctive longitudinal dark markings on the anterior portion of the dorsal plate that fade posteriorly). T. bondi can be differentiated from T. racupalpa and T. irapalpa by having a more ovoid dorsum (dorsum length/width ♀ = 1.35–1.41 in T. bondi, 1.17–1.28 in others; ♂ = 1.32–1.45 in T. bondi, 1.20–1.30 in others) and shorter pedipalpal tibiae (♀ = 90–98 in T. bondi, 100–125 in others, ♂ = 77–83 in T. bondi, 87–110 in others). T. bondi can be differentiated from the Elongata Group by having a less elongate dorsum (length/width ♀ = 1.35–1.41 in T. bondi, 1.45–2.08 in Elongata Group; ♂ = 1.32–1.45 in T. bondi, 1.51–1.70 in Elongata Group) and having a stockier rostrum (length/width = 2.76–3.13 in T. bondi, 3.24–4.00 in Elongata Group). T. bondi can be differentiated from the Neoanomala Group by having a shorter medial suture (♀ = 10–15 in T. bondi, 22–40 in Neoanomala Group; ♂ = 55–70 in T. bondi, 75–108 in Neoanomala Group) and anterior venter/genital field width (♀ = 1.15–1.25 in T. bondi, 1.31–1.45 in Neoanomala Group; ♂ = 1.95–2.05 in T. bondi, 2.09–2.66 in Neoanomala Group).

Description

Female (Figure 33) (n = 5) (holotype measurements in parentheses when available) with characters of the genus with following specifications.

Dorsum — (620–670 (620) long; 440–490 (440) wide) ovoid with bluish-purple coloration separated into anterior and posterior portions and faint orange medially. Anterio-medial platelets (132.5–147.5 (132.5) long; 55–62.5 (55) wide). Anterio-lateral platelets (192.5–200 (192.5) long; 67.5–77.5 (67.5) wide) free from dorsal plate. Dgl-4 approaching midway between muscle scars and dorsum edge (distance between Dgl-4 270–290 (270)). Dorsal plate proportions: dorsum length/width 1.35–1.41 (1.41); dorsal width/distance between Dgl-4 1.63–1.72 (1.63); anterio-medial platelet length/width 2.20–2.52 (2.41); anterio-lateral platelet length/width 2.58–2.85 (2.85); anterio-lateral/anterio-medial length 1.34–1.41 (1.45).

Gnathosoma — Subcapitulum (355–380 (355) long (ventral); 255–295 (255) long (dorsal); 135–150 (135) tall) faint bluish purple coloration. Rostrum (145–155 (145) long; 50–55 (52.5) wide). Chelicerae (354–385 (354) long) with curved fangs 52–75 (52) long). Subcapitular proportions: ventral length/height 2.50–2.63 (2.63); rostrum length/width 2.76–3.00 (2.76). Pedipalps with tuberculate ventral extensions on femora and genua. Palpomeres: trochanter 47.5–52.5 (47.5) long); femur (120–140 (120) long); genu (67.5–77.5 (67.5) long); tibia (90–97.5 (92.5) long; 22.5–25 (22.5) wide); tarsus (17.5–20 (17.5) long). Palpomere proportions: femur/genu 1.74–1.83 (1.78); tibia/femur 0.67–0.77 (0.77); tibia length/width 3.90–4.11 (4.11).

Venter — (760–840 (760) long; 509–580 (509) wide) with faint bluish-purple coloration. Gnathosomal bay (170–190 (170) long; 90–102.5 (102.5) wide). Cxgl-4 subapical. Medial suture (10–15 (10) long). Genital plates (175–180 (175) long; 150–160 (150) wide). Additional measurements: Cx-1 (306–330 (306) long (total); 108–155 (108) long (medial)); Cx-3 (365–410 (382) wide); anterior venter (177.5–187.5 (187.5) long). Ventral proportions: gnathosomal bay length/width 1.66–2.11 (1.66); anterior venter/genital field length 0.99–1.07 (1.07); anterior venter length/genital field width 1.15–1.25 (1.25); anterior venter/medial suture 12–18.75 (18.75).

Male (Figure 34) (n = 5) (allotypic measurements in parentheses when available) with characters of the genus with following specifications.

Dorsum — (515–550 (550) long; 380–410 (380) wide) ovoid with bluish-purple coloration separated into anterior and posterior portions and faint orange medially. Anterio-medial platelets (112.5–122.5 (122.5) long; 47.5–57.5 (57.5) wide). Anterio-lateral platelets (162.5–190 (180) long; 55–67.5 (67.5) wide) free from dorsal plate. Dgl-4 approaching midway between muscle scars and dorsum edge (distance between Dgl-4 235–260 (250)). Dorsal plate proportions: dorsum length/width 1.32–1.45 (1.45); dorsal width/distance between Dgl-4 1.48–1.62 (1.52); anterio-medial platelet length/width 2.13–2.47 (2.13); anterio-lateral platelet length/width 2.65–3.17 (2.67); anterio-lateral/anterio-medial length 1.44–1.57 (1.47).

Gnathosoma — Subcapitulum (295–305 (305) long (ventral); 220–230 (225) long (dorsal); 107.5–112.5 (112.5) tall) faint bluish purple coloration. Rostrum (120–125 (125) long; 40–40 (40) wide). Chelicerae (285–295 (290) long) with curved fangs 50–55 (55) long). Subcapitular proportions: ventral length/height 2.68–2.79 (2.71); rostrum length/width 3.00–3.13 (3.13). Pedipalps with tuberculate ventral extensions on femora and genua. Palpomeres: trochanter 40–42.5 (42.5) long); femur (102.5–107.5 (105) long); genu (60–62.5 (60) long); tibia (77.5–82.5 (77.5) long; 21.25–23.75 (22.5) wide); tarsus (17.5–20 (20) long). Palpomere proportions: femur/genu 1.68–1.75 (1.75); tibia/femur 0.74–0.80 (0.74); tibia length/width 3.44–3.88 (3.44).

Venter — (660–670 (665) long; 420–490 (460) wide) with faint bluish-purple coloration. Gnathosomal bay (140–150 (142.5) long; 75–90 (80) wide). Cxgl-4 subapical. Medial suture (55–70 (60) long). Genital plates (132.5–142.5 (140) long; 107.5–112.5 (110) wide). Additional measurements: Cx-1 (270–290 (285) long (total); 125–140 (135) long (medial)); Cx-3 (310–340 (335) wide); anterior venter (215–220 (220) long). Ventral proportions: gnathosomal bay length/width 1.61–1.93 (1.78); anterior venter/genital field length 1.54–1.66 (1.57); anterior venter length/genital field width 1.95–2.05 (2.00); anterior venter/medial suture 3.07–3.91 (3.67).

Immatures unknown.

Etymology

Specific epithet (bondi) named in honor of arachnologist Jason Bond, for his research on species delimitation and integrative taxonomy, which has been an inspiration to JRF, and for his thoughtful career advice, which was greatly appreciated.

Distribution

Known only from Great Smoky Mountains National Park, Haywood County, North Carolina (Figure 32).

Figure 32. 

Torrenticola bondi sp. n. distribution.

Figure 33. 

Torrenticola bondi sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 34. 

Torrenticola bondi sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Remarks

Torrenticola bondi groups with other members of the Raptor Complex in all analyses with high support. Only one specimen could be acquired for use in our analyses, so differences in COI sequence across specimens could not be investigated, but this single specimen was greater than 5% different in COI sequence from sister species. In all analyses, T. bondi grouped with members of the Elongata Identification Group (T. elongata and T. gorti). However, the position of this clade varied with analysis.

This species hypothesis is supported by high divergence between species (3–15%) and by the morphological characters outlined in the diagnosis.

Torrenticola caerulea Fisher & Dowling, sp. n.

Material examined

HOLOTYPE (♀): from USA, Tennessee, Wayne County, beside service road parallel to Natchez Trace Parkway (35°15'9"N, 87°37'53"W), 27 Sep 2010, by IM Smith, IMS100160, DNA 1882.

PARATYPES (5 ♀; 3 ♂): Tennessee, USA: 2 ♀ from Wayne County, beside Natchez Trace Parkway at Lower Glenrock Branch Picnic Area, (35°15'15"N, 87°37'37"W), 2 June 1992, by IM Smith, IMS920021 • 1 ♂ (ALLOTYPE) from Wayne County, Lower Glenrock Picnic Area off Natchez Trace Parkway, (35°15'15"N, 87°37'37"W), 5 October 2005, by IM Smith, IMS050119A • 1 ♀ and 2 ♂ from Wayne County, Lower Glenrock Picnic Area off Natchez Trace Parkway, (35°15'15"N, 87°37'37"W), 5 October 2005, by IM Smith, IMS050119A • 2 ♀ from Wayne County, beside service road parallel to Natchez Trace Parkway (35°15'9"N, 87°37'53"W), 27 Sep 2010, by IM Smith, IMS100160.

Type deposition

Holotype (♀), allotype (♂), and some paratypes (3 ♀; 1 ♂) deposited in the CNC; other paratypes (2 ♀; 1 ♂) deposited in the ACUA.

Diagnosis

Torrenticola caerulea are similar to other members of the Rusetria “Eastern 2-Plates” group (T. biscutella, T. delicatexa, T. feminellai, T. indistincta, T. malarkeyorum, T. microbiscutella, T. pendula, T. sellersorum, T. tysoni, T. ululata, and T. whitneyae) in having anterio-lateral platelets fused to the dorsal plate, having dorsal coloration separated into anterior and posterior portions (except T. ululata and T. indistincta), and being distributed in the east. T. caerulea can be differentiated from all other Eastern 2-Plates by having faint blue coloration. T. caerulea can be differentiated from T. ululata and T. feminellai by dorsal coloration and pattern. T. caerulea can be differentiated from T. tysoni by having a less elongate rostrum (length/width = 2.67–2.96 in T. caerulea, 3.06–3.50 in T. tysoni). T. caerulea can be differentiated from T. pendula by having a more elongate gnathosomal bay (1.40–2.09 in T. caerulea, 2.42–2.90 in T. pendula) and more elongate pedipalpal tibiae (3.11–3.83 in T. caerulea, 2.78–3.05 in T. pendula). T. caerulea can be differentiated from T. microbiscutella by having a less elongate dorsum (length/width = 1.32–1.56 in T. caerulea, 1.63–1.75 in T. microbiscutella). T. caerulea can be differentiated from T. whitneyae by having more elongate pedipalpal tibiae (length/width = 3.11–3.83 in T. caerulea, 2.42–2.95 in T. whitneyae) and by anterior venter/genital field width (♀ = 0.9–1.04 in T. caerulea, 0.67–0.80 in T. whitneyae; ♂ = 1.71–1.83 in T. caerulea, 1.5–1.54 in T. whitneyae). Female T. caerulea can be differentiated from female T. biscutella by having slightly more elongate pedipalpal tibae (length/width = 3.5–3.83 in T. caerulea, 3.35–3.45 in T. biscutella). Male T. caerulea can be differentiated from male T. biscutella by anterior venter/medial suture (2.4–2.57 in T. caerulea, 2.68–2.83 in T. biscutella). Female T. caerulea can be differentiated from female T. malarkeyorum, T. sellersorum, T. delicatexa, and T. indistincta by having a thinner genital field (120–145 in T. caerulea, 150–205 in others). Male T. caerulea can be differentiated from male T. indistincta by having a smaller dorsum (length = 405–460 in T. caerulea, 480–645 in T. indistincta; width = 260–305 in T. caerulea, 315–470 in T. indistincta). Body proportions of male T. caerulea do not differ from male T. malarkeyorum, T. sellersorum, and T. delicatexa, but can be differentiated by dorsal coloration.

Description

Female (Figure 36) (n = 5) (holotype measurements in parentheses when available) with characters of the genus with following specifications.

Dorsum — (550–600 (580) long; 400–440 (440) wide) ovoid with faint blue coloration anteriorly and posteriorly, broadly connected medially. Anterio-medial platelets (125–130 (128.75) long; 40–45 (42.5) wide). Anterio-lateral platelets (145–157.5 (145) long; 57.5–65 (62.5) wide) fused to dorsal plate. Dgl-4 much closer to the edge of the dorsum than to the muscle scars (distance between Dgl-4 275–320 (320)). Dorsal plate proportions: dorsum length/width 1.32–1.48 (1.32); dorsal width/distance between Dgl-4 1.25–1.51 (1.38); anterio-medial platelet length/width 2.78–3.25 (3.03); anterio-lateral platelet length/width 2.32–2.48 (2.32); anterio-lateral/anterio-medial length 1.12–1.21 (1.13).

Gnathosoma — Subcapitulum (310–330 (320) long (ventral); 225–245 (240) long (dorsal); 137.5–156.25 (155) tall) colorless. Rostrum (120–135 (125) long; 45–47.5 (45) wide). Chelicerae (310–330 (320) long) with curved fangs (62–70 (65) long). Subcapitular proportions: ventral length/height 2.06–2.25 (2.06); rostrum length/width 2.67–2.84 (2.78). Pedipalps with tuberculate ventral extensions on femora and genua. Palpomeres: trochanter (40–48.75 (48.75) long); femur (115–120 (120) long); genu (67.5–72.5 (72.5) long); tibia (85–87.5 (87.5) long; 22.5–25 (25) wide); tarsus (17.5–20 (20) long). Palpomere proportions: femur/genu 1.66–1.78 (1.66); tibia/femur 0.71–0.76 (0.73); tibia length/width 3.50–3.83 (3.50).

Venter — (620–750 (660) long; 400–580 (660) wide) with faint blue coloration. Gnathosomal bay (145–175 (145) long; 82.5–116.25 (95) wide). Cxgl-4 subapical. Medial suture absent. Genital plates (155–165 (155) long; 140–145 (145) wide). Additional measurements: Cx-1 (257.5–305 (257.5) long (total); 125–135 (125) long (medial)); Cx-3 (310–380 (380) wide); anterior venter (130–150 (137.5) long). Ventral proportions: gnathosomal bay length/width 1.40–1.94 (1.53); anterior venter/genital field length 0.82–0.97 (0.89); anterior venter length/genital field width 0.90–1.04 (0.95).

Male (Figure 36) (n = 3) (allotypic measurements in parentheses when available) with characters of the genus with following specifications.

Dorsum — (405–460 (460) long; 260–305 (300) wide) ovoid with faint blue coloration anteriorly and posteriorly, broadly connected medially. Anterio-medial platelets (95–106.25 (106.25) long; 35–37.5 (35) wide). Anterio-lateral platelets (117.5–130 (125) long; 40–47.5 (45) wide) fused to dorsal plate. Dgl-4 much closer to the edge of the dorsum than to the muscle scars (distance between Dgl-4 190–240 (230)). Dorsal plate proportions: dorsum length/width 1.51–1.56 (1.53); dorsal width/distance between Dgl-4 1.27–1.37 (1.30); anterio-medial platelet length/width 2.53–3.04 (3.04); anterio-lateral platelet length/width 2.74–2.94 (2.78); anterio-lateral/anterio-medial length 1.18–1.24 (1.18).

Gnathosoma — Subcapitulum (220–242.5 (227.5) long (ventral); 165–185 (165) long (dorsal); 87.5–97.5 (97.5) tall) colorless. Rostrum (88.75–92.5 (92.5) long; 30–35 (33.75) wide). Chelicerae (225–232.5 (227.5) long) with curved fangs (42.5–50 (45) long). Subcapitular proportions: ventral length/height 2.33–2.51 (2.33); rostrum length/width 2.64–2.96 (2.74). Pedipalps with tuberculate ventral extensions on femora and genua. Palpomeres: trochanter (37.5–37.5 (37.5) long); femur (83.75–87.5 (87.5) long); genu (52.5–55 (55) long); tibia (67.5–70 (67.5) long; 20–22.5 (20) wide); tarsus (15–20 (17.5) long). Palpomere proportions: femur/genu 1.59–1.60 (1.59); tibia/femur 0.77–0.81 (0.77); tibia length/width 3.11–3.38 (3.38).

Venter — (485–550 (550) long; 305–340 (330) wide) with faint blue coloration. Gnathosomal bay (120–127.5 (125) long; 57.5–65 (60) wide). Cxgl-4 subapical. Medial suture (65-75 (75)). Genital plates (110–117.5 (117.5) long; 90–105 (102.5) wide). Additional measurements: Cx-1 (220–220 (220) long (total); 85–95 (95) long (medial)); Cx-3 (240–260 (260) wide); anterior venter (165–180 (180) long). Ventral proportions: gnathosomal bay length/width 1.96–2.09 (2.08); anterior venter/genital field length 1.50–1.57 (1.53); anterior venter length/genital field width 1.71–1.83 (1.76).

Immatures unknown.

Etymology

Specific epithet (caerulea) refers to the overall and diagnostic bluish appearance of this species (caeruleus, L. sky-blue).

Distribution

Known only from Wayne County, Tennessee (Figure 35). T. caerulea has been collected so rarely that comments about distribution are speculative, but given our collection efforts, it is reasonable to speculate that this species is at least restricted to the southern Appalachians.

Figure 35. 

Torrenticola caerulea sp. n. distribution.

Figure 36. 

Torrenticola caerulea sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 37. 

Torrenticola caerulea sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Remarks

Torrenticola caerulea groups with other members of the Rusetria Complex with high support and specimens of this species were less than 1% different in COI sequence from each other. In all analyses, T. caerulea groups with two other morphologically similar species: T. biscutella and T. malarkeyorum. These three species are 3–5% different from each other in COI sequence. The three of these species are morphologically similar to the more distantly-related T. delicatexa, but T. caerulea can be differentiated from all of these by color. The range of T. caerulea overlaps with each of these except for T. biscutella, which is not known from east of the Mississippi River.

This species hypothesis is supported by biogeography, low COI divergence within the species (0–2%) and high divergence between species (3–15%), and by the morphological characters outlined in the diagnosis.

Torrenticola cardia Fisher & Dowling, sp. n.

Material examined

HOLOTYPE (♀): from USA, New York, Greene County, beside Rt. 23A, 9.6 km west of Rt. 296 (Hunter), (42°14'14"N, 74°19'19"W), 22 June 1990, by IM Smith, IMS900052

PARATYPES (9 ♀; 8 ♂): New York, USA: 3 ♀ and 3 ♂ from Cayuga County, beside Route 38A at Niles, (42°50'50"N, 76°25'25"W), 22 July 1990, by IM Smith, IMS900113A • (allotype) 1 ♂ from Greene, beside Rt. 23A, 9.6 km west of Rt. 296 (Hunter), (42°14'14"N, 74°19'19"W), 22 June 1990, by IM Smith, IMS900052 • 2 ♀ and 1 ♂ from Greene County, beside Rt. 23A, 9.6 km west of Rt. 296 (Hunter), (42°14'14"N, 74°19'19"W), 22 June 1990, by IM Smith, IMS900052 • 1 ♀ from Schuyler County, beside Town Line Road off Route 228, 0.6 km south of Perry City, (42°29'29"N, 76°42'42"W), 21 July 1990, by IM Smith, IMS900112A • Ohio, USA: 1 ♀ and 1 ♂ from Hocking County, beside road near Ash Cave, (39°24'24"N, 82°33'33"W), 5 May 1993, by IM Smith, DR Cook, IMS930001A • Virginia, USA: 2 ♀ and 2 ♂ from Bath County, beside Rt. 687, 2.4 km south of Bacova, (38°2'2"N, 79°51'51"W), 15 July 1990, by IM Smith, IMS900097

Type deposition

Holotype (♀), allotype (♂), and some paratypes (5 ♀; 4 ♂) deposited in the CNC; other paratypes (4 ♀; 3 ♂) deposited in the ACUA.

Diagnosis

Torrenticola cardia are similar to other members of the Tricolor Complex (T. bittikoferae, T. dimorpha, T. hoosieri, T. kringi, T. larvata, T. mohawk, T. pearsoni, T. olliei, T. sierrensis, T. tricolor, T. trimaculata, and T. unimaculata,) in having a short, conical rostrum. T. cardia can be differentiated from most Torrenticola, including other members of the Tricolor Complex, by having a distinct dorsal pattern. T. cardia are most similar to other members of the Tricolor Complex that have bold patterning (T. larvata, T. tricolor, T. unimaculata, T. trimaculata, T. kringi, and T. mohawk). T. cardia can be differentiated from T. tricolor, T. trimaculata, T. kringi, and T. mohawk by having a more ovoid dorsum (length/width ♀ = 1.39–1.47 in T. cardia, 1.15–1.35 in others; ♂ = 1.43–1.54 in T. cardia, 1.19–1.39 in others). T. cardia can be differentiated from T. unimaculata by dorsal pattern. Female T. cardia can be differentiated from female T. larvata by having a shorter subcapitulum (♀ = 265–273 in T. cardia, 275–288 in T. larvata) and a larger genital field (length ♀ = 190–198 in T. cardia, 182–188 in T. larvata; width ♀ = 160–175 in T. cardia, 145–153 in T. larvata). Male T. cardia can be differentiated from male T. larvata by having less elongate pedipalpal tibiae (length/width ♂ = 2.82–3.05 in T. cardia, 3.10–3.20 in T. larvata) and a larger dorsum (length ♂ = 625–670 in T. cardia, 550–610 in T. larvata; width ♂ = 405–445 in T. cardia, 350–400 in T. larvata).

Description

Female (Figure 39) (n = 5) (holotype measurements in parentheses when available) with characters of the genus with following specifications.

Dorsum — (710–785 (750) long; 510–555 (510) wide) ellipsoid with reddish-purple, bluish-purple or bright orange spot medially extending in a strip anteriorly often to the anterior-medial platelets. Anterio-medial platelets (130–140 (140) long; 70–75 (75) wide). Anterio-lateral platelets (195–207.5 (197.5) long; 70–80 (80) wide) free from dorsal plate. Dgl-4 much closer to the edge of the dorsum than to the muscle scars (distance between Dgl-4 390–410 (395)). Dorsal plate proportions: dorsum length/width 1.39–1.47 (1.47); dorsal width/distance between Dgl-4 1.29–1.35 (1.29); anterio-medial platelet length/width 1.86–2.00 (1.87); anterio-lateral platelet length/width X=2.47–2.93 (2.47); anterio-lateral/anterio-medial length 1.39–1.50 (1.41).

Gnathosoma — Subcapitulum (265–272.5 (272.5) long (ventral); 180–190 (190) long (dorsal); 125–132.5 (130) tall) with reddish-purple or bluish purple coloration. Rostrum (90–100 (100) long; 40–42.5 (40) wide) short and conical. Chelicerae (250–265 (260) long) with curved fangs (60–60 (60) long). Subcapitular proportions: ventral length/height 2.04–2.12 (2.10); rostrum length/width 2.24–2.50 (2.50). Pedipalps with stocky, tuberculate ventral extensions on femora and genua. Palpomeres: trochanter (38.75–42.5 41.25) long); femur (97.5–107.5 (102.5) long); genu (65–70 (67.5) long); tibia (80–91.25 (80) long; 26.25–27.5 (27.5) wide); tarsus (22.5–25 (22.5) long). Palpomere proportions: femur/genu 1.50–1.56 (1.52); tibia/femur 0.78–0.87 (0.78); tibia length/width 2.91–3.32 (2.91).

Venter — (830–925 (850) long; 565–600 (565) wide) with reddish-purple or bluish-purple coloration restricted to the edges of the gnathosomal bay, coxal plates, and genital plates. Gnathosomal bay (150–162.5 (155) long; 72.5–80 (72.5) wide). Cxgl-4 subapical. Medial suture (25–50 (50) long). Genital plates (190–197.5 (190) long; 160–175 (162.5) wide). Additional measurements: Cx-1 (290–310 (290) long (total); 135–165 (135) long (medial)); Cx-3 (355–380 (355) wide); anterior venter (185–212.5 (205) long). Ventral proportions: gnathosomal bay length/width 1.94–2.17 (2.14); anterior venter/genital field length 0.94–1.09 (1.08); anterior venter length/genital field width 1.12–1.26 (1.26); anterior venter/medial suture 4.10–7.60 (4.10).

Male (Figure 40) (n = 5) (allotypic measurements in parentheses when available) with characters of the genus with following specifications.

Dorsum — (625–670 (670) long; 405–445 (445) wide) ellipsoid with reddish-purple, bluish-purple or bright orange spot medially extending in a strip anteriorly often to the anterior-medial platelets. Anterio-medial platelets (120–130 (125) long; 65–72.5 (70) wide). Anterio-lateral platelets (170–197.5 (197.5) long; 75–80 (75) wide) free from dorsal plate. Dgl-4 much closer to the edge of the dorsum than to the muscle scars (distance between Dgl-4 325–380 (380)). Dorsal plate proportions: dorsum length/width 1.43–1.54 (1.51); dorsal width/distance between Dgl-4 1.17–1.28 (1.17); anterio-medial platelet length/width 1.72–2.00 (1.79); anterio-lateral platelet length/width 2.25–2.63 (2.63); anterio-lateral/anterio-medial length 1.35–1.58 (1.58).

Gnathosoma — Subcapitulum (227.5–250 (250) long (ventral); 165–180 (170) long (dorsal); 90–112.5 (110) tall) with reddish-purple or bluish purple coloration. Rostrum (77.5–92.5 (90) long; 32.5–37.5 (37.5) wide) short and conical. Chelicerae (212.5–230 (225) long) with curved fangs (50–55 (50) long). Subcapitular proportions: ventral length/height 2.17–2.56 (2.27); rostrum length/width 2.27–2.47 (2.40). Pedipalps with stocky, tuberculate ventral extensions on femora and genua. Palpomeres: trochanter (36.25–38.75 (36.25) long); femur (87.5–97.5 (92.5) long); genu (60–67.5 (62.5) long); tibia (73.75–82.5 (77.5) long; 25–27.5 (27.5) wide); tarsus (20–25 (22.5) long). Palpomere proportions: femur/genu 1.38–1.48 (1.48); tibia/femur 0.82–0.90 (0.84); tibia length/width 2.82–3.05 (2.82).

Venter — (710–780 (780) long; 460–495 (490) wide) with reddish-purple or bluish-purple coloration restricted to the edges of the gnathosomal bay, coxal plates, and genital plates. Gnathosomal bay (125–137.5 (135) long; 67.5–72.5 (70) wide). Cxgl-4 subapical. Medial suture (107.5–130 (125) long). Genital plates (150–175 (175) long; 97.5–115 (105) wide). Additional measurements: Cx-1 (265–280 (280) long (total); 140–150 (145) long (medial)); Cx-3 (325–355 (350) wide); anterior venter (265–290 (290) long). Ventral proportions: gnathosomal bay length/width 1.85–1.93 (1.93); anterior venter/genital field length 1.66–1.81 (1.66); anterior venter length/genital field width 2.52–2.77 (2.76); anterior venter/medial suture 2.21–2.51 (2.32).

Immatures unknown.

Etymology

Specific epithet (cardia) refers to the dorsal coloration, which is either a heart-shaped or resembles a bleeding heart (kardiá, G. heart).

Distribution

Appalachians (Figure 38).

Figure 38. 

Torrenticola cardia sp. n. distribution.

Figure 39. 

Torrenticola cardia sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 40. 

Torrenticola cardia sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Remarks

Unfortunately, we were unable to acquire fresh material of Torrenticola cardia and therefore this species is not included in our phylogenetic analyses. However, we were able to examine morphology with material preserved in GAW. The overall appearance, short conical rostrum that is downturned in the male, and distribution, are consistent with placing this species in the Tricolor Complex and Tricolor Identification Group.

Torrenticola copipalpa Fisher & Dowling, sp. n.

Material examined

HOLOTYPE (♀): from USA, Oregon, Lane County, Gate Creek (44°8'48"N, 122°34'20"W), 11 Aug 2013, by JC O’Neill, & WA Nelson, JNOW 13-0811-001.

PARATYPES (20 ♀; 26 ♂): California, USA: 1 ♀ from Alpine County, Markleeville Creek (38°41'39"N, 119°46'41"W), 30 Aug 2013, by JR Fisher, JRF 13-0830-001 • 1 ♀ from Del Norte County, Six Rivers National Forest, Middle Fork Smith River (41°51'20"N, 123°53'10"W), 15 Aug 2013, by JR Fisher, JRF 13-0815-002 • 5 ♂ from El Dorado County, El Dorado National Forest, Taylor Creek (38°55'59"N, 120°3'21"W), 27 Aug 2013, by JR Fisher, JRF 13-0827-003 • 1 ♀ from Mendocino County, Cottaneva Creek, beside Route 1, 21.8 kilometers southwest of Route 101, 5 Aug 1987, by IM Smith, IMS870129A • 2 ♂ from Trinity County, small cascading trickle beside Route 36, 5.2 kilometers west of Forest Glen Station, 6 Aug 1987, by IM Smith, IMS870132 • 1 ♂ from Trinity County, South Fork of Trinity River, beside Route 36 at Forest Glen campground, 6 Aug 1987, by IM Smith, IMS870131 • Oregon, USA: 2 ♂ from Coos County, Gaylord, Coquille Myrtle Grove State Park, Coquille River, 2 Jul 1983, by IM Smith, IMS830014 • 3 ♀ and 3 ♂ from Coos County, Siskiyou National Forest, Road 33 between Powers & Agness, Coal Creek, 2 Jul 1983, by IM Smith, IMS830015 • 2 ♂ from Coos County, Siskiyou National Forest, Road 33 between Powers & Agness, Daphne Grove campground, Coquille River, 2 Jul 1983, by IM Smith, IMS830016 • 1 ♂ from Coos County, Siskiyou National Forest, Road 33 between Powers & Agness, Daphne Grove campground, 2 Jul 1983, by IM Smith, IMS830017 • 1 ♀ and 1 ♂ from Curry County, Port Orford, Butler Bar campground, Elk River, 25 Jun 1976, by IM Smith, IMS760162 • 1 ♀ from Curry County, Port Orford, Butler Bar campground, Elk River, 25-26 Jun 1976, by IM Smith, IMS760163 • 2 ♂ from Curry County, Port Orford, small spring run beside road from Humbug Mountain State Park to McGribble campground, 25 Jun 1976, by IM Smith, IMS760161 • 3 ♂ from Curry County, Port Orford, Humbug Mountain State Park Picnic Area, beside Route 1, Brush Creek, 1 Jul 1983, by IM Smith, IMS830012 • 1 ♀ from Curry County, Port Orford, Humbug Mountain State Park Picnic Area, beside Route 1, Brush Creek, 3 Jul 1983, by IM Smith, IMS830020A • 2 ♀ from Curry County, Quosatana Creek (42°29'21"N, 124°14'2"W), 14 Aug 2013, JR Fisher, JRF 13-0814-003 • 1 ♀ from Curry County, Rogue River National Forest, Elk River (42°42'46"N, 124°18'41"W), 13 Aug 2013, by JR Fisher, JRF 13-0813-003 • 4 ♀ and 3 ♂ from Curry County, Sixes, Sixes River, beside road at mouth of Edson Creek, 4 Jul 1983, by IM Smith, IMS830021A • 1 ♂ (ALLOTYPE) from Lane County, Gate Creek (44°8'48"N, 122°34'20"W), 11 Aug 2013, by JC O’Neill, & WA Nelson, JNOW 13-0811-001 • 4 ♀ from Lane County, Gate Creek (44°8'48"N, 122°34'20"W), 11 Aug 2013, by JC O’Neill, & WA Nelson, JNOW 13-0811-001.

Type deposition

Holotype (♀), allotype (♂), and other paratypes (15 ♀; 20 ♂) deposited in the CNC; other paratypes (5 ♀; 5 ♂) deposited in ACUA.

Diagnosis

Torrenticola copipalpa are similar to members of the Miniforma group (T. manni, T. miniforma, T. oliveri, T. pacificensis, T. pinocchio, and T. rockyensis) in having short, stocky pedipalps (except T. oliveri and T. pinocchio); similar pedipalpal extensions (unique to members of this group); and being among the smallest Torrenticola in the west (dorsum 500–625 long) (except T. oliveri). T. copipalpa are best differentiated from all other Miniforma group (except T. pinocchio) by having broad, flat pedipalp femoral tubercles (conical/tuberculate in all others). T. copipalpa can be differentiated from T. pinocchio by having a less elongate rostrum (length/width = 2.5–3.0 in T. copipalpa, 4.5–4.9 in T. pinocchio) and less elongate pedipalpal tibiae (length/width = 2.4–2.9 in T. copipalpa, 3.1–3.5 in T. pinocchio).

Description

Female (Figure 42) (n = 6) (holotype measurements in parentheses when available) with characters of the genus with following specifications.

Dorsum — (555–605 (605) long; 380–420 (420) wide) ovoid and usually colorless, occasionally with faint purple coloration without distinct pattern. Anterio-medial platelets (115–127.5 (127.5) long; 47.5–57.5 (57.5) wide). Anterio-lateral platelets (162.5–180 (180) long; 53.75–62.5 (62.5) wide) free from dorsal plate. Dgl-4 much closer to the edge of the dorsum than to the muscle scars (distance between Dgl-4 290–335 (335)). Dorsal plate proportions: dorsum length/width 1.39–1.47 (1.44); dorsal width/distance between Dgl-4 1.25–1.32 (1.25); anterio-medial platelet length/width 2.13–2.42 (2.22); anterio-lateral platelet length/width 2.87–3.04 (2.88); anterio-lateral/anterio-medial length 1.35–1.46 (1.41).

Gnathosoma — Subcapitulum (312.5–337.5 (337.5) long (ventral); 228–257.5 (257.5) long (dorsal); 117.5–130 (125) tall) colorless. Rostrum (122.5–135 (130) long; 42.5–47.5 (45) wide). Chelicerae (313–341 (340) long) with curved fangs (50–59 (55) long). Subcapitular proportions: ventral length/height 2.58–2.70 (2.70); rostrum length/width 2.72–2.94 (2.89). Pedipalps short and stocky (especially tibiae) with broad, dentate, and anteriorly-directed ventral extensions on femora and dentate, flanged ventral extensions on genua. Palpomeres: trochanter (30–35 (35) long); femur (90–100 (97.5) long); genu (62.5–67.5 (67.5) long); tibia (52.5–58.75 (57.5) long; 20–22.5 (21.25) wide); tarsus (15–17.5 (15) long). Palpomere proportions: femur/genu 1.44–1.51 (1.41); tibia/femur 0.55–0.61 (0.59); tibia length/width 2.59–2.71 (2.71).

Venter — (690–760 (760) long; 438–520 (520) wide) colorless. Gnathosomal bay (136.25–152.5 (152.5) long; 75–82.5 (80) wide). Cxgl-4 subapical. Medial suture (40–45 (45) long). Genital plates (152.5–165 (165) long; 137.5–160 (160) wide). Additional measurements: Cx-1 (261–290 (290) long (total); 108–155 (155) long (medial)); Cx-3 (268–320 (320) wide); anterior venter (187.5–210 (210) long). Ventral proportions: gnathosomal bay length/width 1.65–1.97 (1.91); anterior venter/genital field length 1.19–1.28 (1.27); anterior venter length/genital field width 1.31–1.40 (1.31); anterior venter/medial suture 4.53–4.88 (4.67).

Male (Figure 43) (n = 6) (allotypic measurements in parentheses when available) with characters of the genus with following specifications.

Dorsum — (500–570 (520) long; 355–390 (360) wide) ovoid and usually colorless, occasionally with faint purple coloration without distinct pattern. Anterio-medial platelets (105–117.5 (105) long; 45–56.25 (45) wide). Anterio-lateral platelets (155–167.5 (155) long; 50–60 (50) wide) free from dorsal plate. Dgl-4 much closer to the edge of the dorsum than to the muscle scars (distance between Dgl-4 285–315 (290)). Dorsal plate proportions: dorsum length/width 1.39–1.54 (1.44); dorsal width/distance between Dgl-4 1.18–1.37 (1.24); anterio-medial platelet length/width 2.09–2.37 (2.33); anterio-lateral platelet length/width 2.79–3.10 (3.10); anterio-lateral/anterio-medial length 1.32–1.48 (1.48).

Gnathosoma — Subcapitulum (280–307.5 (295) long (ventral); 215–253 (220) long (dorsal); 105–115 (105) tall) colorless. Rostrum (110–120 (115) long; 40–46.25 (40) wide). Chelicerae (280–328 (295) long) with curved fangs (45–65 (55) long). Subcapitular proportions: ventral length/height 2.67–2.81 (2.81); rostrum length/width 2.54–2.88 (2.88). Pedipalps short and stocky (especially tibiae) with broad, dentate, and anteriorly-directed ventral extensions on femora and dentate, flanged ventral extensions on genua. Palpomeres: trochanter (32.5–32.5 (32.5) long); femur (82.5–92.5 (87.5) long); genu (57.5–65 (60) long); tibia (52.5–57.5 (52.5) long; 18.75–21.25 (21.25) wide); tarsus (15–17.5 (15) long). Palpomere proportions: femur/genu 1.35–1.54 (1.46); tibia/femur 0.60–0.64 (0.60); tibia length/width 2.47–2.88 (2.47).

Venter — (610–700 (670) long; 420–496 (440) wide) colorless. Gnathosomal bay (110–140 (130) long; 67.5–75 (75) wide). Cxgl-4 subapical. Medial suture (77.5–97.5 (77.5) long). Genital plates (130–137.5 (130) long; 100–112.5 (105) wide). Additional measurements: Cx-1 (235–263 (245) long (total); 115–130 (130) long (medial)); Cx-3 (270–300 (280) wide); anterior venter (210–232.5 (217.5) long). Ventral proportions: gnathosomal bay length/width 1.63–1.87 (1.73); anterior venter/genital field length 1.62–1.77 (1.67); anterior venter length/genital field width 2.04–2.21 (2.07); anterior venter/medial suture 2.36–2.88 (2.81).

Immatures unknown.

Etymology

Specific epithet (copipalpa) refers to the blade-like pedipalp femoral tubercles (copis, L. small knife; palpus, L. hand, feeler), which distinguish them from similar, co-occurring species.

Distribution

Northern California and western Oregon (Figure 41).

Figure 41. 

Torrenticola copipalpa sp. n. distribution.

Figure 42. 

Torrenticola copipalpa sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 43. 

Torrenticola copipalpa sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Remarks

Torrenticola copipalpa groups with other members of the Miniforma Complex with high support and specimens of this species are less than 1% different in COI sequence from each other. In all analyses, T. copipalpa groups with three other morphologically similar species: T. pacificensis, T. manni, and T. rockyensis. These three species are greater than 4% different from each other. This species overlaps with T. miniforma in California and with T. pacificensis in west-central Oregon.

Based upon overall similarity, the pedipalp genu extensions, and western distribution, we were able to place this species in the Miniforma Identification Group.

This species hypothesis is supported by biogeography, low COI divergence within the species (0–2%) and high divergence between species (3–15%), and by the morphological characters outlined in the diagnosis.

Torrenticola daemon Fisher & Dowling, sp. n.

Material examined

HOLOTYPE (♀): from USA, Alabama, Clay County, beside Forest Route 649, 0.8 km northeast of road from Campbell Springs to Forest Route 600, (33°22'22"N, 85°52'52"W), 3 July 1990, by IM Smith, IMS900075A.

PARATYPES (4 ♀; 4 ♂): Alabama, USA: 1 ♂ (ALLOTYPE) from Clay County, beside Forest Route 649, 0.8 km northeast of road from Campbell Springs to Forest Route 600, (33°22'22"N, 85°52'52"W), 3 July 1990, by IM Smith, IMS900075A • 4 ♀ and 3 ♂ from Clay County, beside Forest Route 649, 0.8 km northeast of road from Campbell Springs to Forest Route 600, (33°22'22"N, 85°52'52"W), 3 July 1990, by IM Smith, IMS900075A

Type deposition

Holotype (♀), allotype (♂), and some paratypes (2 ♀; 2 ♂) deposited in the CNC; other paratypes (2 ♀; 1 ♂) deposited in the ACUA.

Diagnosis

Torrenticola daemon are similar to other members of the Raptor Group (T. danielleae, T. elusiva, T. gnoma, T. irapalpa, T. ivyae, T. longitibia, T. mjolniri, T. racupalpa, and T. raptor) in having round bodies; Dgl-4 close to muscles scars; long, thin subcapitular rostra; and long, thin pedipalp tibiae. T. daemon can be differentiated from all other Raptor Group by having Dgl-4 closer to the edge of the dorsum (dorsum width/distance between Dgl-4 ♀ = 1.59–1.67 in T. daemon, 1.80–3.29 in others; ♂ = 1.45–1.65 in T. daemon, 1.66–2.73 in others), except T. irapalpa (♀ = 1.81–2.09, ♂ = 1.58–1.86) and T. danielleae (♀ = 1.57–1.70, ♂ = 1.42–1.52). T. daemon can be differentiated from T. longitibia, T. mjolniri, T. elusiva, T. racupalpa, T. raptor, T. danielleae, and T. ivyae by having a less elongate rostrum (length/width = 2.91–3.31 in T. daemon, 3.43–4.40 in others). Female T. daemon can be differentiated from female T. irapalpa by having Dgl-4 closer to the dorsal edge (dorsal width/distance between Dgl-4 ♀ = 1.59–1.67 in T. daemon, 1.81–2.09 in T. irapalpa) and a more elongate gnathosomal bay (length/width ♀ = 1.95–2.42 in T. daemon, 1.35–1.86 in T. irapalpa). Additionally, T. daemon can be differentiated from T. irapalpa by dorsal coloration and pattern.

Description

Female (Figure 45) (n = 5) (holotype measurements in parentheses when available) with characters of the genus with following specifications.

Dorsum — (590–655 (640) long; 460–500 (490) wide) circular with faint reddish-purple coloration separated into anterior and posterior portions, with bright reddish-purple coloration on the anterior-medial platelets, occasionally extending onto the dorsal plate. Anterio-medial platelets (127.5–147.5 (145) long; 57.5–67.5 (65) wide). Anterio-lateral platelets (182.5–210 (210) long; 72.5–82.5 (82.5) wide) free from dorsal plate. Dgl-4 approximately halfway between the edge of the dorsum and the muscle scars (distance between Dgl-4 285–300 (300)). Dorsal plate proportions: dorsum length/width 1.26–1.34 (1.31); dorsal width/distance between Dgl-4 1.59–1.67 (1.63); anterio-medial platelet length/width 2.04–2.28 (2.23); anterio-lateral platelet length/width 2.28–2.66 (2.55); anterio-lateral/anterio-medial length 1.29–1.51 (1.45).

Gnathosoma — Subcapitulum (370–395 (395) long (ventral); 285–300 (300) long (dorsal); 145–157.5 (157.5) tall) colorless, occasionally with faint reddish-purple coloration. Rostrum (155–165 (165) long; 50–55 (55) wide). Chelicerae (375–390 (390) long) with curved fangs (65–75 (75) long). Subcapitular proportions: ventral length/height 2.39–2.57 (2.51); rostrum length/width 2.91–3.20 (3.00). Pedipalps with tuberculate ventral extensions on femora and genua. Palpomeres: trochanter (50–52.5 (50) long); femur (137.5–146.25 (145) long); genu (75–80 (80) long); tibia (97.5–105 (105) long; 22.5–25 (25) wide); tarsus (20–22.5 (20) long). Palpomere proportions: femur/genu 1.81–1.90 (1.81); tibia/femur 0.67–0.72 (0.72); tibia length/width 4.05–4.33 (4.20).

Venter — (710–800 (800) long; 515–570 (545) wide) colorless. Gnathosomal bay (182.5–200 (190) long; 80–97.5 (97.5) wide). Cxgl-4 subapical. Medial suture (10–15 (15) long). Genital plates (160–175 (175) long; 145–150 (150) wide). Additional measurements: Cx-1 (310–345 (345) long (total); 125–155 (155) long (medial)); Cx-3 (350–375 (375) wide); anterior venter (155–185 (185) long). Ventral proportions: gnathosomal bay length/width 1.95–2.42 (1.95); anterior venter/genital field length 0.97–1.09 (1.06); anterior venter length/genital field width 1.07–1.23 (1.23); anterior venter/medial suture 10.33–17.50 (12.33).

Male (Figure 46) (n = 4) (allotypic measurements in parentheses when available) with characters of the genus with following specifications.

Dorsum — (545–570 (570) long; 410–425 (425) wide) circular with faint reddish-purple coloration separated into anterior and posterior portions. Anterio-medial platelets (115–125 (122.5) long; 52.5–58.75 (58.75) wide). Anterio-lateral platelets (175–188.75 (188.75) long; 67.5–75 (75) wide) free from dorsal plate. Dgl-4 approximately halfway between the edge of the dorsum and the muscle scars (distance between Dgl-4 255–290 (275)). Dorsal plate proportions: dorsum length/width 1.31–1.34 (1.34); dorsal width/distance between Dgl-4 1.45–1.65 (1.55); anterio-medial platelet length/width 2.09–2.22 (2.09); anterio-lateral platelet length/width 2.50–2.78 (2.52); anterio-lateral/anterio-medial length 1.40–1.63 (1.54).

Gnathosoma — Subcapitulum (320–335 (335) long (ventral); 247.5–255 (255) long (dorsal); 120–122.5 (120) tall) colorless, occasionally with faint reddish-purple coloration. Rostrum (132.5–137.5 (137.5) long; 40–45 (45) wide). Chelicerae (310–330 (330) long) with curved fangs (60–65 (60) long). Subcapitular proportions: ventral length/height 2.67–2.79 (2.79); rostrum length/width 3.06–3.31 (3.06). Pedipalps with tuberculate ventral extensions on femora and genua. Palpomeres: trochanter (42.5–46.25 (45) long); femur (117.5–122.5 (120) long); genu (67.5–70 (67.5) long); tibia (90–92.5 (91.25) long; 21.25–22.5 (22.5) wide); tarsus (20–20 (20) long). Palpomere proportions: femur/genu 1.74–1.78 (1.78); tibia/femur 0.76–0.77 (0.76); tibia length/width 4.00–4.35 (4.06).

Venter — (655–705 (705) long; 460–470 (470) wide) colorless. Gnathosomal bay (165–165 (165) long; 70–80 (75) wide). Cxgl-4 subapical. Medial suture (50–55 (55) long). Genital plates (145–155 (155) long; 115–120 (120) wide). Additional measurements: Cx-1 (290–315 (290) long (total); 130–150 (130) long (medial)); Cx-3 (320–340 (340) wide); anterior venter (210–225 (225) long). Ventral proportions: gnathosomal bay length/width 2.06–2.36 (2.20); anterior venter/genital field length 1.42–1.55 (1.45); anterior venter length/genital field width 1.75–1.91 (1.88); anterior venter/medial suture 4.00–4.20 (4.09).

Immatures unknown.

Etymology

Specific epithet (daemon) refers to the diagnostic red coloration on the anterio-medial platelets, which resemble the red eyes of an evil demon (daemon, L. originally benevolent or benign nature spirits, but were characterized as dangerous or evil by the writings of Plato and later used in Christian literature, popularizing the idea of demons as evil; noun in apposition).

Distribution

Alabama (Figure 44).

Figure 44. 

Torrenticola daemon sp. n. distribution.

Figure 45. 

Torrenticola daemon sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 46. 

Torrenticola daemon sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Remarks

Unfortunately, we were unable to acquire fresh material of Torrenticola daemon and therefore this species is not included in our phylogenetic analyses. However, we were able to examine morphology with material preserved in GAW. The overall appearance, elongate subcapitular rostra, elongate pedipalpal tibiae, and Dgl-4 close to the muscle scars, are consistent with placing this species in the Raptor Complex and Raptor Identification Group.

Torrenticola danielleae Fisher & Dowling, sp. n.

Material examined

HOLOTYPE (♀): from USA, Georgia, Floyd County, The Pocket Campground beside road from Everett Springs to Villanow, (34°35'35"N, 85°5'5"W), 2 July 1990, by IM Smith, IMS900073A.

PARATYPES (4 ♀; 5 ♂): Georgia, USA: 2 ♀ and 1 ♂ from Chattooga County, beside road from Everett Springs to Villanow 1.8 km south of The Pocket Recreation Area, (34°34'34"N, 80°5'5"W), 4 July 1990, by IM Smith, IMS900076 • 1 ♂ (ALLOTYPE) from Floyd County, The Pocket Campground beside road from Everett Springs to Villanow, (34°35'35"N, 85°5'5"W), 2 July 1990, by IM Smith, IMS900073A • 2 ♀ and 3 ♂ from Floyd County, The Pocket Campground beside road from Everett Springs to Villanow, (34°35'35"N, 85°5'5"W), 2 July 1990, by IM Smith, IMS900073A

Type deposition

Holotype (♀), allotype (♂), and some paratypes (2 ♀; 2 ♂) deposited in the CNC; other paratypes (2 ♀; 2 ♂) deposited in the ACUA.

Diagnosis

Torrenticola danielleae are similar to other members of the Raptor Group (T. daemon, T. elusiva, T. gnoma, T. irapalpa, T. ivyae, T. longitibia, T. mjolniri, T. racupalpa, and T. raptor) in having round bodies; Dgl-4 close to muscles scars; long, thin subcapitular rostra; and long, thin pedipalp tibiae. T. danielleae can be differentiated from all other Raptor Group by having Dgl-4 closer to the edge of the dorsum (dorsum width/distance between Dgl-4 ♀ = 1.57–1.70 in T. danielleae, 1.80–3.29 in others; ♂ = 1.42–1.52 in T. danielleae, 1.58–2.73 in others), except T. daemon (♀ = 1.59–1.67 ♂ = 1.45–1.65). T. danielleae can be differentiated from T. daemon by having a more elongate rostrum (length/width = 3.43–3.75 in T. danielleae, 2.91–3.31 in T. daemon) and dorsal pattern.

Description

Female (Figure 48) (n = 5) (holotype measurements in parentheses when available) with characters of the genus with following specifications.

Dorsum — (635–680 (635) long; 510–545 (510) wide) circular with reddish-purple coloration posteriorly extending in a strip anteriorly to the edge of the dorsal plate. Anterio-medial platelets (135–155 (147.5) long; 55–65 (62.5) wide). Anterio-lateral platelets (170–195 (180) long; 70–80 (80) wide) free from dorsal plate. Dgl-4 approximately halfway between the edge of the dorsum and the muscle scars (distance between Dgl-4 320–335 (320)). Dorsal plate proportions: dorsum length/width 1.20–1.30 (1.25); dorsal width/distance between Dgl-4 1.57–1.7 (1.59); anterio-medial platelet length/width 2.25–2.73 (2.36); anterio-lateral platelet length/width 2.25–2.79 (2.25); anterio-lateral/anterio-medial length 1.15–1.44 (1.22).

Gnathosoma — Subcapitulum (340–370 (365) long (ventral); 260–280 (270) long (dorsal); 135–145 (145) tall) colorless. Rostrum (150–160 (155) long; 40–45 (42.5) wide). Chelicerae (350–380 (380) long) with curved fangs (50–62.5 (62.5) long). Subcapitular proportions: ventral length/height 2.52–2.65 (2.52); rostrum length/width 3.44–3.75 (3.65). Pedipalps with long, tuberculate ventral extensions on femora and genua. Palpomeres: trochanter (40–45 (42.5) long); femur (127.5–141.25 (137.5) long); genu (67.5–75 (75) long); tibia (92.5–107.5 (102.5) long; 20–22.5 (21.25) wide); tarsus (17.5–20 (20) long). Palpomere proportions: femur/genu 1.77–1.89 (1.83); tibia/femur 0.73–0.76 (0.75); tibia length/width 4.63–5.00 (4.82).

Venter — (780–835 (780) long; 560–610 (560) wide) with reddish-purple coloration, occasionally faint. Gnathosomal bay (170–185 (185) long; 75–85 (80) wide). Cxgl-4 subapical. Medial suture (25–27.5 (27.5) long). Genital plates (165–185 (185) long; 155–160 (160) wide). Additional measurements: Cx-1 (300–340 (315) long (total); 135–155 (135) long (medial)); Cx-3 (345–360 (355) wide); anterior venter (170–195 (185) long). Ventral proportions: gnathosomal bay length/width 2.00–2.31 (2.31); anterior venter/genital field length 1.00–1.12 (1.00); anterior venter length/genital field width 1.10–1.24 (1.16); anterior venter/medial suture 6.73–7.80 (6.73).

Male (Figure 49) (n = 5) (allotypic measurements in parentheses when available) with characters of the genus with following specifications.

Dorsum — (490–530 (510) long; 370–385 (370) wide) circular with reddish-purple coloration posteriorly extending in a strip anteriorly to the edge of the dorsal plate. Anterio-medial platelets (115–125 (120) long; 45–52.5 (52.5) wide). Anterio-lateral platelets (150–175 (150) long; 57.5–65 (65) wide) free from dorsal plate. Dgl-4 approximately halfway between the edge of the dorsum and the muscle scars (distance between Dgl-4 250–265 (260)). Dorsal plate proportions: dorsum length/width 1.27–1.39 (1.38); dorsal width/distance between Dgl-4 1.42–1.52 (1.42); anterio-medial platelet length/width 2.29–2.72 (2.29); anterio-lateral platelet length/width 2.31–2.92 (2.31); anterio-lateral/anterio-medial length 1.25–1.43 (1.25).

Gnathosoma — Subcapitulum (285–295 (290) long (ventral); 210–220 (215) long (dorsal); 100–105 (105) tall) colorless. Rostrum (120–125 (120) long; 35–35 (35) wide). Chelicerae (270–300 (280) long) with curved fangs (40–50 (50) long). Subcapitular proportions: ventral length/height 2.74–2.85 (2.76); rostrum length/width 3.43–3.57 (3.43). Pedipalps with long, tuberculate ventral extensions on femora and genua. Palpomeres: trochanter (33.75–37.5 (36.25) long); femur (100–110 (107.5) long); genu (60–62.5 (62.5) long); tibia (80–82.5 (82.5) long; 20–20 (20) wide); tarsus (17.5–20 (17.5) long). Palpomere proportions: femur/genu 1.67–1.79 (1.72); tibia/femur 0.74–0.83 (0.77); tibia length/width 4.00–4.13 (4.13).

Venter — (620–670 (650) long; 410–430 (420) wide) with reddish-purple coloration, occasionally faint. Gnathosomal bay (135–145 (142.5) long; 62.5–70 (67.5) wide). Cxgl-4 subapical. Medial suture (60–65 (60) long). Genital plates (132.5–145 (140) long; 102.5–107.5 (105) wide). Additional measurements: Cx-1 (260–280 (280) long (total); 125–135 (135) long (medial)); Cx-3 (290–310 (310) wide); anterior venter (205–220 (205) long). Ventral proportions: gnathosomal bay length/width 2.07–2.23 (2.11); anterior venter/genital field length 1.46–1.57 (1.46); anterior venter length/genital field width 1.91–2.05 (1.95); anterior venter/medial suture 3.15–3.52 (3.42).

Immatures unknown.

Etymology

Specific epithet (danielleae) named in honor of Danielle Fisher—lab technician, environmental educator, colleague, friend, wife (of JRF), and mother of Ivy, our beautiful daughter—for her tireless and immense contributions to this research, and for bettering the lives of all those around her. Thank you, Danielle.

Distribution

Southern Appalachians, northeastern Georgia (Figure 47).

Figure 47. 

Torrenticola danielleae sp. n. distribution.

Figure 48. 

Torrenticola danielleae sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 49. 

Torrenticola danielleae sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Remarks

Unfortunately, we were unable to acquire fresh material of Torrenticola danielleae and therefore this species is not included in our phylogenetic analyses. However, we were able to examine morphology with material preserved in GAW. The overall appearance, elongate subcapitular rostra, and elongate pedipalpal tibiae, are consistent with placing this species in the Raptor Complex and the Raptor Identification Group.

Torrenticola delicatexa Habeeb, 1955

T. amplexa delicatexa: Habeeb 1955: 4; 1957: 1.

T. delicatexa: Habeeb 1961: 2; 1967: 3 • Viets 1987: 759.

Material examined

Type series. HOLOTYPE (♂): from USA, New Jersey, Sussex County, Flatbrookeville, Flatbrook, 12 Oct 1953, by H Habeeb, HH530113.

PARATYPES (1 ♀; 0 ♂): New Jersey, USA: 1 ♀ (ALLOTYPE) from Sussex County, Flatbrookeville, Flatbrook, 12 Oct 1953, by H Habeeb, HH530113.

OTHER MATERIAL (19 ♀; 7 ♂): North Carolina, USA: 1 ♀ from Haywood County, Great Smokey Mountains National Park, Cataloochee River (35°38'45"N, 83°4'34"W), 6 Sep 2009, by IM Smith, IMS090099 • Maine, USA: 1 ♀ and 1 ♂ from Franklin County, Small Falls picnic area beside Route 4, Sandy River (44°52'N, 70°31'W), 5 Jul 1989, by IM Smith, IMS890069 • New Hampshire, USA: 1 ♀ and 1 ♂ from Woodstock County, beside Route 118, Jackman Brook (44°0'N, 71°45'W), 11 Sep 1992, by IM Smith, IMS920036 • Pennsylvania, USA: 1 ♀ from Fayette County, Dunbar Creek (39°57'50"N, 79°35'8.70"W), 10 Aug 2014, by MJ Skvarla, MS 14-0810-001 • 2 ♀ and 2 ♂ from Fayette County, Ohiopyle State Park, Laurel Run (39°50'58"N, 79°30'51"W), 10 Aug 2014, by MJ Skvarla, MS 14-0810-005 • 2 ♀ from Somerset County, Laurel Hill State Park, Laurel Hill Creek (40°1'6"N, 79°14'4"W), 8 Aug 2014, by MJ Skvarla, MS 14-0808-001 • Quebec, Canada: 1 ♀ from Stanstead County, 1 kilometer south of Rock Island, Tomifobia River, Tompkin Stream, (45°0'31"N, 72°7'6"W), 20 Aug 1996, by IM Smith & M MacKenzie, IMS960056 • Tennessee, USA: 1 ♀ from Blount County, Great Smokey Mountains National Park, Abrams River (35°35'31"N, 83°51'21"W), 17 Sep 2010, by IM Smith, IMS100141 • 1 ♀ from Blount County, Great Smokey Mountains National Park, Little River (35°40'55"N, 83°39'6"W), 8 Sep 2009, by IM Smith, IMS090102 • 2 ♀ from Sevier County, Great Smokey Mountains National Park, middle prong of Little Pigeon River (35°43'34"N, 83°24'2"W), 10 Sep 2010, by IM Smith, IMS100127 • 1 ♀ from Sevier County, Great Smokey Mountains National Park, middle prong of Little Pigeon River (35°43'34"N, 83°24'2"W), 10 Sep 2010, by IM Smith, IMS100128 • 1 ♀ from Sevier County, Great Smokey Mountains National Park, Sugarlands Nature Trail, spring (35°40'47"N, 83°31'52"W), 18 Sep 2010, by IM Smith, IMS100147 • South Carolina, USA: 1 ♀ from Greenville County, Matthews Creek, 24 Apr 2014, by D Eargle, JRF 14-0424-001 • Vermont, USA: 1 ♀ and 1 ♂ from Addison County, beside road from Lincoln, Middlebury River, (44°0'N, 73°1'W), 6 Jul 1989, by IM Smith, IMS890075 • Virginia, USA: 1 ♀ and 1 ♂ from Montgomery County, Blacksburg, beside Route 321 at Caldwell, Craig Creek (37°20'0"N, 80°20'0"W), 12 Jul 1990, by IM Smith, IMS900089A • 1 ♀ and 1 ♂ from Patrick County, Round Meadow Creek (36°42'59"N, 80°25'29"W), 10 Jun 2006, by IM Smith, IMS060005A.

Type deposition

Holotype (♀) and allotype (♂) deposited in CNC.

Diagnosis

Torrenticola delicatexa are similar to other members of Rusetria “Eastern 2-Plates” group (T. biscutella, T. caerulea, T. feminellai, T. indistincta, T. malarkeyorum, T. microbiscutella, T. pendula, T. sellersorum, T. tysoni, T. ululata, andT. whitneyae) in having anterio-lateral platelets fused to the dorsal plate, having dorsal coloration separated into anterior and posterior portions (except T. indistincta and T. ululata), and being distributed in the east. T. delicatexa can be differentiated from T. ululata, T. indistincta, and T. feminellai by dorsal coloration and pattern. T. delicatexa can be differentiated from T. tysoni by having a stockier rostrum (length/width = 2.33–3.00 in T. delicatexa, 3.06–3.50 in A34). T. delicatexa can be differentiated from T. pendula by having a stockier gnathosomal bay (length/width = 1.28–2.22 in T. delicatexa, 2.42–2.90 in T. pendula), and a longer dorsum (♀ = 560–620 in T. delicatexa, 630–650 in T. pendula; ♂ = 420–465 in T. delicatexa, 500 in T. pendula). T. delicatexa can be differentiated from T. microbiscutella by having a less elongate dorsum (length/width = 1.38–1.56 in T. delicatexa, 1.63–1.75 in T. microbiscutella). Female T. delicatexa can be differentiated from female T. malarkeyorum, T. biscutella, and T. caerulea by having a longer genital field (175–185 in T. delicatexa, 153–170 in others). Male T. delicatexa can be differentiated from male T. biscutella by having a more ovoid dorsum (length/width = 1.44–1.56 in T. delicatexa; 1.37–1.42 in T. biscutella). Female T. delicatexa can be differentiated from female T. sellersorum by having a slightly more ovoid dorsum (length/width = 1.38–1.44 in T. delicatexa, 1.23–1.37 in T. sellersorum). Male T. delicatexa do not have any measurement differences with male T. malarkeyorum, T. caerulea, and T. sellersorum; however, they can be differentiated by dorsal coloration. T. delicatexa can be differentiated from T. whitneyae by having a slightly more ovoid dorsum (length/width ♀ = 1.38–1.44 in T. delicatexa, 1.26–1.38 in T. whitneyae; ♂ = 1.44–1.56 in T. delicatexa, 1.35–1.37 in T. whitneyae) and by dorsal coloration. Additionally, male T. delicatexa can be differentiated from male T. whitneyae by having more elongate pedipalpal tibiae (length/width = 2.89–3.63 in T. delicatexa, 2.48–2.70 in T. whitneyae).

Re-description

Male (Figure 51) (n = 5) (holotype measurements in parentheses when available) with characters of the genus with following specifications.

Dorsum — (410–465 (465) long; 270–320 (320) wide) ovoid with highly variable coloration, reddish-purple to purple (occasionally bluish-purple) separated into anterior and posterior portions. Anterio-medial platelets (83.75–102.5 (102.5) long; 30–32.5 (32.5) wide). Anterio-lateral platelets (122.5–132.5 (132.5) long; 40–55 (52.5) wide) fused to dorsal plate. Dgl-4 much closer to the edge of the dorsum than to the muscle scars (distance between Dgl-4 205–255 (255)). Dorsal plate proportions: dorsum length/width 1.44–1.56 (1.45); dorsal width/distance between Dgl-4 1.23–1.37 (1.25); anterio-medial platelet length/width 2.79–3.17 (3.15); anterio-lateral platelet length/width 2.36–3.06 (2.52); anterio-lateral/anterio-medial length 1.29–1.46 (1.29).

Gnathosoma — Subcapitulum (235–247.5 (247.5) long (ventral); 172.5–193.75 (194) long (dorsal); 97.5–107.5 (107.5) tall) colorless. Rostrum (90–100 (100) long; 32.5–37.5 (37.5) wide). Chelicerae (220-230 long) with curved fangs (45–52.5 (50) long). Subcapitular proportions: ventral length/height 2.30–2.45 (2.30); rostrum length/width 2.67–2.81 (2.67). Pedipalps with tuberculate ventral extensions on femora and genua. Palpomeres: trochanter (33.75–40 (40) long); femur (82.5–92.5 (92.5) long); genu (50–55 (55) long); tibia (62.5–72.5 (65) long; 20–22.5 (22.5) wide); tarsus (15–17.5 (17.5) long). Palpomere proportions: femur/genu 1.59–1.75 (1.68); tibia/femur 0.70–0.83 (0.70); tibia length/width 2.89–3.63 (2.89).

Venter — (490–540 (540) long; 311–435 (435) wide) usually colorless; occasionally with faint reddish-purple coloration. Gnathosomal bay (120–135 (135) long; 57.5–72.5 (72.5) wide). Cxgl-4 subapical. Medial suture (55–75 (62.5) long). Genital plates (105–110 (106.25) long; 95–102.5 (102.5) wide). Additional measurements: Cx-1 (220–235 (235) long (total); 78–100 (100) long (medial)); Cx-3 (240–297.5 (297.5) wide); anterior venter (160–175 (170) long). Ventral proportions: gnathosomal bay length/width 1.86–2.22 (1.86); anterior venter/genital field length 1.52–1.60 (1.60); anterior venter length/genital field width 1.64–1.79 (1.66); anterior venter/medial suture 2.33–2.91 (2.72).

Female (Figure 52) (n = 6) (allotypic measurements in parentheses when available) with characters of the genus with following specifications.

Dorsum — (560–620 (620) long; 390–440 (435) wide) ovoid with highly variable coloration, reddish-purple to purple (occasionally bluish-purple) separated into anterior and posterior portions. Anterio-medial platelets (125–140 (140) long; 36.25–45 (42.5) wide). Anterio-lateral platelets (152.5–172.5 (162.5) long; 57.5–70 (70) wide) fused to dorsal plate. Dgl-4 much closer to the edge of the dorsum than to the muscle scars (distance between Dgl-4 295–350 (330)). Dorsal plate proportions: dorsum length/width 1.38–1.44 (1.43); dorsal width/distance between Dgl-4 1.26–1.33 (1.32); anterio-medial platelet length/width 2.83–3.52 (3.29); anterio-lateral platelet length/width 2.26–2.88 (2.32); anterio-lateral/anterio-medial length 1.16–1.33 (1.16).

Gnathosoma — Subcapitulum (305–345 (345) long (ventral); 216–260 (260) long (dorsal); 145–165 (160) tall) colorless. Rostrum (115–135 (135) long; 45–55 (45) wide). Chelicerae (312–350 (350) long) with curved fangs (54–75 (70) long). Subcapitular proportions: ventral length/height 1.97–2.22 (2.16); rostrum length/width 2.33–3.00 (3.00). Pedipalps with tuberculate ventral extensions on femora and genua. Palpomeres: trochanter (42.5–52.5 (47.5) long); femur (105–130 (128.75) long); genu (67.5–75 (72.5) long); tibia (81.25–90 (87.5) long; 22.5–30 (30) wide); tarsus (20–22.5 (22.5) long). Palpomere proportions: femur/genu 1.56–1.78 (1.78); tibia/femur 0.65–0.77 (0.68); tibia length/width 2.92–3.61 (2.92).

Venter — (640–690 (690) long; 431–540 (540) wide) usually colorless; occasionally with faint reddish-purple coloration. Gnathosomal bay (132.5–175 (157.5) long; 85–117.5 (117.5) wide). Cxgl-4 subapical. Medial suture absent. Genital plates (175–197.5 (197.5) long; 150–172.5 (170) wide). Additional measurements: Cx-1 (216–295 (295) long (total); 93–135 (135) long (medial)); Cx-3 (304–400 (400) wide); anterior venter (115–135 (135) long). Ventral proportions: gnathosomal bay length/width 1.28–2.06 (1.34); anterior venter/genital field length 0.64–0.76 (0.68); anterior venter length/genital field width 0.71–0.83 (0.79).

Immatures unknown.

Etymology

Habeeb (1955) did not offer an explanation for the specific epithet (delicatexa) and we are unable to offer helpful speculation.

Distribution

Appalachians (Figure 50).

Figure 50. 

Torrenticola delicatexa distribution.

Figure 51. 

Torrenticola delicatexa female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 52. 

Torrenticola delicatexa male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Remarks

In all analyses, Torrenticola delicatexa groups with members of the Rusetria Complex with high support and specimens of this species were less than 2% different in COI sequence from each other. However, one specimen from Tennessee (DNA#1839) was 5% different; this specimen was collected from the same river as the other specimens and was indistinguishable morphologically. We refrain from speculating on this COI variation, but consider this specimen as an outlier, and thus within our hypothesis for T. delicatexa. The position of this species was only strongly supported in our combined analysis, where it is recovered as sister to two other species: T. ululata and T. glomerabilis. However, T. delicatexa does not resemble these species morphologically, and instead is quite similar to three more distantly-related species (T. biscutella, T. malarkeyorum, and T. caerulea).

This species hypothesis is supported by low COI divergence within the species (0–2%) and high divergence between species (3–15%), and by the morphological characters outlined in the diagnosis.

Torrenticola dentirostra Fisher & Dowling, sp. n.

Material examined

HOLOTYPE (♀): from USA, Georgia, Chattooga County, beside road from Everett Springs to Villanow 1.4 km south of The Pocket Recreation Area, 4 July 1990, by IM Smith, IMS900077.

PARATYPES (4 ♀; 2 ♂): Georgia, USA: 1 ♂ (ALLOTYPE) from Chattooga County, beside road from Everett Springs to Villanow 1.4 km south of The Pocket Recreation Area, 4 July 1990, by IM Smith, IMS900077 • 2 ♀ and 1 ♂ from Chattooga County, beside road from Everett Springs to Villanow 1.4 km south of The Pocket Recreation Area, 4 July 1990, by IM Smith, IMS900077 • Tennessee, USA: 1 ♀ from Monroe County, beside Forest Route 35, 2.0 km northeast of road from Rt. 165 to Miller Chapel Baptist Church, (35°21'21"N, 84°9'9"W), 5 July 1990, by IM Smith, IMS900078 • Virginia, USA: 1 ♀ from Alleghany County, Covington; beside Rt. 18, 0.5 km north of Rt. 657, (37°44'44"N, 80°2'2"W), 13 July 1990, by IM Smith, IMS900091A.

Type deposition

Holotype (♀), allotype (♂), and some paratypes (2 ♀) deposited in the CNC; other paratypes (2 ♀; 1 ♂) deposited in the ACUA.

Diagnosis

Torrenticola dentirostra are similar to other members of the Nigroalba Group (T. flangipalpa, T. nigroalba, and T. solisorta) in being small, slightly elongate, and having purple dorsal coloration restricted posteriorly. T. dentirostra can be differentiated from most Torrenticola (except T. erectirostra, T. karambita, and T. robisoni) by having a dentate bump midway on the dorsal edge of the rostrum. T. dentirostra can be differentiated from T. karambita, T. erectirostra, and T. robisoni by having a straight rostrum (others have upturned rostra) and by dorsal pattern.

Description

Female (Figure 54) (n = 5) (holotype measurements in parentheses when available) with characters of the genus with following specifications.

Dorsum — (470–560 (470) long; 340–390 (340) wide) ovoid with faint purple coloration restricted posteriorly. Anterio-medial platelets (107.5–120 (107.5) long; 45–47.5 (45) wide). Anterio-lateral platelets (157.5–177.5 (157.5) long; 55–62.5 (55) wide) free from dorsal plate. Dgl-4 closer to the edge of the dorsum than to the muscle scars (distance between Dgl-4 240–275 (240)). Dorsal plate proportions: dorsum length/width 1.38–1.45 (1.38); dorsal width/distance between Dgl-4 1.38–1.43 (1.42); anterio-medial platelet length/width 2.32–2.67 (2.39); anterio-lateral platelet length/width 2.84–3.05 (2.86); anterio-lateral/anterio-medial length 1.43–1.59 (1.47).

Gnathosoma — Subcapitulum (275–330 (275) long (ventral); 200–247.5 (200) long (dorsal); 77.5–102.5 (77.5) tall) colorless. Rostrum (100–120 (100) long; 35–40 (35) wide) with dentate bump midway on dorsal edge. Chelicerae (255–320 (255) long) with curved fangs (45–55 (45) long). Subcapitular proportions: ventral length/height 3.22–3.73 (3.55); rostrum length/width 2.80–3.00 (2.86). Pedipalps with tuberculate ventral extensions dentate apically on femora and genua. Palpomeres: trochanter (30–35 (30) long); femur (92.5–110 (92.5) long); genu (55–67.5 (55) long); tibia (80–92.5 (80) long; 17.5–20 (17.5) wide); tarsus (12.5–15 (13.75) long). Palpomere proportions: femur/genu 1.62–1.68 (1.68); tibia/femur 0.84–0.88 (0.86); tibia length/width 4.57–4.93 (4.57).

Venter — (610–685 (610) long; 390–425 (390) wide) mostly colorless with faint purple genital plates. Gnathosomal bay (85–110 (85) long; 60–80 (60) wide). Cxgl-4 far from apex and ventral. Medial suture (60–70 (60) long). Genital plates (142.5–165 (142.5) long; 120–135 (120) wide). Additional measurements: Cx-1 (230–260 (230) long (total); 140–160 (140) long (medial)); Cx-3 (270–300 (270) wide); anterior venter (210–240 (210) long). Ventral proportions: gnathosomal bay length/width 1.30–1.50 (1.42); anterior venter/genital field length 1.42–1.64 (1.47); anterior venter length/genital field width 1.70–1.83 (1.75); anterior venter/medial suture 3.29–3.69 (3.50).

Male (Figure 55) (n = 2) (allotypic measurements in parentheses when available) with characters of the genus with following specifications.

Dorsum — (480–505 (505) long; 360–360 (360) wide ovoid with faint purple coloration restricted posteriorly. Anterio-medial platelets (110–110 (110) long; 40–42.5 (42.5) wide). Anterio-lateral platelets (162.5–165 (165) long; 50–52.5 (52.5) wide) free from dorsal plate. Dgl-4 closer to the edge of the dorsum than to the muscle scars (distance between Dgl-4 240–245 (240)). Dorsal plate proportions: dorsum length/width 1.33–1.40 (1.40); dorsal width/distance between Dgl-4 1.47–1.50 (1.50); anterio-medial platelet length/width 2.59–2.75 (2.59); anterio-lateral platelet length/width 3.14–3.25 (3.14); anterio-lateral/anterio-medial length 1.48–1.50 (1.50).

Gnathosoma — Subcapitulum (285–287.5 (287.5) long (ventral); 210–217.5 (217.5) long (dorsal); 77.5–80 (77.5) tall) colorless. Rostrum (100–105 (100) long; 32.5–35 (32.5) wide) with dentate bump midway on dorsal edge. Chelicerae (255–265 (265) long) with curved fangs (40–45 (45) long). Subcapitular proportions: ventral length/height 3.56–3.71 (3.71); rostrum length/width 3.00–3.08 (3.08). Pedipalps with tuberculate ventral extensions dentate apically on femora and genua. Palpomeres: trochanter (30–32.5 (32.5) long); femur (90–92.5 (90) long); genu (57.5–57.5 (57.5) long); tibia (82.5–85 (85) long; 17.5–18.75 (18.75) wide); tarsus (12.5–13.75 (13.75) long). Palpomere proportions: femur/genu 1.57–1.61 (1.57); tibia/femur 0.89–0.94 (0.94); tibia length/width 4.53–4.71 (4.53).

Venter — (610–645 (645) long; 390–400 (400) wide) mostly colorless with faint purple genital plates. Gnathosomal bay (100–107.5 (407.5) long; 70–70 (70) wide). Cxgl-4 far from apex and ventral. Medial suture (75–85 (85) long). Genital plates (130–135 (135) long; 102.5–102.5 (102.5) wide). Additional measurements: Cx-1 (240–250 (250) long (total); 140–147.5 (147.5) long (medial)); Cx-3 (275–280 (275) wide); anterior venter (235–247.5 (247.5) long). Ventral proportions: gnathosomal bay length/width 1.43–1.54 (1.54); anterior venter/genital field length 1.81–1.83 (1.83); anterior venter length/genital field width 2.29–2.41 (2.41); anterior venter/medial suture 2.91–3.13 (2.91).

Immatures unknown.

Etymology

Specific epithet (dentirostra) refers to the diagnostic tooth-like serrations on the dorsal surface of the rostrum (dentis, L. tooth; rostrum, L. snout).

Distribution

Southern Appalachians (Figure 53).

Figure 53. 

Torrenticola dentirostra sp. n. distribution.

Figure 54. 

Torrenticola dentirostra sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 55. 

Torrenticola dentirostra sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Remarks

Unfortunately, we were unable to acquire fresh material of Torrenticola dentirostra and therefore this species is not included in our phylogenetic analyses. However, we were able to examine morphology with material preserved in GAW. The overall similarity, small size, elongate subcapitular rostra, elongate pedipalpal tibiae, and dorsal purple coloration restricted to posterior half, place this species in the Raptor Complex and the Nigroalba Identification Group.

Torrenticola dimorpha Fisher & Dowling, sp. n.

Material examined

HOLOTYPE (♀): from USA, Texas, Bandera County, Lost Maples State Natural Area, north of Vanderpool; picnic area, (29°48'48"N, 99°34'34"W), 10 October 2010, by IM Smith, IMS100184.

PARATYPES (8 ♀; 5 ♂): Texas, USA: 1 ♂ (ALLOTYPE) from Bandera County, Lost Maples State Natural Area, north of Vanderpool; picnic area, (29°48'48"N, 99°34'34"W), 10 October 2010, by IM Smith, IMS100184 • 1 ♀ and 1 ♂ from Bandera County, Lost Maples State Natural Area, north of Vanderpool; picnic area, (29°48'48"N, 99°34'34"W), 10 October 2010, by IM Smith, IMS100184 • 2 ♀ and 3 ♂ from Bandera County, Lost Maples State Natural Area north of Vanderpool; picnic area, (29°48'48"N, 99°34'34"W), 7 October 2010, by IM Smith, IMS100178 • 2 ♀ from Bandera County, Lost Maples State Natural Area; picnic area, (29°48'48"N, 99°34'34"W), 27 May 1997, by IM Smith, IMS970008 • 1 ♀ from Bandera County, Vanderpool; beside Rt. 187, 0.7 km south of entrance to Lost Maples State Natural Area, (29°48'48"N, 99°34'34"W), 2 May 2009, by IM Smith, IMS090007 • 1 ♀ from Kinney County, Brackettville; beside Rt. 90, 12.1 km west of Rt. 131, (29°20'20"N, 100°32'32"W), 4 May 2003, by IM Smith, IMS030007 • 1 ♀ from Uvalde County, Garner State Park; river crossing site, (29°35'35"N, 99°44'44"W), 28 May 1998, by IM Smith, IMS980027A.

Type deposition

Holotype (♀), allotype (♂), and some paratypes (5 ♀; 2 ♂) deposited in the CNC; other paratypes (3 ♀; 2 ♂) deposited in the ACUA.

Diagnosis

Torrenticola dimorpha are similar to other members of the Tricolor Complex (T. bittikoferae, T. cardia, T. hoosieri, T. kringi, T. larvata, T. mohawk, T. pearsoni, T. olliei, T. sierrensis, T. tricolor, T. trimaculata, and T. unimaculata) in having a short, conical rostrum. T. dimorpha can be differentiated from all other Torrenticola by having a dorsal plate with a medial extension covering nearly half the length of the anterio-medial platelets and by males having large, highly modified pedipalps, which are expanded vertically and laterally. Additionally, T. dimorpha can be differentiated from most other Tricolor Complex (except T. bittikoferae, T. hoosieri, and T. pearsoni) by being colorless, whereas most other members have bold patterning.

Description

Female (Figure 57) (n = 5) (holotype measurements in parentheses when available) with characters of the genus with following specifications.

Dorsum — (590–670 (650) long; 455–520 (490) wide) colorless and ovoid. Anterior dorsal plate with medial extension covering nearly half the length of the anterio-medial platelets. Anterio-medial platelets (117.5–135 (135) long; 52.5–57.5 (57.5) wide). Anterio-lateral platelets (145–172.5 (172.5)) long; 55–65 (62.5) wide) free from dorsal plate. Dgl-4 approximately halfway in between the edge of the dorsum and the muscle scars (distance between Dgl-4 320–360 (360)). Dorsal plate proportions: dorsum length/width 1.29–1.33 (1.33); dorsal width/distance between Dgl-4 1.36–1.48 (1.36); anterio-medial platelet length/width 2.24–2.35 (2.35); anterio-lateral platelet length/width 2.46–2.78 (2.76); anterio-lateral/anterio-medial length 1.19–1.33 (1.28).

Gnathosoma — Subcapitulum (230–260 (255) long (ventral); 150–170 (170) long (dorsal); 110–132.5 (132.5) tall) colorless. Posterior dorsal apodeme long. Rostrum (65–70 (70) long; 42.5–50 (45) wide) very short. Chelicerae (210–245 (235) long) with curved fangs (60–70 (70) long). Subcapitular proportions: ventral length/height 1.90–2.09 (1.92); rostrum length/width 1.40–1.56 (1.56). Pedipalps with long, tuberculate ventral extensions on femora and no ventral extensions on genua. Palpomeres: trochanter (37.5–40 (37.5) long); femur (76.25–85 (82.5) long); genu (52.5–62.5 (60) long); tibia (65–72.5 (70) long; 22.5–22.5 (22.5) wide); tarsus (20–22.5 (22.5) long). Palpomere proportions: femur/genu 1.36–1.48 (1.38); tibia/femur 0.84–0.87 (0.85); tibia length/width 2.89–3.22 (3.11).

Venter — (720–800 (770) long; 505–600 (580) wide) colorless. Gnathosomal bay (120–145 (145) long; 72.5–85 (85) wide). Cxgl-4 apical. Medial suture (25–30 (30)). Genital plates (170–195 (185) long; 162.5–180 (180) wide). Additional measurements: Cx-1 (240–270 (270) long (total); 110–130 (125) long (medial)); Cx-3 (315–350 (350) wide); anterior venter (137.5–155 (155) long). Ventral proportions: gnathosomal bay length/width 1.60–1.79 (1.71); anterior venter/genital field length 0.78–0.84 (0.84); anterior venter length/genital field width 0.85–0.86 (0.86).

Male (Figure 58) (n = 5) (allotypic measurements in parentheses when available) with characters of the genus with following specifications.

Dorsum — (465–510 (500) long; 310–330 (320) wide) colorless and ovoid. Anterior dorsal plate with medial extension covering nearly half the length of the anterio-medial platelets. Muscle scars absent or very faint. Anterio-medial platelets (102.5–115 (107.5) long; 42.5–45 (45) wide). Anterio-lateral platelets (177.5–197.5 (195) long; 42.5–55 (55) wide) free from dorsal plate. Dgl-4 much closer to the edge of the dorsum than to the muscle scars (distance between Dgl-4 225–245 (245)). Dorsal plate proportions: dorsum length/width 1.50–1.58 (1.56); dorsal width/distance between Dgl-4 1.31–1.41 (1.31); anterio-medial platelet length/width 2.34–2.63 (2.39); anterio-lateral platelet length/width 3.38–4.65 (3.55); anterio-lateral/anterio-medial length 1.64–1.81 (1.81).

Gnathosoma — Subcapitulum (227.5–255 (240) long (ventral); 160–177.5 (165) long (dorsal); 100–110 (100) tall) colorless. Rostrum (45–50 (47.5) long; 32.5–37.5 (32.5) wide) very short. Chelicerae (195–235 (205) long) with curved fangs (55–65 (60) long). Subcapitular proportions: ventral length/height 2.17–2.40 (2.40); rostrum length/width 1.29–1.46 (1.46). Pedipalps highly modified and expanded, with long, tuberculate ventral extensions on femora and no ventral extensions on genua. Palpomeres: trochanter (53.75–57.5 (55) long); femur (107.5–112.5 (112.5) long); genu (92.5–103.75 (103.75) long); tibia (77.5–85 (85) long; 30–35 (35) wide); tarsus (20–25 (22.5) long). Palpomere proportions: femur/genu 1.08–1.22 (1.08); tibia/femur 0.69–0.77 (0.76); tibia length/width 2.36–2.67 (2.43).

Venter — (550–620 (580) long; 350–370 (370) wide) colorless and highly modified with coxae II–IV forming a large ventral plate that covers the insertions of legs IV. Suture dividing coxae III and IV incomplete. Apodemes expanded internally. Gnathosomal bay (110–115 (112.5) long; 90–110 (90) wide) expanded to accommodate large pedipalps. Cxgl-4 apical. Medial suture (155–165 (155) long). Genital plates (100–120 (105) long; 105–110 (105) wide) triangular. Additional measurements: Cx-1 (230–260 (250) long (total); 130–145 (130) long (medial)); Cx-3 (335–345 (335) wide); anterior venter (297.5–315 (300) long). Ventral proportions: gnathosomal bay length/width 1.05–1.25 (1.25); anterior venter/genital field length 2.63–2.98 (2.86); anterior venter length/genital field width 2.70–2.86 (2.86); anterior venter/medial suture 1.88–2.03 (1.94).

Immatures unknown.

Etymology

Specific epithet (dimorpha) refers to the sexual dimorphism in terms of size and morphology (di, G. two; morphḗ, G. form), the resulting morphology is so unlike all other Torrenticolidae that upon first glance specimens appear to be a different family altogether.

Distribution

Texas (probably also extending southward into Mexico) (Figure 56).

Figure 56. 

Torrenticola dimorpha sp. n. distribution.

Figure 57. 

Torrenticola dimorpha sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 58. 

Torrenticola dimorpha sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Remarks

Torrenticola dimorpha groups with other members of the Tricolor Complex with high support and specimens of this species were less than 1% different in COI sequence from each other. In the combined analysis, T. dimorpha groups with T. larvata and these species are greater than 16% different in COI sequence from each other. This clade (T. dimorpha + T. larvata) is sister to all other eastern members of the Tricolor Complex.

This species is so unique that upon initial observation, specimens appear to be members of a different genus, especially the males. However, the short conical rostrum that is downturned in males is characteristic of other members of the Tricolor Identification Group.

This species hypothesis is supported by biogeography, high divergence between species, and by highly distinctive morphological characteristics outlined in the diagnosis. The high degree of sexual size dimorphism (males are 15–30% smaller than females) is only matched by most members of the Rusetria Complex, where males are usually 20–30% smaller than females.

Torrenticola dolichodactyla Fisher & Dowling, sp. n.

Material examined

HOLOTYPE (♂): from USA, New Mexico, Catron County, Cottonwood Campground beside Rt. 180 south of Rt. 12, (33°37'37"N, 108°54'54"W), 12 July 1987, by IM Smith, IMS870086.

PARATYPES (0 ♀; 0 ♂):

Type deposition

Holotype (♂) deposited in the CNC.

Diagnosis

Torrenticola dolichodactyla are similar to other members of the Rala Group (T. rala, T. lamellipalpis, T. boettgeri, T. kurtvietsi, T. keesdavidsi, and T. anoplopalpa) by being colorless, having incomplete hind coxal margins and being distributed in the southwest. T. dolichodactyla can be differentiated from all other Rala Group by having longer pedipalpal tarsi (♂ = 52.5 in T. dolichodactyla, 10–20 in others) and Dgl-4 closer to the muscle scars (dorsum width/distance between Dgl-4 ♂ = 1.74 in T. dolichodactyla, 1.16–1.48 in others).

Description

Female unknown.

Male (Figure 60) (n = 1) (holotype only) with characters of the genus with following specifications.

Dorsum — (735 long; 590 wide) circular and colorless. Anterio-medial platelets (177.5 long; 77.5 wide). Anterio-lateral platelets (227.5 long; 97.5 wide) free from dorsal plate. Dgl-4 halfway between the muscle scars and the edge of the dorsum (distance between Dgl-4 340). Dorsal plate proportions: dorsum length/width 1.25; dorsal width/distance between Dgl-4 1.74; anterio-medial platelet length/width 2.29; anterio-lateral platelet length/width 2.33; anterio-lateral/anterio-medial length 1.28.

Gnathosoma — Subcapitulum (325 long (ventral); 235 long (dorsal); 135 tall) colorless. Rostrum (120 long; 50 wide). Chelicerae (325 long) with curved fangs (65 long). Subcapitular proportions: ventral length/height 2.41; rostrum length/width 2.40. Pedipalps with elongate tarsi and tuberculate ventral extensions on femora and genua. Palpomeres: trochanter (55 long); femur (140 long); genu (80 long); tibia (110 long; 32.5 wide); tarsus (52.5 long). Palpomere proportions: femur/genu 1.75; tibia/femur 0.79; tibia length/width 3.38.

Venter — (850 long; 720 wide) colorless. Gnathosomal bay (170 long; 115 wide). Cxgl-4 apical. Medial suture (95 long). Genital plates (220 long; 170 wide). Additional measurements: Cx-1 (350 long (total); 175 long (medial)); Cx-3 (480 wide); anterior venter (305 long). Ventral proportions: gnathosomal bay length/width 1.48; anterior venter/genital field length 1.39; anterior venter length/genital field width 1.79; anterior venter/medial suture 3.21.

Immatures unknown.

Etymology

Specific epithet (dolichodactyla) refers to elongate pedipalp tarsus (dolichos, G. long; daktylos, G. finger), which is the most elongate of all Torrenticolidae.

Distribution

New Mexico (probably also Arizona) (Figure 59).

Figure 59. 

Torrenticola dolichodactyla sp. n. distribution.

Figure 60. 

Torrenticola dolichodactyla sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Remarks

Unfortunately, we were unable to acquire fresh material of Torrenticola dolichodactyla and therefore this species is not included in our phylogenetic analyses. However, we were able to examine morphology with material preserved in GAW. The overall appearance, incomplete hind coxal margins, distribution, and lack of color are consistent with placing this species in the Rala Identification Group.

Torrenticola dunni Fisher & Dowling, sp. n.

Material examined

Type series. HOLOTYPE (♀): from USA, Tennessee, Sevier County, Great Smokey Mountains National Park, Cosby Recreation Area (35°46'54"N, 83°13'2"W), 16 Sep 2010, by IM Smith, IMS100140, DNA 1289.

PARATYPES (9 ♀; 5 ♂): North Carolina, USA: 1 ♂ (ALLOTYPE) from Haywood County, Great Smokey Mountains National Park, Rough Fork Creek (35°37'31"N, 83°6'46"W), 20 Sep 2010, by IM Smith, IMS100148 • 2 ♀ and 2 ♂ from Haywood County, Great Smokey Mountains National Park, Rough Fork Creek (35°37'31"N, 83°6'46"W), 20 Sep 2010, by IM Smith, IMS100148 • 1 ♀ from Haywood County, Great Smokey Mountains National Park, Waterville (35°44'59"N, 83°6'42"W), 16 Sep 2010, by IM Smith, IMS100138 • South Carolina, USA: 2 ♀ from Greenville County, Matthews Creek, 24 Apr 2014, by D Eargle, JRF 14-0424-001 • Tennessee, USA: 1 ♀ from Blount County, Great Smokey Mountains National Park, Abrams River (35°35'31"N, 83°51'21"W), 17 Sep 2010, by IM Smith, IMS100141 • 1 ♀ and 1 ♂ from Sevier County, Great Smokey Mountains National Park (35°40'47"N, 83°31'48"W), 3 Sep 2009, by IM Smith, IMS090096 • 2 ♀ and 1 ♂ from Sevier County, Great Smokey Mountains National Park (35°43'33"N, 83°24'1"W), 12 Sep 2010, by IM Smith, IMS100131 • Virginia, USA: 1 ♀ from Smyth County, Mount Rogers National Recreation Area, beside Route 600, Little Laurel Creek, 10 Jul 1990, by IM Smith, IMS900086.

Type deposition

Holotype (♀), allotype (♂), and most paratypes (4 ♀; 2 ♂) deposited in the CNC; other paratypes (5 ♀; 2 ♂) deposited in the ACUA.

Diagnosis

Torrenticola dunni are similar to other members of the Rusetria “4-Plates” group (T. dunni, T. glomerabilis, T. kittatinniana, T. pollani, T. rufoalba and T. shubini) and T. skvarlai in having anterio-lateral platelets free from the dorsal plate, dorsal coloration separated into anterior and posterior portions, and indistinct hind coxal margins. T. dunni can be differentiated from T. pollani by having a larger dorsum (length ♀ = 605–680 in T. dunni, 535–560 in T. pollani; ♂ = 500–540 in T. dunni, 440–492 in T. pollani; width ♀ = 440–490 in T. dunni, 410–420 in T. pollani; ♂ = 350–370 in T. dunni, 310–340 in T. pollani) ; and a stockier rostrum (length/width = 2.80–3.14 in T. dunni, 3.27–3.82 in T. pollani). Female T. dunni can be differentiated from female T. shubini by having a thinner rostrum (length/width = 2.8–3.0 in T. dunni, 2.5–2.7 in T. shubini). Male T. dunni can be differentiated from male T. shubini by having a longer anterior venter (277–285 in T. dunni, 215–238 in T. shubini). T. dunni can be differentiated from T. glomerabilis by having Dgl-4 closer to the dorsal edge (dorsal width/distance between Dgl-4 = 1.2–1.4 in T. dunni, 1.5–1.7 in T. glomerabilis) and stockier tibiae (length/width ♀ = 3.27–3.50 in T. dunni, 4.11–4.50 in T. glomerabilis, ♂ = 3.25–3.44 in T. dunni, 3.55–4.38 in T. glomerabilis). Female T. dunni can be differentiated from female T. kittatinniana by having a longer pedipalp genu (70–75 in T. dunni, 64 in T. kittatinniana); a longer subcapitulum (ventral length = 330–355 in T. dunni, 310 in T. kittatinniana); and anterio-medial platelets more elongate (length/width = 2.33–2.54 in T. dunni, 2.83 in T. kittatinniana). Male T. dunni can be differentiated from male T. kittatinniana by having a longer anterior venter (277–285 in T. dunni, 235 in T. kittatinniana) and wider dorsum (350–370 in T. dunni, 340 in T. kittatinniana). T. dunni can be differentiated from T. rufoalba by having a larger dorsum (length ♀ = 605–680 in T. dunni, 550 in T. rufoalba; ♂ = 500–540 in T. dunni, 440 in T. rufoalba; width ♀ = 440–490 in T. dunni, 400 in T. rufoalba; ♂ = 350–370 in T. dunni; 320 in T. rufoalba). T. dunni can be differentiated from T. skvarlai by having a conical pedipalpal femoral tubercle, whereas T. skvarlai has a broad and flat pedipalpal femoral tubercle, and by having a longer anterior venter (♀ = 160–190 in T. dunni, 140–152.5 in T. skvarlai; ♂ = 277.5–285 in T. dunni, 177.5–205 in T. skvarlai).

Description

Female (Figure 62) (n = 5) (holotype measurements in parentheses when available) with characters of the genus with following specifications.

Dorsum— (605–680 (655) long; 440–490 (460) wide) ovoid with purple or reddish-purple coloration separated into anterior and posterior portions, and occasionally with faint orange medially. Anterio-medial platelets (117.5–125 (125) long; 46.25–52.5 (50) wide). Anterio-lateral platelets (172.5–197.5 (192.5) long; 62.5–68.75 (62.5) wide) free from dorsal plate. Dgl-4 much closer to the edge of the dorsum than to the muscle scars (distance between Dgl-4 315–380 (340)). Dorsal plate proportions: dorsum length/width 1.37–1.42 (1.42); dorsal width/distance between Dgl-4 1.29–1.40 (1.35); anterio-medial platelet length/width 2.33–2.54 (2.50); anterio-lateral platelet length/width 2.76–3.08 (3.08); anterio-lateral/anterio-medial length 1.44–1.61 (1.54).

Gnathosoma — Subcapitulum (330–355 (345) long (ventral); 250–265 (255) long (dorsal); 132.5–150 (150) tall) colorless. Rostrum (130–140 (135) long; 45–50 (45) wide). Chelicerae (325–355 (350) long) with curved fangs (60–65 (60) long). Subcapitular proportions: ventral length/height 2.30–2.53 (2.30); rostrum length/width 2.80–3.00 (3.00). Pedipalps with tuberculate ventral extensions on femora and genua. Palpomeres: trochanter (42.5–50 (48.75) long); femur (117.5–132.5 (131.25) long); genu (70–75 (75) long); tibia (85–95 (90) long; 25–27.5 (27.5) wide); tarsus (20–20 (20) long). Palpomere proportions: femur/genu 1.68–1.83 (1.75); tibia/femur 0.69–0.72 (0.69); tibia length/width 3.27–3.50 (3.27).

Venter — (710–810 (780) long; 540–600 (600) wide) with faint bluish-purple or reddish purple coloration or colorless. Gnathosomal bay (157.5–175 (175) long; 92.5–115 (115) wide). Cxgl-4 subapical. Medial suture (20–25 (20) long). Genital plates (160–185 (177.5) long; 145–160 (152.5) wide). Additional measurements: Cx-1 (290–330 (330) long (total); 140–160 (155) long (medial)); Cx-3 (365–410 (400) wide); anterior venter (160–190 (190) long). Ventral proportions: gnathosomal bay length/width 1.50–1.70 (1.52); anterior venter/genital field length 0.99–1.07 (1.07); anterior venter length/genital field width 1.08–1.25 (1.25); anterior venter/medial suture 7.60–9.50 (9.50).

Male (Figure 63) (n = 5) (allotypic measurements in parentheses when available) with characters of the genus with following specifications.

Dorsum — (500–540 (540) long; 350–370 (360) wide) ovoid with purple or reddish-purple coloration separated into anterior and posterior portions, and occasionally with faint orange medially. Anterio-medial platelets (95–102.5 (100) long; 37.5–42.5 (41.25) wide). Anterio-lateral platelets (165–172.5 (172.5) long; 55–60 (55) wide) free from dorsal plate. Dgl-4 much closer to the edge of the dorsum than to the muscle scars (distance between Dgl-4 265–295 (285)). Dorsal plate proportions: dorsum length/width 1.35–1.53 (1.50); dorsal width/distance between Dgl-4 1.24–1.32 (1.26); anterio-medial platelet length/width 2.41–2.56 (2.42); anterio-lateral platelet length/width 2.88–3.14 (3.14); anterio-lateral/anterio-medial length 1.66–1.74 (1.73).

Gnathosoma — Subcapitulum (275–285 (285) long (ventral); 205–215 (215) long (dorsal); 102.5–115 (105) tall) colorless. Rostrum (105–112.5 (110) long; 35–38.75 (35) wide). Chelicerae (265–280 (275) long) with curved fangs (50–55 (52.5) long). Subcapitular proportions: ventral length/height 2.43–2.71 (2.71); rostrum length/width 2.90–3.14 (3.14). Pedipalps with tuberculate ventral extensions on femora and genua. Palpomeres: trochanter (40–47.5 (40) long); femur (105–107.5 (107.5) long); genu (62.5–66.25 (65) long); tibia 77.5–85 (77.5) long; 22.5–25 (22.5) wide); tarsus (17.5–20 (20) long). Palpomere proportions: femur/genu 1.58–1.72 (1.65); tibia/femur 0.72–0.79 (0.72); tibia length/width 3.25–3.44 (3.44).

Venter — (640–660 (655) long; 440–470 (460) wide) with faint bluish-purple or reddish purple coloration or colorless. Gnathosomal bay (125–135 (130) long; 75–82.5 (82.5) wide). Cxgl-4 subapical. Medial suture (120–135 (125) long). Genital plates (130–137.5 (135) long; 85–90 (87.5) wide). Additional measurements: Cx-1 (260–275 (275) long (total); 135–145 (145) long (medial)); Cx-3 (330–350 (335) wide); anterior venter (277.5–285 (285) long). Ventral proportions: gnathosomal bay length/width 1.56–1.80 (1.58); anterior venter/genital field length 2.04–2.19 (2.11); anterior venter length/genital field width 3.11–3.35 (3.26); anterior venter/medial suture 2.06–2.33 (2.28).

Immatures unknown.

Etymology

Specific epithet (dunni) named in honor of Rob Dunn of North Carolina State University, for his exceptional writings and research that personalize ecology by bringing nature indoors; and particularly for his storytelling ability, in which he wonderfully conveys that we humans, rather than being separate from nature, are indeed just as wild as what we perceive outdoors—a sentiment exemplified by his book, Wild Life of Our Bodies: Predators, Parasites, and Partners That Shape Who We Are Today (2011).

Distribution

Southeastern Appalachians (Figure 61).

Figure 61. 

Torrenticola dunni sp. n. distribution.

Figure 62. 

Torrenticola dunni sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 63. 

Torrenticola dunni sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Remarks

In all analyses, Torrenticola dunni groups with two other members of the Rusetria Complex with high support and specimens are less than 1% different in COI sequence from each other. In all analyses, T. dunni groups with two other species: T. pollani and T. shubini. These species are greater than 5% different from each other in COI sequence and the ranges of T. T. dunni overlaps with T. shubini, but the ranges of these species do not overlap with T. pollani. Given our collection efforts across the Appalachians, it is reasonable to speculate that T. dunni is restricted to the southern Appalachians.

Based upon overall similarity, dorso-lateral platelet fusion, and distribution, we were able to place this species within the Eastern 2-Plate Identification Group

This species hypothesis is supported by biogeography, low COI divergence within the species (0–2%) and high divergence between species (3–15%), and by the morphological characters outlined in the diagnosis.

Torrenticola ellipsoidalis (Marshall, 1943)

Atractides ellipsoidalis: Marshall 1943: 308.

T. ellipsoidalis: Mitchell 1954: 40 • Habeeb 1955: 2 • Viets 1956: 241 • Crowell 1961: 330 • Habeeb 1967: 3 • Conroy 1968: 28 • Habeeb 1974: 1 • Conroy and Scudder 1975: 307 • Quaglia and Conroy 1984: 89 • Viets 1987: 759.

Torrenticola rectiforma Habeeb, 1974: 1.

Material examined

Type series. HOLOTYPE (♀): from USA, California, Nevada County, north of Lake Tahoe, Martis Creek, Jun 1933, by PR Needham, RM330010.

PARATYPES (1 ♀; 0 ♂): California, USA: 1 ♀ from Nevada County, north of Lake Tahoe, Martis Creek, Jun 1933, by PR Needham, RM330010.

OTHER MATERIAL (94 ♀; 94 ♂): Alberta, Canada: 1 ♀ and 2 ♂ from Waterton Lakes National Park, Cameron Creek, beside Akamina Parkway (49°5'N, 113°52'W), 2 Aug 1985, by IM Smith, IMS850133 • 2 ♀ and 2 ♂ from Waterton Lakes National Park, Cameron Creek, beside Akamina Parkway, west of Rowe Creek, 27 Jun 1980, by IM Smith, IMS800086A • 1 ♀ and 2 ♂ from Waterton Lakes National Park, Cameron Creek, beside Akamina Parkway, 12-15 Jun 1980, by IM Smith, IMS800069A & IMS800069B • 2 ♀ and 2 ♂ from Waterton Lakes National Park, Little Prairie Picnic Area, Cameron Creek, 5-9 Jun 1980, by IM Smith, IMS800053B • British Columbia, Canada: 1 ♀ and 1 ♂ from Bella Coola Valley, Tweedsmuir Provincial Park, Atnarko River, at campground, 28 Jul 1983, by IM Smith & AB Smith, IMS830054A • 1 ♀ and 1 ♂ from Bella Coola Valley, Tweedsmuir Provincial Park, Atnarko Slough, beside Highway 20, west of Youngs Creek Picnic Area, 4 Aug 1983, by IM Smith, IMS830064 • 1 ♀ from Bella Coola Valley, Tweedsmuir Provincial Park, Belarko, Atnarko River, 24-26 Jul 1983, by IM Smith, IMS830049B • 1 ♀ and 2 ♂ from Bella Coola Valley, Tweedsmuir Provincial Park, Hotnarko River, at end of Atnarko tote road, 27 Jul 1983, by IM Smith, IMS830052 • 2 ♀ and 3 ♂ from Bella Coola Valley, Tweedsmuir Provincial Park, Hotnarko River, at end of Atnarko tote road, 31 Jul 1983, by IM Smith, IMS830059A & IMS830059B • 3 ♂ from Bella Coola Valley, Tweedsmuir Provincial Park, Youngs Creek Picnic Area, Atnarko Slough, 24-27 Jul 1983, by IM Smith, IMS830048A • 2 ♀ and 1 ♂ from Fernie, Lizard Creek, beside Highway 3, 1.8 km west of Fernie Mountain Provincial Park, 16 Aug 2012, by IM Smith, IMS120073 • 1 ♀ and 1 ♂ from Tweedsmuir Provincial Park, Youngs Creek, beside Highway 20, between Heckman Pass & Bella Coola Valley, 5 Aug 1983, by IM Smith, IMS830065 • 2 ♀ and 2 ♂ from Vancouver Island, Caycuse, Nixon Creek, 8 Jul 1976, by IM Smith, IMS760197 & IMS760198 • 1 ♀ and 2 ♂ from Vancouver Island, beside Highway 4, 35.6 kilometers east of Pacific Rim Road, 9 Jul 1976, by IM Smith, IMS760206 • 1 ♀ and 1 ♂ from Vancouver Island, Lake Cowichan, Cowichan River, above Skutz Falls, 9 Jul 1979, by IM Smith, IMS790035 • 1 ♀ and 1 ♂ from Vancouver Island, Lake Cowichan, Skutz Falls, Skutz Creek, near Cowichan River, 9 Jul 1979, by IM Smith, IMS790036A & IMS790036B • 1 ♀ and 2 ♂ from Vancouver Island, Lake Cowichan, South Shore Road, north of Mesachie Lake, Robertson River, 4 Jul 1976, by IM Smith, IMS760183A • 1 ♀ from Vancouver Island, Lake Cowichan, South Shore Road, north of Mesachie Lake, tributary of Robertson River, 4-10 Jul 1976, by IM Smith, IMS760182 • 1 ♀ and 2 ♂ from Vancouver Island, Lost Shoe Creek, beside Highway 4, 1.3 kilometers east of Pacific Rim Road, 9 Jul 1976, by IM Smith, IMS760202 • 1 ♀ and 2 ♂ from Vancouver Island, Malahat, Goldstream Provincial Park, Goldstream River, 26 Jun 1979, by IM Smith, IMS790028A • 1 ♂ from Vancouver Island, spring run beside North Shore Road, 1.7 kilometers north of Lake Cowichan, 7 Jun 1979, by IM Smith, IMS790008A • 3 ♀ and 2 ♂ from Vancouver Island, North Shore Road, 3.2 kilometers south of Youbou, 4 Jul 1976, by IM Smith, IMS760190 • 1 ♀ from Vancouver Island, Youbou, Shaw Creek, North Shore Road, 4.3 kilometers south of north end of Cowichan Lake, 8 Jul 1976, by IM Smith, IMS760196 • 1 ♀ from Vancouver Island, spring run beside South Shore Road, 2.3 kilometers north of Lake Cowichan, 6 Jun 1979, by IM Smith, IMS790007 • 1 ♀ and 1 ♂ from Vancouver Island, Ucluelet, beside Highway 4, 16.6 kilometers east of Pacific Rim Road, 18-19 Jul 1979, by IM Smith, IMS790047 • California, USA: 1 ♀ and 1 ♂ from Humboldt County, Prairie Creek State Park, Prairie Creek, 12 Jul 1964, by H Habeeb, HH640021 • 1 ♀ from Humboldt County, Trinidad, stream beside Patrick Point Road, near Bishop Pine Lodge, 7 Aug 1987, by IM Smith, IMS870134 • 1 ♀ from Inyo County, Inyo National Forest, Bishop Creek (37°17'23"N, 118°33'14"W), 2 Sep 2013, by JR Fisher, JRF 13-0902-003 • 1 ♂ from Mendocino County, Cottaneva Creek, beside Route 1, 21.8 kilometers southwest of Route 101, 5 Aug 1987, by IM Smith, IMS870129A • 2 ♀ and 3 ♂ from Mendocino County, small stream at beach access road, off Route 1, 2.6 kilometers south of Westport, 5 Aug 1987, by IM Smith, IMS870128A • 1 ♀ from Mono County, Humboldt-Toiyabe National Forest, Leavitt Creek (38°18'40"N, 119°34'49"W), 31 Aug 2013, by JR Fisher, JRF 13-0831-004 • 1 ♂ from Monterey County, Los Padres National Forest, Salmon Creek (35°48'57"N, 121°21'29"W), 6 Sep 2013, by JR Fisher, JRF 13-0906-003 • 2 ♀ and 2 ♂ from Monterey County, Nacimiento River, beside Nacimiento-Ferguson Road at Nacimiento campground, 30 Jul 1987, by IM Smith, IMS870120A • 2 ♀ and 2 ♂ from Monterey County, Salmon Creek, beside Route 1, south of Gorda, 28 Jul 1987, by IM Smith, IMS870114A • 1 ♀ from Nevada County, beside Route 89, north of Hobart Mills, 13 Jun 1976, by IM Smith, IMS760109 • 1 ♀ and 2 ♂ from San Bernardino County, Claremont, Mt Baldy, 3.5 kilometers above Mt Baldy Village, 24 Jul 1987, by IM Smith, IMS870107 • 1 ♀ from San Bernardino County, Claremont, Mt Baldy, San Antonio Falls, above Monker Flats, 24 Jul 1987, by IM Smith, IMS870105 • 3 ♀ and 2 ♂ from San Bernardino County, Claremont, Mt Baldy, stream below San Antonio Falls, above Monker Flats, 24 Jul 1987, by IM Smith, IMS870106 • 2 ♀ and 2 ♂ from Shasta County, Battle Creek, beside Route 44, 5.6 kilometers west of Viola, 10 Aug 1987, by IM Smith, IMS870139A • 1 ♀ and 1 ♂ from Trinity County, small cascading trickle beside Route 36, 5.2 kilometers west of Forest Glen Station, 6 Aug 1987, by IM Smith, IMS870132 • 3 ♂ from Tulare County, Stony Creek at Stony Creek Picnic Area, east of Sequoia National Park, 1 Aug 1987, by IM Smith, IMS870124A • 1 ♀ and 1 ♂ from Ventura County, Ojai, North Fork of Ventura River, beside Route 33, just above Wheeler Gorge, 25-26 Jul 1987, by IM Smith, IMS870109A • Idaho, USA: 1 ♀ from Custer County, Challis National Forest, Stanley Creek (44°15'12"N, 115°0'19"W), 30 Jul 2012, by JR Fisher, WA Nelson, & JC O’Neill, ROW 12-0730-005 • 1 ♂ from Custer County, Salmon River (44°12'31"N, 114°55'51"W), 29 Jul 2012, by JR Fisher, WA Nelson, & JC O’Neill, ROW 12-0729-003 • 2 ♀ and 3 ♂ from Lemhi County, North Fork of Salmon River, beside Route 93, 15 kilometers north of North Fork, 1 Jul 1985, by IM Smith, IMS850062 • Montana, USA: 2 ♀ from Flathead County, stream beside Route 487 near north end of Whitefish Lake, 29 Jun 1985, by IM Smith, IMS850057 • 3 ♀ and 4 ♂ from Lake County, stream beside Route 83, 39.5 kilometers north of Condon, 30 Jun 1985, by IM Smith, IMS850059A • 1 ♀ from Missoula County, Lolo National Forest, Lolo Creek (46°46'7"N, 114°27'53"W), 7 Aug 2012, by JR Fisher, WA Nelson, & JC O’Neill, ROW 12-0807-003 • 1 ♂ from Ravalli County, Bitterroot National Forest, East Fork Bitterroot River (45°51'40"N, 114°1'46"W), 3 Aug 2012, by JR Fisher, WA Nelson, & JC O’Neill, ROW 12-0803-005 • Oregon, USA: 1 ♀ and 1 ♂ from Benton County, Marys Peak near Philomath, Parker Creek, 27-28 Jun 1983, by IM Smith & AB Smith, IMS830006 • 1 ♀ from Clackamas County, Rhododendron Pioneer Tollgate campground, Zigzag River, 27 Jun 1976, by IM Smith, IMS760164 • 1 ♀ and 2 ♂ from Curry County, Port Orford, Butler Bar campground, Elk River, 25 Jun 1976, by IM Smith, IMS760162 • 1 ♂ from Coos County, Siskiyou National Forest, Road 33 between Powers & Agness, Daphne Grove campground, Coquille River, 2 Jul 1983, by IM Smith, IMS830016 • 1 ♀ and 1 ♂ from Coos County, Siskiyou National Forest, Road 33 between Powers & Agness, Daphne Grove campground, 2 Jul 1983, by IM Smith, IMS830017 • 2 ♀ and 5 ♂ from Curry County, Port Orford, Humbug Mountain State Park Picnic Area, beside Route 1, Brush Creek, 1 Jul 1983, by IM Smith, IMS830012 • 4 ♀ and 4 ♂ from Curry County, Port Orford, Humbug Mountain State Park Picnic Area, Brush Creek, beside Route 1, 3 Jul 1983, by IM Smith, IMS830020A & IMS830020B • 1 ♀ from Curry County, Port Orford, Humbug Mountain State Park Picnic Area, beside Route 1, 1 Jul 1983, by IM Smith, IMS830013 • 1 ♀ from Curry County, Rogue River National Forest, Elk River (42°42'46"N, 124°18'41"W), 13 Aug 2013, by JR Fisher, JRF 13-0813-003 • 2 ♀ and 2 ♂ from Curry County, Siskiyou National Forest, road 33 between Powers and Agness, North Fork of Foster Creek, 2 Jul 1983, by IM Smith, IMS830019 • 1 ♀ from Curry County, Sixes, Sixes River, beside road at mouth of Edson Creek, 4 Jul 1983, by IM Smith, IMS830021A • 1 ♀ from Grant County, Prairie City, Lunch Creek, beside Route 26. east of Dixie Pass, 17-20 Jun 1976, by IM Smith, IMS760125 • 1 ♀ and 1 ♂ from Grant County, Prairie City, Strawberry Forest Camp, Strawberry Creek, 17-20 Jun 1976, by IM Smith, IMS760126 • 2 ♀ from Lane County, Gate Creek (44°8'48"N, 122°34'20"W), 11 Aug 2013, by JC O’Neill, & WA Nelson, JNOW 13-0811-001 • 3 ♀ and 1 ♂ from Marion County, Marion Forks Riverside campground, North Fork of Santiam River, 22 Jun 1976, by IM Smith, IMS760145 • 3 ♀ and 4 ♂ from Multnomah County, Columbia River Scenic Highway, Horsetail Falls, 27 Jun 1983, by IM Smith, IMS830005 • 1 ♂ from Tillamook County, Siuslaw National Forest, Alder Creek (45°9'27"N, 123°47'60"W), 6 Aug 2013, by JC O’Neill, & WA Nelson, JNOW 13-0806-002 • Washington, USA: 2 ♀ from Lewis County, Gifford Pinchot National Forest (46°39'49"N, 121°41'11"W), 23 Jul 2013, by JC O’Neill, & WA Nelson, JNOW 13-0723-005 • 2 ♀ from Mason County, Olympic National Forest, Cabin Creek (47°35'44"N, 123°7'39"W), 22 Jul 2013, by JC O’Neill, & WA Nelson, JNOW 13-0722-004 • 2 ♀ from Snohomish County, Mount Baker National Forest, Marten River (48°4'19"N, 121°36'24"W), 28 Jul 2013, by JC O’Neill, & WA Nelson, JNOW 13-0728-002 • Wyoming, USA: 1 ♂ from Johnson County, Bighorn Mountains, Clear Creek, west of Buffalo Mosier Gulch Picnic Area, 28 Jul 2012, by IM Smith, IMS120041 • 1 ♀ and 1 ♂ from Johnson County, Crazy Woman Creek, beside Route 16, 14.9 kilometers west of road to South Fork campground, 18 Aug 1987, by IM Smith, IMS870158A • 1 ♀ and 2 ♂ from Washakie County, Ten Sleep, Ten Sleep Creek, beside Route 16, 4 kilometers west of Route 435, 18 Aug 1987, by IM Smith, IMS870157 • 2 ♀ and 1 ♂ from Washakie County, Ten Sleep Creek, Ten Sleep Wigwam Rearing Station, 26 Jul 2012, by IM Smith, IMS120044.

Type deposition

Holotype (♀) and paratype (1 ♀) deposited in the CNC.

Diagnosis

Torrenticola ellipsoidalis are similar to other members of the Ellipsoidalis Group (T. multiforma, T. occidentalis, and T. leviathan), in being among the largest Torrenticola in the west (dorsum length ♀ = 700–885; ♂ = 665–850), although T. sierrensis are also large (dorsum length ♀ = 700–880; ♂ = 590–735) but can easily be distinguished from the Ellipsoidalis Group by being circular instead of ellipsoid or rectangular (dorsum length/width = 1.17–1.28 in T. sierrensis, 1.30–1.67 in Ellipsoidalis Group). T. ellipsoidalis can be differentiated from T. multiforma by having stockier subcapitular rostra (length/width = 1.8–2.1 in T. ellipsoidalis, 2.5–2.8 in T. multiforma). T. ellipsoidalis can be differentiated from T. occidentalis (only known from females) by having a longer medial suture (40–57.5 in T. ellipsoidalis, 20 in T. occidentalis) and by having stockier anterio-lateral platelets (length/width = 2.00–2.39 in T. ellipsoidalis, 2.54 in T. occidentalis). T. ellipsoidalis can be differentiated from T. leviathan by having less elongate pedipalpal tibiae (length/width = 2.6–3.3 in T. ellipsoidalis, 3.4–4.2 in T. leviathan) and stockier anterio-medial platelets (length/width ♀ = 1.43–1.72 in T. ellipsoidalis, 1.94–2.14 in T. leviathan; ♂ = 1.53–2.00 in T. ellipsoidalis, 2.15 in T. leviathan).

Re-description

Female (Figure 65) (n = 8) (holotype measurements in parentheses when available) with characters of the genus with following specifications.

Dorsum — (765–885 (800) long; 520–605 (540) wide) rectangular and usually colorless, occasionally with faint purple coloration without distinct pattern. Anterio-medial platelets (127.5–147.5 137.5) long; 80–97.5 (80) wide). Anterio-lateral platelets (207.5–235 (217.5) long; 90–115 (92.5) wide) free from dorsal plate. Dgl-4 much closer to the edge of the dorsum than to the muscle scars (distance between Dgl-4 390–470 (410)). Dorsal plate proportions: dorsum length/width 1.41–1.64 (1.48); dorsal width/distance between Dgl-4 1.27–1.40 (1.32); anterio-medial platelet length/width 1.43–1.72 (1.72); anterio-lateral platelet length/width 2.00–2.39 (2.35); anterio-lateral/anterio-medial length 1.48–1.77 (1.58).

Gnathosoma — Subcapitulum (285–315 (310) long (ventral); 194–219 (215) long (dorsal); 145–165 (165) tall) colorless. Rostrum (115–127.5 (117.5) long; 57.5–62.5 (60) wide) short and conical. Chelicerae (261–289 long) with curved fangs (61–74 long). Subcapitular proportions: ventral length/height 1.82–2.07 (1.88); rostrum length/width 1.84–2.09 (1.96). Pedipalps with tuberculate ventral extensions on femora and genua. Palpomeres: trochanter (37.5–42.5 (40) long); femur (101.25–107.5 (102.5) long); genu (70–77.5 (77.5) long); tibia (80–90 (85) long; 26.25–27.5 (27.5) wide); tarsus (17.5–21.25 (20) long). Palpomere proportions: femur/genu 1.32–1.46 (1.32); tibia/femur 0.76–0.85 (0.83); tibia length/width 2.91–3.27 (3.09).

Venter — (885–1000 (935) long; 605–700 (605) wide) colorless. Gnathosomal bay (157.5–180 (177.5) long; 80–105 (85) wide). Cxgl-4 subapical. Medial suture (40–57.5 (47.5) long). Genital plates (201.25–222.5 (205) long; 167.5–195 (172.5) wide). Additional measurements: Cx-1 (308–337.5 (335) long (total); 122–162.5 (162.5) long (medial)); Cx-3 (393–440 (405) wide); anterior venter (210–237.5 (225) long). Ventral proportions: gnathosomal bay length/width 1.50–2.12 (2.09); anterior venter/genital field length 1.01–1.13 (1.10); anterior venter length/genital field width 1.13–1.30 (1.30); anterior venter/medial suture 3.83–5.94 (4.74).

Male (Figure 66) (n = 6) with characters of the genus with following specifications.

Dorsum — (725–850 long; 450–565 wide) rectangular and usually colorless, occasionally with faint purple coloration without distinct pattern. Anterio-medial platelets (122.5–165 long; 72.5–95 wide). Anterio-lateral platelets (195–230 long; 85–107.5 wide) free from dorsal plate. Dgl-4 much closer to the edge of the dorsum than to the muscle scars (distance between Dgl-4 350–460). Dorsal plate proportions: dorsum length/width 1.38–1.67; dorsal width/distance between Dgl-4 1.22–1.43; anterio-medial platelet length/width 1.53–2.00; anterio-lateral platelet length/width 2.07–2.36; anterio-lateral/anterio-medial length 1.39–1.67.

Gnathosoma — Subcapitulum (280–290 long (ventral); 196–203.75 long (dorsal); 138.75–155 tall) colorless. Rostrum (102.5–115 long; 52.5–60 wide) short and conical. Chelicerae (263–280 long) with curved fangs (60–74 long). Subcapitular proportions: ventral length/height 1.87–2.04; rostrum length/width 1.86–2.02. Pedipalps with tuberculate ventral extensions on femora and genua. Palpomeres: trochanter (35–41.25 long); femur 92.5–100 long); genu (65–72.5 long); tibia (72.5–80 long; 25–30 wide); tarsus (17.5–20 long). Palpomere proportions: femur/genu 1.31–1.46; tibia/femur 0.74–0.84; tibia length/width 2.64–3.1.

Venter — (840–980 long; 469–653 wide) colorless. Gnathosomal bay (147.5–177.5 long; 77.5–90 wide). Cxgl-4 subapical. Medial suture (70–90 long). Genital plates (177.5–236.25 long; 131.25–162.5 wide). Additional measurements: Cx-1 (283–345 long (total); 117–167.5 long (medial)); Cx-3 (348–432.5 wide); anterior venter (245–270 long). Ventral proportions: gnathosomal bay length/width 1.69–2.15; anterior venter/genital field length 1.14–1.44; anterior venter length/genital field width 1.66–1.96; anterior venter/medial suture 2.72–3.71.

Immatures unknown.

Etymology

Marshall (1943) presumably named the specific epithet (ellipsoidalis) after the elongate body of this species, as she wrote, “the body is an ellipse.”

Distribution

Western (Figure 64). T. ellipsoidalis was previously recorded from Martis Creek and Gibbon River, Wyoming (Marshall 1943); and from Torch River, Saskatchewan (Quaglia & Conroy 1984). We expand the range into most of western North America. However, T. ellipsoidalis is not known from the southwest.

Figure 64. 

Torrenticola ellipsoidalis distribution. Red dots indicate material examined. Blue crosses indicate additional previous records (Marshall 1943; Quaglia & Conroy 1984).

Figure 65. 

Torrenticola ellipsoidalis female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 66. 

Torrenticola ellipsoidalis male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Remarks

Torrenticola ellipsoidalis groups other members of the Miniforma Complex with high support and most specimens are 0–3% different in COI sequence. This is higher sequence variability than in many species hypotheses presented herein. However, given the topology in the COI tree (Figure 10, 16) and morphological similarity, it seems apparent that the variability represents a continuum across a large distribution, rather than isolated species. An exception is that a single specimen (DNA#1930) is 2.9–3.6% different from the rest. This specimen is indistinguishable from other specimens and is collected from the same location. We do not find evidence to propose it as a separate species and therefore include it within T. ellipsoidalis.

In all analyses, T. ellipsoidalis groups with T. multiforma and T. regalis, which are greater than 10% different from each other. Based upon overall similarity, body size, and distribution, we place this species within the Ellipsoidalis Identification Group.

Upon examining the types of T. ellipsoidalis and T. rectiforma, all characters for both species overlap with members of only one clade in our analyses. Furthermore, the main character Habeeb (1974) used to differentiate T. rectiforma from T. ellipsoidalis was body size, which is known to be a highly variable character. Therefore, we synonymize T. rectiforma as the junior synonym of T. ellipsoidalis.

Torrenticola elongata Fisher & Dowling, sp. n.

Material examined

HOLOTYPE (♀): from USA, Mississippi, Tishomingo County, Tishomingo State Park, Rock Quarry Branch (34°36'N, 88°11'W), 18 Sep 1991, by IM Smith, IMS910049.

PARATYPES (1 ♀; 2 ♂): Mississippi, USA: 1 ♂ (ALLOTYPE) from Tishomingo County, Tishomingo State Park, Rock Quarry Branch (34°36'43"N, 88°12'4"W), 20 Sep 2009, by IM Smith, IMS090115, DNA 1593 • 1 ♀ and 1 ♂ from Tishomingo County, Tishomingo State Park, Rock Quarry Branch (34°36'43"N, 88°12'4"W), 20 Sep 2009, by IM Smith, IMS090115.

Type deposition

Holotype (♀), allotype (♂) deposited in the CNC; other paratypes (1 ♀; 1 ♂) deposited in the ACUA.

Diagnosis

Torrenticola elongata are similar to species with similar dorsal patterning, such as the Rusetria “4-Plate” group (T. dunni, T. glomerabilis, T. kittatinniana, T. pollani, T. rufoalba and T. shubini), Neoanomala Group (T. interiorensis and T. neoanomala), and T. bondi, T. erectirostra, T. robisoni, T. gorti, T. reduncarostra, T. irapalpa, T. racupalpa, T. skvarlai, and T. arktonyx. T. elongata can be differentiated from all other Torrenticola with similar dorsal patterning by having a more elongate dorsum (length/width ♀ = 1.92–2.08 in T. elongata, 1.17–1.58 in others; ♂ = 1.70–1.70 in T. elongata, 1.20–1.68 in others). Additionally, they can be differentiated from Rusetria 4-Plates and T. skvarlai by having distinct hind coxal margins, they can be differentiated from T. erectirostra and T. robisoni by having a straight, anteriorly-directed rostrum (upturned in others), and they can be differentiated from T. arktonyx by having an unmodified dorsal plate (T. arktonyx has distinctive longitudinal dark markings on the anterior portion of the dorsal plate that fade posteriorly).

Description

Female (Figure 68) (n = 2) (holotype measurements in parentheses when available) with characters of the genus with following specifications.

Dorsum — (540–565 (540) long; 260–295 (260) wide) ovoid and elongate with purple coloration separated into anterior and posterior portions. Anterio-medial platelets (105–105 (105) long; 47.5–50 (47.5) wide). Anterio-lateral platelets (150–157.5 (150) long; 42.5–45 (45) wide) free from dorsal plate. Dgl-4 closer to the edge of the dorsum than to the muscle scars (distance between Dgl-4 180–200 (180)). Dorsal plate proportions: dorsum length/width 1.92–2.08 (1.92); dorsal width/distance between Dgl-4 1.44–1.48 (1.48); anterio-medial platelet length/width 2.10–2.21 (2.21); anterio-lateral platelet length/width 3.33–3.71 (3.33); anterio-lateral/anterio-medial length 1.43–1.50 (1.43).

Gnathosoma — Subcapitulum (285–295 (285) long (ventral); 210–222 (210) long (dorsal); 101.25–102.5 (101.25) tall) colorless. Rostrum (115–117.5 (117.5) long; 32.5–36.25 (32.5) wide) elongate. Chelicerae ((290) long) with curved fangs (45–47.5 (45) long). Subcapitular proportions: ventral length/height 2.81–2.88 (2.81); rostrum length/width 3.24–3.54 (3.24). Pedipalps with tuberculate ventral extensions on femora and on genua. Palpomeres: trochanter (35–37.5 (37.5) long); femur (96.25–100 (96.25) long); genu (55–57.5 (55) long); tibia (61.25–62.5 (61.25) long; 20–21.25 (20) wide); tarsus (15–17.5 (17.5) long). Palpomere proportions: femur/genu 1.74–1.75 (1.75); tibia/femur 0.63–0.64 (0.64); tibia length/width 2.94–3.06 (3.06).

Venter — (690–690 (690) long; 300–350 (300) wide) colorless. Gnathosomal bay (130–135 (130) long; 56.25–60 (56.25) wide). Cxgl-4 subapical. Medial suture (52.5–60 (60) long). Genital plates (142.5–145 (142.5) long; 120–122.5 (120) wide). Additional measurements: Cx-1 (250–270 (250) long (total); 120–125 (120) long (medial)); Cx-3 (240–258 (240) wide); anterior venter (205–207.5 (205) long). Ventral proportions: gnathosomal bay length/width 2.25–2.31 (2.31); anterior venter/genital field length 1.43–1.44 (1.44); anterior venter length/genital field width 1.69–1.71 (1.71); anterior venter/medial suture 3.42–3.95 (3.42).

Male (Figure 69) (n = 2) (allotypic measurements in parentheses when available) with characters of the genus with following specifications.

Dorsum — (450–460 (460) long; 265–270 (270) wide) ovoid and elongate with purple coloration separated into anterior and posterior portions. Anterio-medial platelets (92.5–100 (100) long; 45–47.5 (45) wide). Anterio-lateral platelets (130–142.5 (130) long; 43.75–45 (43.75) wide) free from dorsal plate. Dgl-4 closer to the edge of the dorsum than to the muscle scars (distance between Dgl-4 175–180 (180)). Dorsal plate proportions: dorsum length/width 1.70–1.70 (1.70); dorsal width/distance between Dgl-4 1.50–1.51 (1.50); anterio-medial platelet length/width 1.95–2.22 (2.22); anterio-lateral platelet length/width 2.97–3.17 (2.97); anterio-lateral/anterio-medial length 1.30–1.54 (1.30).

Gnathosoma — Subcapitulum (250–255 (255) long (ventral); 184–187 (184) long (dorsal); 85–87.5 (85) tall) colorless. Rostrum (97.5–102.5 (102.5) long; 30–30 (30) wide) elongate. Chelicerae (243–262 (243) long) with curved fangs (34–42 (42) long). Subcapitular proportions: ventral length/height 2.83–3.00 (3.00); rostrum length/width 3.25–3.42 (3.42). Pedipalps with tuberculate ventral extensions on femora and on genua. Palpomeres: trochanter (31.25–37.5 (31.25) long); femur (83.75–85 (83.75) long); genu (47.5–47.5 (47.5) long); tibia (57.5–60 (57.5) long; 18.75–20 (20) wide); tarsus (15–15 (15) long). Palpomere proportions: femur/genu 1.76–1.79 (1.76); tibia/femur 0.69–0.71 (0.69); tibia length/width 2.88–3.20 (2.88).

Venter — (565–570 (570) long; 325–329 (329) wide) colorless. Gnathosomal bay (105–110 (105) long; 50–55 (55) wide). Cxgl-4 subapical. Medial suture (55–80 (55) long). Genital plates (107.5–113.75 (113.75) long; 92.5–92.5 (92.5) wide). Additional measurements: Cx-1 (207–232 (231) long (total); 82–98 (98) long (medial)); Cx-3 (257–266 (266) wide); anterior venter (215–220 (215) long). Ventral proportions: gnathosomal bay length/width 1.91–2.20 (1.91); anterior venter/genital field length 1.89–2.05 (1.89); anterior venter length/genital field width 2.32–2.38 (2.32); anterior venter/medial suture 2.75–3.91 (3.91).

Immatures unknown.

Etymology

Specific epithet (elongata) refers to elongated bodies of this species, which is more pronounced than in all other North American Torrenticola (elongatus, L. prolonged).

Distribution

Known only from Tishomingo County, Mississippi (Figure 67).

Figure 67. 

Torrenticola elongata sp. n. distribution.

Figure 68. 

Torrenticola elongata sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 69. 

Torrenticola elongata sp. n. male: A dorsal plates, coloration added B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Remarks

Torrenticola elongata groups with other members of the Raptor Complex with high support and specimens are less than 1% different in COI sequence from each other. In all analyses, T. elongata groups with two other species (T. gorti and T. bondi) which are 4% different from each other and have non-overlapping ranges.

Based upon overall similarity, an elongate body, and distribution, we place this species in the Elongata Identification Group.

This species hypothesis is supported by low COI divergence within the species (0–2%) and high divergence between species (3–15%), and by the morphological characters outlined in the diagnosis.

Torrenticola elusiva Fisher & Dowling, sp. n.

Material examined

HOLOTYPE (♀): from Canada, New Brunswick, Charlotte County, Rollingham, Whittier Ridge, Highway 770, 6.6 km east of covered bridge, 3 Oct 2011, by IM Smith, IMS110120, DNA 1857.

Type deposition

Holotype (♀) deposited in the CNC.

Diagnosis

Torrenticola elusiva are similar to other members of the Raptor Group (T. gnoma, T. irapalpa, T. longitibia, T. mjolniri, T. racupalpa, T. raptor, T. danielleae, T. daemon, and T. ivyae) in having round bodies; Dgl-4 close to muscles scars; long, thin subcapitular rostra; and long, thin pedipalp tibiae. T. elusiva can be differentiated from T. racupalpa by having a stockier subcapitulum (ventral length/height = 2.39 in T. elusiva, 2.48–2.73 in T. racupalpa); and by dorsal pattern. T. elusiva can be differentiated from T. irapalpa and T. daemon by having Dgl-4 closer to the muscle scars (dorsal width/distance between Dgl-4 = 2.5 in T. elusiva, 1.59–2.09 in others); a more elongate rostrum (length/width = 3.65 in T. elusiva, 2.66–3.39 in others); and by dorsal coloration and pattern. T. elusiva can be differentiated from T. gnoma by being larger (dorsum length = 645 in T. elusiva, 540–595 in T. gnoma); having a more elongate rostrum (length/width = 3.65 in T. elusiva, 2.74–3.13 in T. gnoma); and dorsal coloration. T. elusiva can be differentiated from T. mjolniri and T. ivyae by having stockier pedipalp tibiae (length/width = 4.42 in T. elusiva, 5.00–6.00 in others); and a stockier rostrum (length/width = 3.65 in T. elusiva, 3.81–4.32 in others). T. elusiva can be differentiated from T. raptor by having Dgl-4 closer to the muscle scars (dorsal width/distance between Dgl-4 = 2.50 in T. elusiva, 1.8–2.02 in T. raptor); shorter anterior venter (163.75 in T. elusiva, 205–240 in T. raptor); and stockier pedipalp tibiae (length/width = 4.42 in T. elusiva, 6–7.54 in T. raptor). T. elusiva can be differentiated from T. danielleae by having Dgl-4 closer to the muscle scars (dorsal width/distance between Dgl-4 = 2.5 in T. elusiva, 1.57–1.70 in T. danielleae) and by dorsal coloration. T. elusiva cannot be confidently differentiated from T. longitibia because T. elusiva is only known from a single female and T. longitibia is only known from two males; however, T. elusiva is only known from Charlotte County, New Brunswick, whereas T. longitibia is only known from Monroe County, Tennessee. Additionally, two character systems that vary minimally between sexes are rostrum and pedipalp tibiae proportions, which do differ between T. elusiva and T. longitibia as follows: pedipalp tibia stockier (4.42 in T. elusiva, 5.5–5.5 in T. longitibia) and rostrum stockier (3.65 in T. elusiva, 4.15–4.23 in T. longitibia).

Description

Female (Figure 71) (n = 1) (holotype only) with characters of the genus with following specifications.

Dorsum — (645 long; 500 wide) circular with bluish-purple coloration posteriorly with a broad anterior extension reaching the anterior edge of the dorsal plate. Anterio-medial platelets (152.5 long; 70 wide). Anterio-lateral platelets (182.5 long; 87.5 wide) free from dorsal plate. Dgl-4 much closer to the muscle scars than to edge of dorsum (distance between Dgl-4 200). Dorsal plate proportions: dorsum length/width 1.29; dorsal width/distance between Dgl-4 2.50; anterio-medial platelet length/width 2.18; anterio-lateral platelet length/width 2.09; anterio-lateral/anterio-medial length 1.20.

Gnathosoma — Subcapitulum (340 long (ventral); 259 long (dorsal); 142.5 tall) colorless. Rostrum (155 long; 42.5 wide) elongate. Chelicerae (333 long) with curved fangs (59 long). Subcapitular proportions: ventral length/height 2.39; rostrum length/width 3.65. Pedipalps elongate (especially tibiae) with tuberculate ventral extensions with dentate tip on femora and tuberculate ventral extensions on genua. Palpomeres: trochanter (48.75 long); femur (132.5 long); genu (72.5 long); tibia (105 long; 23.75 wide); tarsus (17.5 long). Palpomere proportions: femur/genu 1.83; tibia/femur 0.79; tibia length/width 4.42.

Venter — (730 long; 554 wide) colorless. Gnathosomal bay (176.25 long; 87.5 wide). Cxgl-4 subapical. Medial suture (17.5 long). Genital plates (167.5 long; 150 wide). Additional measurements: Cx-1 (288 long (total); 115 long (medial)); Cx-3 (384 wide); anterior venter (163.75 long). Ventral proportions: gnathosomal bay length/width 2.01; anterior venter/genital field length 0.98; anterior venter length/genital field width 1.09; anterior venter/medial suture 9.36.

Male unknown.

Immatures unknown.

Etymology

Specific epithet (elusiva) refers to the fact that we were only able to find a single specimen of this species, despite extensive searching among the abundant samples taken from the type locality in New Brunswick.

Distribution

Known only from Charlotte County, New Brunswick (Figure 70).

Figure 70. 

Torrenticola elusiva sp. n. distribution.

Figure 71. 

Torrenticola elusiva sp. n. female: A dorsal plates, coloration added B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Remarks

Torrenticola elusiva groups with other members of the Raptor Complex in all analyses with high support. Only one specimen could be acquired for use in our analyses, so differences in COI sequence across specimens could not be investigated, but this single specimen was greater than 4% different in COI sequence from sister species. Furthermore, this species is known from only a single female, so morphological variation could not be investigated. However, this specimen was different enough in terms of morphology and sequence to warrant a separate description. We place this species within the Raptor Identification group based upon similarity with those species.

This species hypothesis is supported by high divergence between species (3–15%), and by the morphological characters outlined in the diagnosis.

Torrenticola erectirostra Fisher & Dowling, sp. n.

Material examined

HOLOTYPE (♀): from Canada, New Brunswick, York County, Stanley, Nashwaak River, Stanley Municipal Park, 19 Jun 2012, by IM Smith, IMS120031, DNA 2962.

PARATYPES (9 ♀; 9 ♂): New Brunswick, Canada: 3 ♂ from Charlotte County, Rollingham, Digdeguash River, beside Highway 770 at covered bridge, 30 Jun 1989, by IM Smith, IMS890053 • 1 ♀ and 2 ♂ from Charlotte County, Digdeguash River, beside Sorrel Ridge Road west of Whittier Road, 10 Jun 2012, by IM Smith, IMS120015 • 4 ♀ and 4 ♂ from York County, Magaguadavic River, beside Highway 3 just east of Thomaston Corners, 1 Jul 1989, by IM Smith, IMS890055A • 1 ♂ (ALLOTYPE) from York County, Stanley, Davis Brook, beside Highway 3, 3.5 km south of Highway 4 at Thomaston Corner, 11 Jun 2012, by IM Smith, IMS120017, DNA 2964 • 2 ♀ and 1 ♂ from New Brunswick, York County, Stanley, Davis Brook, beside Highway 3, 3.5 km south of Highway 4 at Thomaston Corner, 11 Jun 2012, by IM Smith, IMS120017 • Maine, USA: 1 ♀ from Aroostook County, Ashland, beside Route 11, Aroostook River (46°38'N 68°24'W), 4 Jul 1989, by IM Smith, IMS890067 • New York, USA: 1 ♂ from Cayuga County, Dutch Hollow Brook, beside Route 38A at Niles, 22 Jul 1990, by IM Smith, IMS900113A • Virginia, USA: 1 ♀ from Amherst County, beside Blue Ridge, Otter Creek (37°36'57"N, 79°19'27"W), 7 Sep 2007, by IM Smith, IMS070056A.

Type deposition

Holotype (♀), allotype (♂), and most paratypes (6 ♀; 4 ♂) deposited in the CNC; other paratypes (3 ♀; 4 ♂) deposited in the ACUA.

Diagnosis

Torrenticola erectirostra are similar to other members of the Erectirostra Group (T. karambita and T. robisoni), species with similar dorsal patterning, such as Rusetria “4-Plate” group (T. dunni, T. glomerabilis, T. kittatinniana, T. pollani, T. rufoalba and T. shubini), Elongata Group (T. gorti and T. elongata), Neoanomala Group (T. interiorensis and T. neoanomala), T. bondi, T. irapalpa, T. racupalpa, and T. skvarlai. They can be differentiated from all other Torrenticola, except T. karambita and T. robisoni, by having a dentate, upturned rostrum that is wide when viewed ventrally. T. erectirostra can be differentiated from T. karambita by having dorsal coloration (T. karambita is colorless) and a slightly more elongate rostrum (length/width ♀ = 1.72–1.91 in T. erectirostra, 1.57–1.62 in T. karambita; ♂ = 2.0–2.2 in T. erectirostra, 1.6–1.95 in T. karambita). T. erectirostra can be differentiated from T. robisoni by having less elongate anterio-lateral platelets (length/width ♀ = 2.52–2.69 in T. erectirostra, 2.96–3.00 in T. robisoni) and by being distributed in the Appalachians, while T. robisoni is in the Interior Highlands.

Description

Female (Figure 73) (n = 5) (holotype measurements in parentheses when available) with characters of the genus with following specifications.

Dorsum— (690–750 (735) long; 480–510 (510) wide) ovoid with bluish-purple or purple coloration separated into anterior and posterior portions with orange medially. Anterio-medial platelets (150–165 (162.5) long; 62.5–75 (75) wide). Anterio-lateral platelets (195–225 (220) long; 77.5–83.75 (82.5) wide) free from dorsal plate. Dgl-4 closer to the edge of the dorsum than to the muscle scars (distance between Dgl-4 325–370 (370)). Dorsal plate proportions: dorsum length/width 1.41–1.47 (1.44); dorsal width/distance between Dgl-4 1.38–1.51 (1.38); anterio-medial platelet length/width 2.17–2.48 (2.17); anterio-lateral platelet length/width 2.52–2.69 (2.67); anterio-lateral/anterio-medial length 1.26–1.36 (1.35).

Gnathosoma — Subcapitulum (315–350 (350) long (ventral); 225–247.5 (247.5) long (dorsal); 130–140 (130) tall) colorless. Rostrum (105–125 125) long; 55–72.5 (72.5) wide) wide and upturned with dentation. Chelicerae (320–345 (345) long) with curved fangs (45–55 (45) long). Subcapitular proportions: ventral length/height 2.42–2.69 (2.69); rostrum length/width 1.72–1.91 (1.72). Pedipalps short and stocky (especially tibiae) with tuberculate ventral extensions on femora and genua. Palpomeres: trochanter (45–55 (50) long); femur (100–107.5 (102.5) long); genu (57.5–67.5 (57.5) long); tibia (50–65 (62.5) long; 27.5–30 (28.75) wide); tarsus (17.5–20 (20) long). Palpomere proportions: femur/genu 1.48–1.78 (1.78); tibia/femur 0.50–0.61 (0.61); tibia length/width 1.82–2.17 (2.17).

Venter — (860–920 (900) long; 580–650 (650) wide) colorless. Gnathosomal bay (210–220 long; 105–150 wide). Cxgl-4 far from apex. Medial suture (17.5–27.5 (22.5) long). Genital plates (187.5–202.5 (202.5) long; 162.5–180 (180) wide). Additional measurements: Cx-1 (330–360 (350) long (total); 140–160 (160) long (medial)); Cx-3 (410–460 (460) wide); anterior venter (192.5–220 (220) long). Ventral proportions: gnathosomal bay length/width 1.47–2.00; anterior venter/genital field length 0.98–1.09 (1.09); anterior venter length/genital field width 1.13–1.25 (1.22); anterior venter/medial suture 7.09–12.14 (9.78).

Male (Figure 74) (n = 5) (allotypic measurements in parentheses when available) with characters of the genus with following specifications.

Dorsum — (580–640 (620) long; 400–430 (400) wide) ovoid with bluish-purple or purple coloration separated into anterior and posterior portions with orange medially. Anterio-medial platelets (130–150 (138.75) long; 52.5–58.75 (58.75) wide). Anterio-lateral platelets (187.5–205 (205) long; 62.5–70 (68.75) wide) free from dorsal plate. Dgl-4 closer to the edge of the dorsum than to the muscle scars (distance between Dgl-4 275–305 (300)). Dorsal plate proportions: dorsum length/width 1.41–1.55 (1.55); dorsal width/distance between Dgl-4 1.33–1.45 (1.33); anterio-medial platelet length/width 2.26–2.73 (2.36); anterio-lateral platelet length/width 2.78–3.20 (2.98); anterio-lateral/anterio-medial length 1.37–1.48 (1.48).

Gnathosoma — Subcapitulum (270–292.5 (285) long (ventral); 175–215 (197.5) long (dorsal); 96.25–110 (105) tall) colorless. Rostrum (90–107.5 (98.75) long; 45–50 (46.25) wide) wide and upturned with dentation. Chelicerae (265–285 (265) long) with curved fangs (45–50 (50) long). Subcapitular proportions: ventral length/height 2.45–2.86 (2.71); rostrum length/width 2.00–2.17 (2.14). Pedipalps short and stocky (especially tibiae) with tuberculate ventral extensions on femora and genua. Palpomeres: trochanter (42.5–47.5 (42.5) long); femur (80–91.25 (87.5) long); genu (55–57.5 (55) long); tibia (50–57.5 (50) long; 23.75–27.5 (23.75) wide); tarsus (15–17.5 (15) long). Palpomere proportions: femur/genu 1.39–1.59 (1.59); tibia/femur 0.57–0.63 (0.57); tibia length/width 2.00–2.11 (2.11).

Venter — (720–780 (750) long; 470–495 (470) wide) colorless. Gnathosomal bay (167.5–177.5 (172.5) long; 100–105 (105) wide). Cxgl-4 far from apex. Medial suture (75–82.5 (75) long). Genital plates (152.5–165 (157.5) long; 112.5–125 (112.5) wide). Additional measurements: Cx-1 (290–330 (310) long (total); 125–150 (140) long (medial)); Cx-3 (360–390 (360) wide); anterior venter (232.5–250 (250) long). Ventral proportions: gnathosomal bay length/width 1.60–1.78 (1.64); anterior venter/genital field length 1.47–1.64 (1.59); anterior venter length/genital field width 1.94–2.22 (2.22); anterior venter/medial suture 2.94–3.33 (3.33).

Immatures unknown.

Etymology

Specific epithet (erectirostra) refers to the upturned rostrum characteristic of members of the Erectirostra Group (erectus, raised up; rostrum, L. snout).

Distribution

Appalachians (Figure 72).

Figure 72. 

Torrenticola erectirostra sp. n. distribution.

Figure 73. 

Torrenticola erectirostra sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 74. 

Torrenticola erectirostra sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Remarks

Torrenticola erectirostra groups with other members of the Raptor Complex with high support and specimens are 2.7% different from each other in COI sequence. This variation in COI is higher than in most species hypotheses proposed herein, especially since those specimens were form the same region (New Brunswick). However, we could not find morphological differences that corresponded to clades in our analysis, and we were only able to examine four sequence, therefore, we consider these specimens to represent a single species hypothesis.

This species groups with T. karambita and T. robisoni to form the Erectirostra Identification Group, which can be readily identified by the shape of the rostrum.

This species hypothesis is supported by biogeography, molecular divergence (although COI variation is greater than most hypotheses herein), and by the morphological characters outlined in the diagnosis.

Torrenticola feminellai Fisher & Dowling, sp. n.

Material examined

HOLOTYPE (♀): from USA, Georgia, Chattooga County, Cloudland; beside Rt. 48 just east of Alabama state line, (34°31'31"N, 85°30'30"W), 29 September 2005, by IM Smith, IMS050110A.

PARATYPES (4 ♀; 6 ♂): Georgia, USA : 2 ♀ and 4 ♂ from Chattooga County, Cloudland; beside Rt. 48 just east of Alabama State line, (34°31'31"N, 85°30'30"W), 28 September 1992, by IM Smith, IMS920056A • 1 ♂ (ALLOTYPE) from Chattooga County, Cloudland; beside Rt. 48 just east of Alabama state line, (34°31'31"N, 85°30'30"W), 29 September 2005, by IM Smith, IMS050110A • 2 ♀ and 1 ♂ from Chattooga County, Cloudland; beside Rt. 48 just east of Alabama state line, (34°31'31"N, 85°30'30"W), 29 September 2005, by IM Smith, IMS050110A.

Type deposition

Holotype (♀), allotype (♂), and some paratypes (2 ♀; 3 ♂) deposited in the CNC; other paratypes (2 ♀; 2 ♂) deposited in the ACUA.

Diagnosis

Torrenticola feminellai are similar to other members of the Rusetria “Eastern 2-Plates” group (T. biscutella, T. caerulea, T. delicatexa, T. indistincta, T. malarkeyorum, T. pendula, T. sellersorum, T. tysoni, T. ululata, T. whitneyae, and T. microbiscutella) in having anterio-lateral platelets fused to the dorsal plate, having dorsal coloration separated into anterior and posterior portions (except T. ululata and T. indistincta), and being distributed in the east. T. feminellai can be differentiated from all other Eastern 2-Plates by having a more elongate rostrum (length/width ♀ = 3.05–3.38 in T. feminellai, 2.33–3.00 in others; ♂ = 3.14–3.38 in T. feminellai, 2.50–3.05 in others), except T. tysoni (3.06–3.50). Additionally, T. feminellai can be differentiated from all other Eastern 2-Plates by having a distinct dorsal pattern.

Description

Female (Figure 76) (n = 5) (holotype measurements in parentheses when available) with characters of the genus with following specifications.

Dorsum — (590–690 (640) long; 470–540 (500) wide) circular with reddish-purple coloration in the shape of an hourglass. Anterio-medial platelets (137.5–150 (150) long; 52.5–55 (55) wide). Anterio-lateral platelets (185–202.5 (202.5) long; 75–80 (75) wide) partially fused to dorsal plate. Dgl-4 approximately halfway between the edge of the dorsum and the muscle scars (distance between Dgl-4 290–310 (290)). Dorsal plate proportions: dorsum length/width 1.24–1.33 (1.28); dorsal width/distance between Dgl-4 1.59–1.74 (1.72); anterio-medial platelet length/width 2.62–2.76 (2.73); anterio-lateral platelet length/width 2.34–2.70 (2.70); anterio-lateral/anterio-medial length 1.25–1.36 (1.35).

Gnathosoma — Subcapitulum (320–357.5 (352.5) long (ventral); 245–275 (267.5) long (dorsal); 140–160 (160) tall) colorless. Rostrum (135–157.5 (145) long; 40–47.5 (47.5) wide). Chelicerae (335–375 (375) long) with curved fangs (65–75 (70) long). Subcapitular proportions: ventral length/height 2.17–2.30 (2.20); rostrum length/width 3.05–3.38 (3.05). Pedipalps with tuberculate ventral extensions on femora and genua. Palpomeres: trochanter (46.25–52.5 (52.5) long); femur (117.5–140 (140) long); genu (65–77.5 (75) long); tibia (90–100 (100) long; 22.5–25 (25) wide); tarsus (17.5–20 (20) long). Palpomere proportions: femur/genu 1.73–1.87 (1.87); tibia/femur 0.71–0.77 (0.71); tibia length/width 3.85–4.11 (4.00).

Venter — (670–760 (700) long; 550–690 (550) wide) colorless. Gnathosomal bay (177.5–195 (195) long; 80–115 (80) wide). Cxgl-4 subapical. Medial suture absent. Genital plates (185–195 (195) long; 167.5–180 (170) wide). Additional measurements: Cx-1 (310–320 (320) long (total); 120–140 (125) long (medial)); Cx-3 (340–410 (360) wide); anterior venter (130–140 (130) long). Ventral proportions: gnathosomal bay length/width 1.61–2.44 (2.44); anterior venter/genital field length 0.67–0.76 (0.67); anterior venter length/genital field width 0.74–0.82 (0.76).

Male (Figure 77) (n = 5) (allotypic measurements in parentheses when available) with characters of the genus with following specifications.

Dorsum — (520–565 (545) long; 370–410 (390) wide) circular with reddish-purple coloration in the shape of an hourglass. Anterio-medial platelets (115–126.25 (122.5) long; 42.5–50 (45) wide). Anterio-lateral platelets (150–180 (155) long; 55–67.5 (60) wide) partially fused to dorsal plate. Dgl-4 approximately halfway between the edge of the dorsum and the muscle scars (distance between Dgl-4 215–250 (245)). Dorsal plate proportions: dorsum length/width 1.36–1.42 (1.40); dorsal width/distance between Dgl-4 1.54–1.72 (1.59); anterio-medial platelet length/width 2.42–2.82 (2.72); anterio-lateral platelet length/width 2.58–2.77 (2.58); anterio-lateral/anterio-medial length 1.27–1.43 (1.27).

Gnathosoma — Subcapitulum (270–300 (290) long (ventral); 200–227.5 (215) long (dorsal); 105–120 (115) tall) colorless. Rostrum (110–120 (120) long; 32.5–37.5 (37.5) wide). Chelicerae (270–310 (295) long) with curved fangs (50–60 (55) long). Subcapitular proportions: ventral length/height 2.50–2.59 (2.52); rostrum length/width 3.14–3.38 (3.20). Pedipalps with tuberculate ventral extensions on femora and genua. Palpomeres: trochanter (35–45 (42.5) long); femur (95–112.5 (105) long); genu (60–65 (62.5) long); tibia (77.5–86.25 (85) long; 20–23.75 (21.25) wide); tarsus (15–17.5 (17.5) long). Palpomere proportions: femur/genu 1.58–1.75 (1.68); tibia/femur 0.76–0.82 (0.81); tibia length/width 3.63–4.00 (4.00).

Venter — (610–685 (640) long; 415–470 (470) colorless. Gnathosomal bay (135–160 (155) long; 62.5–70 (62.5) wide). Cxgl-4 subapical. Medial suture (65–75 (70) long). Genital plates (130–140 (140) long; 115–125 (120) wide). Additional measurements: Cx-1 (245–290 (270) long (total); 105–130 (120) long (medial)); Cx-3 (285–315 (315) wide); anterior venter (190–205 (195) long). Ventral proportions: gnathosomal bay length/width 2.00–2.48 (2.48); anterior venter/genital field length 1.39–1.49 (1.39); anterior venter length/genital field width 1.58–1.72 (1.63); anterior venter/medial suture 2.60–2.93 (2.79).

Immatures unknown.

Etymology

Specific epithet (feminellai) named in honor of Jack Feminella, professor of biology at Auburn University, who believed in me (JRF) enough to employ me as a lab technician in stream ecology, write a winning recommendation letter for graduate school, and was the first to teach me how to conduct self-directed research.

Distribution

Southern Appalachians, Georgia (Figure 75).

Figure 75. 

Torrenticola feminellai sp. n. distribution.

Figure 76. 

Torrenticola feminellai sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 77. 

Torrenticola feminellai sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Remarks

Unfortunately, we were unable to acquire fresh material of Torrenticola feminellai and therefore this species is not included in our phylogenetic analyses. However, we were able to examine morphology with material preserved in GAW. The overall similarity, distribution in the east, and fusion of the dorso-lateral platelets to the dorsal plate, are consistent with placing this species in the Rusetria Complex and the Eastern 2-Plate Identification Group.

Torrenticola flangipalpa Fisher & Dowling, sp. n.

Material examined

HOLOTYPE (♀): from USA, Alabama, Lauderdale County, Natchez Trace Parkway, (34°56'31"N, 87°49'41"W), 27 Sep 2010, by IM Smith, IMS100162, DNA 1310.

PARATYPES (5 ♀; 7 ♂): Alabama, USA: 3 ♂ from Lauderdale County, Natchez Trace Parkway, (34°56'31"N, 87°49'41"W), 24 Sep 2009, by IM Smith, IMS090121 • 1 ♂ (ALLOTYPE) from Lauderdale County, Natchez Trace Parkway, (34°56'31"N, 87°49'41"W), 27 Sep 2010, by IM Smith, IMS100162, DNA 1309 • 1 ♀ and 2 ♂ from Lauderdale County, Natchez Trace Parkway, (34°56'31"N, 87°49'41"W), 27 Sep 2010, by IM Smith, IMS100162 • Arkansas, USA: 1 ♀ from Carroll County, beside Route 62, Kings River, 21 Jul 1960, by DR Cook, DRC600026 • Kentucky, USA: 1 ♀ from Larue County, beside Route 31E, 4 kilometers south of Route 61 at Hodgenville (37°35'N, 85°42'W), 28 May 1992, by IM Smith, IMS920014 • North Carolina, USA: 1 ♀ and 1 ♂ from Haywood County, Great Smokey Mountains National Park, Big Creek (35°45'3.92"N, 83°6'31.67"W), 15 Sep 2009, by AJ Radwell, AJR090008A • Tennessee, USA: 1 ♀ from Sevier County, Great Smokey Mountains National Park, Sugarlands Nature Trail (35°40'47"N, 83°31'52"W), 7 Sep 2009, by IM Smith, IMS090101.

Type deposition

Holotype (♀), allotype (♂), and most paratypes (3 ♀; 4 ♂) deposited in the CNC; other paratypes (2 ♀; 2 ♂) deposited in the ACUA.

Diagnosis

Torrenticola flangipalpa are similar to other members of the Nigroalba Group (T. nigroalba, T. solisorta, and T. dentirostra) in being small, slightly elongate, having purple dorsal coloration restricted posteriorly, and having distinct yet poorly-defined hind coxal margins (can be indistinct, at least in males). T. flangipalpa are best differentiated from other members of the Nigroalba Group by having a flange-like, anteriorly-directed pedipalp femoral extension (this extension is tuberculate in other members of the Nigroalba Group). Additionally, T. flangipalpa can be differentiated from T. nigroalba and T. solisorta by having a longer anterior venter (235–265 in T. flangipalpa; 192–225 in others) and stockier pedipalpal tibiae (length/width ♀ = 4.79–5.0 in T. flangipalpa, 5.38–5.83 in others; length/width ♂ = 4.4–4.86 in T. flangipalpa, 5.08–5.33 in others). T. flangipalpa can be differentiated from T. dentirostra by having a smooth rostrum (T. dentirostra has a dentate bump midway on the dorsal edge of the rostrum). Other Torrenticola with purple dorsal coloration restricted posteriorly, such as T. tahoei and T. oregonensis are larger (dorsum length ♀ = 530–565 in T. flangipalpa, 600–810 in others; ♂ = 480–510 in T. flangipalpa, 560–820 in others) and distributed in the west (T. flangipalpa is only known from Alabama and Tennessee).

Description

Female (Figure 79) (n = 5) (holotype measurements in parentheses when available) with characters of the genus with following specifications.

Dorsum — (530–580 (545) long; 365–425 (380) wide) ovoid with purple or bluish-purple coloration restricted posteriorly. Anterio-medial platelets (112.5130 (127.5) long; 47.5–57.5 (47.5) wide). Anterio-lateral platelets (157.5–175 (172.5) long; 55–62.5 (55) wide) free from dorsal plate. Dgl-4 closer to the edge of the dorsum than to the muscle scars (distance between Dgl-4 255–295 (255)). Dorsal plate proportions: dorsum length/width 1.33–1.45 (1.43); dorsal width/distance between Dgl-4 1.39–1.49 (1.49); anterio-medial platelet length/width 2.25–2.74 (2.68); anterio-lateral platelet length/width 2.86–3.14 (3.14); anterio-lateral/anterio-medial length 1.29–1.40 (1.35).

Gnathosoma — Subcapitulum (305–330 (307.5) long (ventral); 225–255 (225) long (dorsal); 90–95 (90) tall) elongate and colorless. Rostrum (112.5–130 (117.5) long; 37.5–42.5 (37.5) wide) elongate. Chelicerae (285–320 (286) long) with curved fangs (43–50 (44) long). Subcapitular proportions: ventral length/height 3.39–3.47 (3.42); rostrum length/width 2.82–3.06 (3.00). Pedipalps elongate (especially tibiae) with broad, dentate, anteriorly-directed flange on femora and with variable, dentate flange-like extension on genua. Palpomeres: trochanter (32.5–35 (35) long); femur (93.75–102.5 (93.75) long); genu (55–62.5 (55) long); tibia (80–90 (83.75) long; 16.25–18.75 (17.5) wide); tarsus (15–17.5 (15) long). Palpomere proportions: femur/genu 1.60–1.70 (1.70); tibia/femur 0.82–0.90 (0.89); tibia length/width 4.79–5.00 (4.79).

Venter — (680–750 (680) long; 430–495 (436) wide) with faint purple or bluish-purple coloration. Gnathosomal bay (112.5–140 (120) long; 67.5–75 (67.5) wide). Cxgl-4 far from apex. Medial suture (67.5–80 (67.5) long). Genital plates (145–160 (151.25) long; 125–132.5 (125) wide). Additional measurements: Cx-1 (266–310 (266) long (total); 150–170 (156) long (medial)); Cx-3 (278–321 (278) wide); anterior venter (235–255 (245) long). Ventral proportions: gnathosomal bay length/width 1.50–2.00 (1.78); anterior venter/genital field length 1.55–1.76 (1.62); anterior venter length/genital field width 1.86–2.04 (1.96); anterior venter/medial suture 3.06–3.78 (3.63).

Male (Figure 80) (n = 6) (allotypic measurements in parentheses when available) with characters of the genus with following specifications.

Dorsum — (480–510 (480) long; 330–370 (330) wide) ovoid with purple or bluish-purple coloration restricted posteriorly. Anterio-medial platelets (112.5–122.5 (115) long; 41.25–47.5 (41.25) wide). Anterio-lateral platelets (152.5–162.5 (152.5) long; 50–53.75 (50) wide) free from dorsal plate. Dgl-4 closer to the edge of the dorsum than to the muscle scars (distance between Dgl-4 235–260 (235)). Dorsal plate proportions: dorsum length/width 1.38–1.45 (1.45); dorsal width/distance between Dgl-4 1.39–1.44 (1.40); anterio-medial platelet length/width 2.42–2.79 (2.79); anterio-lateral platelet length/width 3.02–3.10 (3.05); anterio-lateral/anterio-medial length 1.33–1.44 (1.33).

Gnathosoma — Subcapitulum (272.5–290 (272.5) long (ventral); 200–209 (200) long (dorsal); 75–87.5 (75) tall) elongate and colorless. Rostrum (102.5–107.5 (105) long; 32.5–35 (35) wide) elongate. Chelicerae (250–272 (260) long) with curved fangs (39–62 (40) long). Subcapitular proportions: ventral length/height 3.31–3.68 (3.63); rostrum length/width 2.93–3.31 (3.00). Pedipalps elongate (especially tibiae) with broad, dentate, anteriorly-directed flange on femora and with variable, dentate flange-like extension on genua. Palpomeres: trochanter (27.5–31.25 (27.5) long); femur (85–90 (87.5) long); genu 47.5–57.5 (55) long); tibia (77.5–85 (80) long; 17.5–18.75 (17.5) wide); tarsus (12.5–15 (12.5) long). Palpomere proportions: femur/genu 1.52–1.79 (1.59); tibia/femur 0.91–0.94 (0.91); tibia length/width 4.40–4.86 (4.57).

Venter — (600–640 (600) long; 356–420 (380) wide) with faint purple or bluish-purple coloration. Gnathosomal bay (100–112.5 (105) long; 65–72.5 (65) wide). Cxgl-4 far from apex. Medial suture (82.5–107.5 (95) long). Genital plates (122.5–127.5 (122.5) long; 92.5–100 (95) wide). Additional measurements: Cx-1 (240–258 (240) long (total); 122–160 (135) long (medial)); Cx-3 (251–291 (265) wide); anterior venter (245–265 (245) long). Ventral proportions: gnathosomal bay length/width 1.48–1.62 (1.62); anterior venter/genital field length 1.98–2.12 (2.00); anterior venter length/genital field width 2.55–2.68 (2.58); anterior venter/medial suture 2.42–3.00 (2.58).

Immatures unknown.

Etymology

Specific epithet (flangipalpa) refers the expanded (i.e., flanged) pedipalp femoral tubercle (flange, English; palpus, L. hand, feeler), which distinguishes this species from other members of the Nigroalba Group.

Distribution

Southeastern (Figure 78).

Figure 78. 

Torrenticola flangipalpa sp. n. distribution.

Figure 79. 

Torrenticola flangipalpa sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 80. 

Torrenticola flangipalpa sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Remarks

Torrenticola flangipalpa groups with other members of the Raptor Complex with high support in all analyses and specimens are less than 1% different in COI sequence from each other. In all analyses, T. flangipalpa groups with two other morphologically similar species: T. nigroalba and T. solisorta. These three species are greater than 12% different from each other in COI sequence.

That clade of three species corresponds to an identification group, Nigroalba Group, the members of which are easily differentiated by their size, coloration, long medial suture in females, and overall appearance.

This species hypothesis is supported by low COI divergence within the species (0–2%) and high divergence between species (3–15%), and by the morphological characters outlined in the diagnosis.

Torrenticola folkertsae Fisher & Dowling, sp. n.

Material examined

HOLOTYPE (♀): from USA, New Hampshire, Coos County, picnic area beside Rt. 110 ca. 1 km east of Stark, (44°36'36"N, 71°24'24"W), 5 July 1989, by IM Smith, IMS890071

PARATYPES (4 ♀; 4 ♂): 1 ♂ (ALLOTYPE) from Coos County, New Hampshire picnic area beside Rt. 110 ca. 1 km east of Stark, (44°36'36"N, 71°24'24"W), 5 July 1989, by IM Smith, IMS890071 • 4 ♀ and 4 ♂ from Coos County, picnic area beside Rt. 110 ca. 1 km east of Stark, (44°36'36"N, 71°24'24"W), 5 July 1989, by IM Smith, IMS890071.

Type deposition

Holotype (♀), allotype (♂), and some paratypes (2 ♀; 2 ♂) deposited in the CNC; other paratypes (2 ♀; 1 ♂) deposited in the ACUA.

Diagnosis

Torrenticola folkertsae are similar to other members of the Partial 2-Plate Group (T. magnexa, T. priapus, and T. pulchra) in having anterio-lateral platelets partially fused to the dorsal plate and being distributed in the east. T. folkertsae can be differentiated from T. magnexa and T. pulchra by having more elongate pedipalpal tibiae (length/width = 4.05–4.83 in T. folkertsae, 2.61–4.00 in others) and by dorsal coloration and pattern. T. folkertsae can be differentiated from T. priapus by having more elongate pedipalpal tibiae (length/width ♀ = 4.5–4.83 in T. folkertsae, 3.9–4.22 in T. priapus ♂ = 4.05–4.33 in T. folkertsae, 3.5–3.78 in T. priapus) and a less elongate rostrum (length/width = 2.55–3.00 in T. folkertsae, 3.17–3.39 in T. priapus).

Description

Female (Figure 82) (n = 5) (holotype measurements in parentheses when available) with characters of the genus with following specifications.

Dorsum — (600–720 (720) long; 505–600 (580) wide) ovoid with faint reddish-purple medially. Anterio-medial platelets (140–167.5 (156.25) long; 57.5–70 (65) wide). Anterio-lateral platelets (180–205 (185) long; 85–95 (85) wide) partially fused to dorsal plate. Dgl-4 closer to the edge of the dorsum than to the muscle scars (distance between Dgl-4 340–380 (380)). Dorsal plate proportions: dorsum length/width 1.11–1.24 (1.24); dorsal width/distance between Dgl-4 1.40–1.75 (1.53); anterio-medial platelet length/width 2.32–2.50 (2.40); anterio-lateral platelet length/width 2.12–2.31 (2.18); anterio-lateral/anterio-medial length 1.12–1.41 (1.18).

Gnathosoma — Subcapitulum (310–345 (345) long (ventral); 240–270 (270) long (dorsal); 145–162.5 (157.5) tall) colorless. Rostrum (135–150 (140) long; 50–55 (55) wide). Chelicerae (320–415 (415) long) with curved fangs (65–70 (70) long). Subcapitular proportions: ventral length/height 2.07–2.19 (2.19); rostrum length/width 2.55–2.86 (2.55). Pedipalps with tuberculate ventral extensions on femora and genua. Palpomeres: trochanter (48.75–52.5 (52.5) long); femur (130–145 (145) long); genu (77.5–87.5 (87.5) long); tibia (108.75–120 (118.75) long; 22.5–26.25 (26.25) wide); tarsus (17.5–22.5 (22.5) long). Palpomere proportions: femur/genu 1.65–1.68 (1.66); tibia/femur 0.82–0.87 (0.82); tibia length/width 4.50–4.83 (4.52).

Venter — (680–880 (860) long; 560–675 (650) wide) colorless. Gnathosomal bay (167.5–185 (182.5) long; 80–95 (95) wide). Cxgl-4 far from apex. Medial suture (15–20 (20) long). Genital plates (175–197.5 (197.5) long; 167.5–175 (167.5) wide). Additional measurements: Cx-1 (300–340 (330) long (total); 127.5–157.5 (145) long (medial)); Cx-3 (380–400 (400) wide); anterior venter (155–180 (180) long). Ventral proportions: gnathosomal bay length/width 1.92–2.13 (1.92); anterior venter/genital field length 0.79–1.00 (0.91); anterior venter length/genital field width 0.91–1.07 (1.07); anterior venter/medial suture 8.38–12.00 (9.00).

Male (Figure 83) (n = 5) (allotypic measurements in parentheses when available) with characters of the genus with following specifications.

Dorsum — (535–580 (570) long; 420–450 (445) wide) ovoid with faint reddish-purple medially. Anterio-medial platelets (125–135 (135) long; 55–65 (55) wide). Anterio-lateral platelets (180–187.5 (180) long; 62.5–77.5 (77.5) wide) partially fused to dorsal plate. Dgl-4 closer to the edge of the dorsum than to the muscle scars (distance between Dgl-4 280–310 (310)). Dorsal plate proportions: dorsum length/width 1.27–1.29 (1.28); dorsal width/distance between Dgl-4 1.44–1.54 (1.44); anterio-medial platelet length/width 2.04–2.45 (2.45); anterio-lateral platelet length/width 2.32–2.88 (2.32); anterio-lateral/anterio-medial length 1.33–1.48 (1.33).

Gnathosoma — Subcapitulum (265–292.5 (292.5) long (ventral); 210–227.5 (227.5) long (dorsal); 110–115 (110) tall) colorless. Rostrum (110–120 (120) long; 40–42.5 (40) wide). Chelicerae (275–295 (295) long) with curved fangs (55–57.5 (55) long). Subcapitular proportions: ventral length/height 2.36–2.66 (2.66); rostrum length/width 2.75–3.00 (3.00). Pedipalps with tuberculate ventral extensions on femora and genua. Palpomeres: trochanter (42.5–46.25 (45) long); femur (110–116.25 (116.25) long); genu (67.5–72.5 (72.5) long); tibia (97.5–102.5 (102.5) long; 22.5–25 (23.75) wide); tarsus (17.5–20 (17.5) long). Palpomere proportions: femur/genu 1.55–1.67 (1.60); tibia/femur 0.88–0.90 (0.88); tibia length/width 4.05–4.33 (4.32).

Venter — (640–685 (680) long; 470–570 (510) wide) colorless. Gnathosomal bay (132.5–150 (142.5) long; 70–80 (75) wide). Cxgl-4 subapical. Medial suture (105–125 (115) long). Genital plates (130–145 (145) long; 125–130 (130) wide). Additional measurements: Cx-1 (255–295 (295) long (total); 130–150 (150) long (medial)); Cx-3 (330–365 (340) wide); anterior venter (250–275 (267.5) long). Ventral proportions: gnathosomal bay length/width 1.81–2.00 (1.90); anterior venter/genital field length 1.79–1.96 (1.84); anterior venter length/genital field width 2.00–2.16 (2.06); anterior venter/medial suture 2.17–2.45 (2.33).

Immatures unknown.

Etymology

Specific epithet (folkertsae) named in honor of Debbie Folkerts, professor of biology at Auburn University, who, together with her late husband George Folkerts, were instrumental to JRF in channeling his passion for natural history and teaching into a career path.

Distribution

New Hampshire (Figure 81).

Figure 81. 

Torrenticola folkertsae sp. n. distribution.

Figure 82. 

Torrenticola folkertsae sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 83. 

Torrenticola folkertsae sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Remarks

Unfortunately, we were unable to acquire fresh material of Torrenticola folkertsae and therefore this species is not included in our phylogenetic analyses. However, we were able to examine morphology with material preserved in GAW. The overall similarity, distribution, and partial fusion of the dorso-lateral platelets to the dorsal plate, are consistent with placing this species in the Rusetria Complex and the Partial 2-Plate Identification Group.

Torrenticola glomerabilis Fisher & Dowling, sp. n.

Material examined

HOLOTYPE (♀): from USA, Tennessee, Sevier County, Great Smokey Mountains National Park, Sugarlands Nature Trail (35°40'47"N, 83°31'51"W), 10 Sep 2010, by IM Smith, IMS100125.

PARATYPES (23 ♀; 25 ♂): Georgia, USA: 2 ♀ and 2 ♂ from Floyd County, beside road from Everett Spring to Villanow, 1.4 kilometers south of The Pocket Campground, tributary of Johns Creek, 4 Jul 1990, by IM Smith, IMS900077 • Kentucky, USA: 2 ♀ and 3 ♂ from Bell County, Middlesboro, near north boundary of Cumberland Gap National Historical Park, Sugar Run, 9 Jul 1990, by IM Smith, IMS900084 • 2 ♀ and 1 ♂ from Bell County, Pineville, Pine Mountain State Resort Park, Laurel Cove, Lower shelter Picnic Area, 9 Jul 1990, by IM Smith, IMS900083 • Pennsylvania, USA: 1 ♀ from Fayette County, Dunbar Creek (39°57'50"N, 79°35'8.70"W), 10 Aug 2014, by MJ Skvarla, MS 14-0810-001 • 1 ♀ from Huntingdon County, Alan Seeger Natural Area, beside road from McAlevys Fort to Route 322, Stone Creek, 19 Jul 1990, by IM Smith, IMS900107 • Tennessee, USA: 1 ♂ (ALLOTYPE) from Sevier County, Great Smokey Mountains National Park, Sugarlands Nature Trail (35°40'47"N, 83°31'51"W), 10 Sep 2010, by IM Smith, IMS100125 • 1 ♂ from Sevier County, Great Smokey Mountains National Park, Laurel Creek (35°39'7"N, 83°42'32"W), 17 Sep 2010, by IM Smith, IMS100145 • 4 ♀ and 5 ♂ from Sevier County, Great Smokey Mountains National Park, Sugarlands Nature Trail (35°40'47"N, 83°31'51"W), 10 Sep 2010, by IM Smith, IMS100125 • Virginia, USA: 1 ♀ and 1 ♂ from Alleghany County, Clifton Forge, beside Route 606, 1.2 kilometers southeast of Forest Route 125, Smith Creek, 13 Jul 1990, by IM Smith, IMS900093 • 1 ♀ and 1 ♂ from Alleghany County, Longdale Furnace, beside Forest Route 108, 1.7 kilometers west of Route 850, Simpson Creek, 14 Jul 1990, by IM Smith, IMS900094 • 1 ♀ and 1 ♂ from Amherst County, Upper Otter Creek Overlook beside Blue Ridge, Otter Creek (37°36'57"N, 79°19'27"W), 7 Sep 2007, by IM Smith, IMS070056A • 2 ♀ and 2 ♂ from Augusta County, beside Forest Road 42, 15.9 kilometers east of Vesuvius, Coles Run, 26 Jul 1990, by IM Smith, IMS900060 • 2 ♂ from Bath County, beside Forest Route 1744 at Route 39, between Warm Springs and Mountain Grove, O’Roarke Draw, 15 Jul 1990, by IM Smith, IMS900098 • 1 ♀ and 2 ♂ from Bath County, beside Forest Route 364, off Route 39 east of Warm Springs, Panther Run, 15 Jul 1990, by IM Smith, IMS900099 • 1 ♀ and 1 ♂ from Giles County, Mechanicsburg, beside Dismal Creek Road, Standrock Brook (37°11'38"N, 80°53'26"W), 9 Sep 2005, by IM Smith, IMS050066 • 2 ♀ and 1 ♂ from Giles County, Mechanicsburg, beside Dismal Creek Road, Standrock Brook (37°11'38"N, 80°53'26"W), 11 Jul 1990, IMS900088 • 3 ♂ from Montgomery County, Blacksburg, beside Route 621 at Caldwell Fields Campground, Craig Creek (37°20'N, 80°20'W), 12 Jul 1990, by IM Smith, IMS900089A • 1 ♀ and 1 ♂ from Page County, beside Route 730, 0.2 kilometers west of Route 675, Passage Creek, 25 Jun 1990, by IM Smith, IMS900059 • 1 ♀ from Rock Bridge County, Vesuvius, beside Route 56, 2.2 kilometers west of Blue Ridge Parkway, Little Marys Creek, 26 Jun 1990, by IM Smith, IMS900062.

Type deposition

Holotype (♀), allotype (♂), and most paratypes (18 ♀; 19 ♂) deposited in the CNC; other paratypes (5 ♀; 5 ♂) deposited in the ACUA.

Diagnosis

Torrenticola glomerabilis are similar to other members of the Rusetria “4-Plate” group (T. dunni, T. glomerabilis, T. kittatinniana, T. pollani, T. rufoalba and T. shubini) and T. skvarlai in having anterio-lateral platelets free from the dorsal plate, dorsal coloration separated into anterior and posterior portions, and indistinct hind coxal margins. T. glomerabilis can be differentiated from T. dunni, T. shubini, T. kittatinniana, by having Dgl-4 closer to the dorsal edge (dorsal width/distance between Dgl-4 = 1.53–1.66 in T. glomerabilis, 1.20–1.42 in others) and stockier tibiae (length/width ♀ = 4.11–4.50 in T. glomerabilis, 3.27–3.60 in others; ♂ = 3.55–4.38 in T. glomerabilis, 2.80–3.45 in others). T. glomerabilis can be differentiated from T. pollani and T. rufoalba by having stockier anterio-medial platelets (length/width ♀ = 1.9–2.3 in T. glomerabilis, 2.5–3.0 in others; ♂ = 1.9–2.2 in T. glomerabilis, 2.3–2.9 in others) and wider dorsum (♀ = 460–490 in T. glomerabilis, 400–420 in others; ♂ = 395–430 in T. glomerabilis, 310–340 in others). T. glomerabilis can be differentiated from T. skvarlai by having a conical pedipalpal femoral tubercle, whereas T. skvarlai has a broad and flat pedipalpal femoral tubercle, and by having a longer anterior venter (♀ = 202–213 in T. glomerabilis, 140–153 in T. skvarlai; ♂ = 240–280 in T. glomerabilis, 177–205 in T. skvarlai).

Description

Female (Figure 85) (n = 5) (holotype measurements in parentheses when available) with characters of the genus with following specifications.

Dorsum — (580–615 (605) long; 460–490 (475) wide) circular with bold bluish-purple or reddish-purple coloration separated into anterior and posterior portions. Anterio-medial platelets (125–132.5 (132.5) long; 55–65 (60) wide). Anterio-lateral platelets (172.5–195 (180) long; 70–82.5 (82.5) wide) free from dorsal plate. Dgl-4 approaching midway between muscle scars and dorsum edge (distance between Dgl-4 280–310 (310)). Dorsal plate proportions: dorsum length/width 1.23–1.32 (1.27); dorsal width/distance between Dgl-4 1.53–1.66 (1.53); anterio-medial platelet length/width 1.96–2.27 (2.21); anterio-lateral platelet length/width 2.17–2.48 (2.18); anterio-lateral/anterio-medial length 1.36–1.53 (1.36).

Gnathosoma — Subcapitulum (320–330 (320) long (ventral); 223–243 (223) long (dorsal); 112.5–120 (117.5) tall) colorless. Rostrum (132.5–137.5 (132.5) long; 40–47.5 (40) wide). Chelicerae (320–330 (321) long) with curved fangs (50–55 (53) long). Subcapitular proportions: ventral length/height 2.72–2.89 (2.72); rostrum length/width 2.89–3.34 (3.31). Pedipalps with tuberculate ventral extensions on femora and genua. Palpomeres: trochanter (37.5–42.5 (40) long); femur (112.5–117.5 (116.25) long); genu (67.5–70 (67.5) long); tibia (88.75–95 (88.75) long; 20–22.5 (20) wide); tarsus (17.5–17.5 (17.5) long). Palpomere proportions: femur/genu 1.67–1.74 (1.72); tibia/femur 0.76–0.83 (0.76); tibia length/width 4.11–4.50 (4.44).

Venter — (710–730 (730) long; 512–550 (513) wide) bold bluish-purple or reddish-purple coloration. Gnathosomal bay (130–155 (155) long; 72.5–95 (72.5) wide). Cxgl-4 subapical. Medial suture (22.5–50 (27.5) long). Genital plates (167.5–177.5 (175) long; 150–157.5 (157.5) wide). Additional measurements: Cx-1 (276–305 (291) long (total); 122–160 (149) long (medial)); Cx-3 (320–370 (321) wide); anterior venter (202.5–212.5 (202.5) long). Ventral proportions: gnathosomal bay length/width 1.37–2.14 (2.14); anterior venter/genital field length 1.14–1.24 (1.16); anterior venter length/genital field width 1.29–1.40 (1.29); anterior venter/medial suture 4.15–9.00 (7.36).

Male (Figure 86) (n = 6) (allotypic measurements in parentheses when available) with characters of the genus with following specifications.

Dorsum— (495–575 (530) long; 395–430 (420) wide) circular with bold bluish-purple or reddish-purple coloration separated into anterior and posterior portions. Anterio-medial platelets (112.5–120 (120) long; 52.5–60 (55) wide). Anterio-lateral platelets (165–187.5 (167.5) long; 60–67.5 (60) wide) free from dorsal plate. Dgl-4 approaching midway between muscle scars and dorsum edge (distance between Dgl-4 235–280 (275)). Dorsal plate proportions: dorsum length/width 1.25–1.34 (1.26); dorsal width/distance between Dgl-4 1.50–1.68 (1.53); anterio-medial platelet length/width 1.96–2.18 (2.18); anterio-lateral platelet length/width 2.54–2.88 (2.79); anterio-lateral/anterio-medial length 1.40–1.67 (1.40).

Gnathosoma — Subcapitulum (260–297.5 (290) long (ventral); 188–225 (212.5) long (dorsal); 93.75–103.75 (93.75) tall) colorless. Rostrum (105–120 (120) long; 35–40 (35) wide). Chelicerae (249–298 (285) long) with curved fangs (40–50 (50) long). Subcapitular proportions: ventral length/height 2.77–3.09 (3.09); rostrum length/width 2.80–3.43 (3.43). Pedipalps with tuberculate ventral extensions on femora and genua. Palpomeres: trochanter (35–35 (35) long); femur (95–105 (102.5) long); genu (57.5–65 (60) long); tibia (80–88.75 (82.5) long; 20–25 (20) wide); tarsus (15–17.5 (15) long). Palpomere proportions: femur/genu 1.58–1.71 (1.71); tibia/femur 0.80–0.87 (0.80); tibia length/width 3.55–4.38 (4.13).

Venter — (600–690 (670) long; 443–540 (460) wide) bold bluish-purple or reddish-purple coloration. Gnathosomal bay (107.5–135 (132.5) long; 70–80 (70) wide). Cxgl-4 subapical. Medial suture (85–107.5 (92.5) long). Genital plates (135–147.5 (140) long; 110–120 (117.5) wide). Additional measurements: Cx-1 (224–280 (280) long (total); 88–160 (150) long (medial)); Cx-3 (292–342 (300) wide); anterior venter (240–280 (260) long). Ventral proportions: gnathosomal bay length/width 1.48–1.93 (1.89); anterior venter/genital field length 1.78–1.90 (1.86); anterior venter length/genital field width 2.13–2.41 (2.21); anterior venter/medial suture 2.47–2.87 (2.81).

Immatures unknown.

Etymology

Specific epithet (glomerabilis) refers to the rounded body of this species compared to all other members of the Rusetria Complex (glomerabilis, L. round).

Distribution

Appalachians (Figure 84).

Figure 84. 

Torrenticola glomerabilis sp. n. distribution.

Figure 85. 

Torrenticola glomerabilis sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 86. 

Torrenticola glomerabilis sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Remarks

Torrenticola glomerabilis groups with other members of the Rusetria Complex with high support and specimens are less than 1% different in COI sequence from each other. In all analyses, T. glomerabilis groups with two other species, T. delicatexa and T. ululata, which are 10–13% different from each other. Of these species, T. glomerabilis is the only one with the lateral platelets free from the dorsal plate. Because of this, we place T. glomerabilis within the Eastern 4-Plate Identification Group, which can be differentiated by having rounder bodies than any other species in the Rusetria Complex.

Torrenticola glomerabilis occupy an interesting position phylogenetically by being nested between the Eastern 2-Plate and Eastern 4-Plate Identification Groups (Figure 6, 12). Their unique shape and interesting phylogenetic affinity flag this species as important to future studies on the evolution of eastern members of the Rusetria Complex.

This species hypothesis is supported by biogeography, low COI divergence within the species (0–2%) and high divergence between species (3–15%), and the morphological characters outlined in the diagnosis.

Torrenticola gnoma Fisher & Dowling, sp. n.

Material examined

HOLOTYPE (♀): from USA, Mississippi, Attala County, Hurricane Creek (33°4'57"N, 89°31'29"W), 12 Sep 2008, by IM Smith, IMS080052.

PARATYPES (6 ♀; 10 ♂): Georgia, USA: 1 ♂ from Lowndes County, Withlacoochee River, beside Route 84 at Brooks County line, 13 Sep 1968, by DR Cook, DRC680075 • Illinois, USA: 1 ♀ and 2 ♂ from Clark County, Big Creek (32°25'59"N, 87°41'15"W), 30 Jul 2014, by MJ Skvarla, MS 14-0730-001 • 1 ♂ from Vermilion County, Fairmount, beside Vermilion County Road 680E, Jordan Creek (40°4'N, 87°50'W), 10 Sep 1991, by IM Smith, IMS910030 • Mississippi, USA: 1 ♂ (ALLOTYPE) from USA, Mississippi, Attala County, Hurricane Creek (33°4'57"N, 89°31'29"W), 12 Sep 2008, by IM Smith, IMS080052 • 1 ♀ and 1 ♂ from Attala County, Hurricane Creek (33°4'58"N, 89°31'31"W), 30 Sep 2010, by IM Smith, IMS100168 • 1 ♀ and 1 ♂ from Grenada County, Leflore, beside Black Creek Road, Black Creek (33°43'N, 90°3'W), 16 Sep 1991, by IM Smith, IMS910044 • 1 ♀ and 1 ♂ from Jefferson County, off Natchez Trace Parkway, Coles Creek (31°41'26"N, 91°10'52"W), 2 Oct 1994, by IM Smith, IMS940029A • 1 ♀ and 1 ♂ from Tishomingo County, Tishomingo State Park, Bear Creek, (34°36'N, 88°11'W), 18 Sep 1991, by IM Smith, IMS910047A • Oklahoma, USA: 2 ♀ and 2 ♂ from Pushmataha County, beside Route 271, South of Albion, Walnut Creek (34°39'N, 95°7'W), 1 Jul 1987, by IM Smith, IMS870063A.

Type deposition

Holotype (♀), allotype (♂), and some paratypes (4 ♀; 5 ♂) deposited in the CNC; other paratypes (2 ♀; 3 ♂) deposited in ACUA.

Diagnosis

Torrenticola gnoma are similar to other members of the Raptor Group (T. irapalpa, T. longitibia, T. mjolniri, T. elusiva, T. racupalpa, T. raptor, T. danielleae, T. daemon, and T. ivyae) in having round bodies; Dgl-4 close to muscles scars; long, thin subcapitular rostra; and long, thin pedipalp tibiae. T. gnoma can be differentiated from T. elusiva by being smaller (dorsum length = 540–595 in T. gnoma, 645 in T. elusiva); having a stockier rostrum (length/width = 2.74–3.13 in T. gnoma, 3.65 in T. elusiva); and by dorsal coloration. T. gnoma can be differentiated from T. racupalpa by having a stockier rostrum (length/width = 2.74–3.13 in T. gnoma, 3.56–3.88 in T. racupalpa) and by dorsal coloration and pattern. T. gnoma can be differentiated from T. irapalpa, T. danielleae, and T. daemon by dorsal coloration and pattern. Additionally, female T. gnoma can be differentiated from female T. irapalpa, T. danielleae, and T. daemon by having Dgl-4 closer to the muscle scars (dorsal width/distance between Dgl-4 ♀ = 2.65–3.29 in T. gnoma, 1.57–2.09 in others). T. gnoma can be differentiated from T. mjolniri, T. longitibia, T. raptor, and T. ivyae by having stockier pedipalp tibiae (length/width = 3.88–4.67 in T. gnoma, 4.75–7.54 in others) and a stockier rostrum (length/width = 2.56–3.23 in T. gnoma, 3.44–4.4 in others).

Description

Female (Figure 88) (n = 5) (holotype measurements in parentheses when available) with characters of the genus with following specifications.

Dorsum — (540–595 (550) long; 440–500 (455) wide) circular with a large spot of coloration medially extending in a thin strip anteriorly to the edge of the dorsal plate, coloration variable from navy blue to purple to pink. Anterio-medial platelets (122.5–137.5 (131.25) long; 55–62.5 (60) wide). Anterio-lateral platelets (152.5–187.5 (167.5) long; 67.5–75 (70) wide) free from dorsal plate. Dgl-4 much closer to the muscle scar than to dorsum edge (distance between Dgl-4 140–185 (155)). Dorsal plate proportions: dorsum length/width 1.17–1.30 (1.21); dorsal width/distance between Dgl-4 2.65–3.29 (2.94); anterio-medial platelet length/width 2.06–2.39 (2.19); anterio-lateral platelet length/width 2.26–2.59 (2.39); anterio-lateral/anterio-medial length 1.11–1.46 (1.28).

Gnathosoma — Subcapitulum (285–305 (290) long (ventral); 225–239 (226) long (dorsal); 115–135 (120) tall) colorless. Rostrum (122.5–130 (125) long; 40–47.5 (40) wide) elongate. Chelicerae (285–310 (286) long) with curved fangs (53–60 (54) long). Subcapitular proportions: ventral length/height 2.26–2.52 (2.42); rostrum length/width 2.74–3.13 (3.13). Pedipalps elongate with tuberculate ventral extensions on femora and genua. Palpomeres: trochanter (40–45 (40) long); femur (110–120 (112.5) long); genu (60–67.5 (62.5) long); tibia (87.5–105 (92.5) long; 20–22.5 (20) wide); tarsus (17.5–20 (17.5) long). Palpomere proportions: femur/genu 1.63–1.88 (1.80); tibia/femur 0.76–0.88 (0.82); tibia length/width 4.17–4.67 (4.63).

Venter — (660–730 (680) long; 500–575 (500) wide) colorless. Gnathosomal bay (142.5–172.5 (152.5) long; 75–92.5 (75) wide). Cxgl-4 subapical. Medial suture (15–27.5 (27.5) long). Genital plates (152.5–165 (157.5) long; 142.5–152.5 (142.5) wide). Additional measurements: Cx-1 (252–285 (257) long (total); 84–122 (108) long (medial)); Cx-3 (317–377 (318) wide); anterior venter (160–167.5 (165) long). Ventral proportions: gnathosomal bay length/width 1.57–2.16 (2.03); anterior venter/genital field length 0.97–1.10 (1.05); anterior venter length/genital field width 1.05–1.16 (1.16); anterior venter/medial suture 6.00–10.67 (6.00).

Male (Figure 89) (n = 5) (allotypic measurements in parentheses when available) with characters of the genus with following specifications.

Dorsum — (420–495 (450) long; 355–375 (375) wide) circular with a large spot of coloration medially extending in a thin strip anteriorly to the edge of the dorsal plate, coloration variable from navy blue to purple to pink. Anterio-medial platelets (1.80–2.30 (2.18) long; 50–62.5 (55) wide). Anterio-lateral platelets (135–152.5 (152.5) long; 60–65 (60) wide) free from dorsal plate. Dgl-4 much closer to the muscle scar than to dorsum edge (distance between Dgl-4 130–180 (165)). Dorsal plate proportions: dorsum length/width 1.17–1.32 (1.20); dorsal width/distance between Dgl-4 2.06–2.73 (2.27); anterio-medial platelet length/width 1.80–2.30 (2.18); anterio-lateral platelet length/width 2.16–2.54 (2.54); anterio-lateral/anterio-medial length 1.17–1.36 (1.27).

Gnathosoma— Subcapitulum (240–265 (265) long (ventral); 175–196 (196) long (dorsal); 97.5–105 (105) tall) colorless. Rostrum (98.75–107.5 (107.5) long; 35–40 (35) wide) elongate. Chelicerae (225–257 (256) long) with curved fangs (41–51 (50) long). Subcapitular proportions: ventral length/height 2.29–2.56 (2.52); rostrum length/width 2.56–3.07 (3.07). Pedipalps elongate with tuberculate ventral extensions on femora and genua. Palpomeres: trochanter (35–47.5 (47.5) long); femur (96.25–103.75 (103.75) long); genu (52.5–57.5 (57.5) long); tibia (77.5–90 (88.75) long; 20–22.5 (20) wide); tarsus (17.5–20 (18.75) long). Palpomere proportions: femur/genu 1.72–1.83 (1.80); tibia/femur 0.81–0.91 (0.86); tibia length/width 3.88–4.44 (4.44).

Venter — (560–590 (581) long; 354–440 (395) wide) colorless. Gnathosomal bay (105–130 (127.5) long; 62.5–77.5 (62.5) wide). Cxgl-4 subapical. Medial suture (67.5–80 (70) long). Genital plates (122.5–135 (127.5) long; 100–110 (105) wide). Additional measurements: Cx-1 (217–269 (255) long (total); 90–119 (115) long (medial)); Cx-3 (264–312 (295) wide); anterior venter (197.5–222.5 (207.5) long). Ventral proportions: gnathosomal bay length/width 1.40–2.04 (2.04); anterior venter/genital field length 1.49–1.65 (1.63); anterior venter length/genital field width 1.80–2.12 (1.98); anterior venter/medial suture 2.53–2.96 (2.96).

Immatures unknown.

Etymology

Specific epithet (gnoma) refers to the dorsal pattern, which, although variable, resembles the head and cap of a gnome (gnoma, L. diminutive fabled being, dwarf).

Distribution

Eastern, but apparently absent from Appalachians and Northeast (Figure 87).

Figure 87. 

Torrenticola gnoma sp. n. distribution.

Figure 88. 

Torrenticola gnoma sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 89. 

Torrenticola gnoma sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Remarks

Torrenticola gnoma groups with other members of the Raptor Complex with high support and specimens of this species were less than 1% different in COI sequence from each other. In the combined analysis, T. gnoma groups with T. irapalpa with high support, but the position of this clade was not recovered. These species are greater than 9% from each other. Based upon overall similarity, distribution, and phylogenetic position, this species is placed within the Raptor Identification Group.

This species hypothesis is supported by low COI divergence within the species (0–2%) and high divergence between species (3–15%), and by the morphological characters outlined in the diagnosis.

Torrenticola gorti Fisher & Dowling, sp. n.

Material examined

HOLOTYPE (♀): from Canada, New Brunswick, York County, Davis Brook, beside Hwy 3, 3.5 km south of Hwy 4 at Thomaston Corner, 11 Jun 2012, by IM Smith, IMS120017, DNA 2970.

PARATYPES (24 ♀; 22 ♂): Alabama, USA: 1 ♀ and 1 ♂ from Cleburne County, beside Route 431, 3.3 kilometers southeast of Calhoun, Jackson Creek (33°36'N, 85°42'W), 2 Jul 1990, by IM Smith, IMS900074 • 2 ♀ and 1 ♂ from DeKalb County, Desoto State Park, beside Trail Y, West Fork of Little River (34°29'N, 85°32'W), 26 Sep 1992, by IM Smith, IMS920053A • Georgia, USA: 2 ♀ and 2 ♂ from Chattooga County, Cloudland, beside Route 48, East Fork of Little River (34°31'25"N, 85°30'23"W), 28 Sep 1992, by IM Smith, IMS920056A • 4 ♀ and 1 ♂ from Floyd County, Johns Creek, beside road from Everett Springs to Villanow, south of The Pocket campground, 4 Jul 1990, by IM Smith, IMS900076 & IMS900077 • Kentucky, USA: 3 ♀ and 2 ♂ from McCreary County, Rock Creek, White Oak Junction, beside Forest Route 556, south of Route 1363, 8 Jul 1990, by IM Smith, IMS900082A & IMS900082B • Maine, USA: 2 ♀ and 3 ♂ from Aroostook County, Ashland, beside Route 11 at bridge, Aroostook River (46°38'N, 68°24'W), 4 Jul 1989, by IM Smith, IMS890067 • New Brunswick, Canada: 1 ♂ (ALLOTYPE) from York County, Davis Brook, beside Hwy 3, 3.5 km south of Hwy 4 at Thomaston Corner, 11 Jun 2012, by IM Smith, IMS120017, DNA 2972 • 2 ♀ and 2 ♂ from York County, Davis Brook, beside Hwy 3, 3.5 km south of Hwy 4 at Thomaston Corner, 11 Jun 2012, by IM Smith, IMS120017 • 1 ♀ and 1 ♂ from York County, Magaguadavic River, beside Highway 3 just east of Thomaston Corners, 1 Jul 1989, by IM Smith, IMS890055A • South Carolina, USA: 1 ♂ from Greenville County, Matthews Creek, 24 Apr 2014, by D Eargle, JRF 14-0424-001 • Tennessee, USA: 4 ♀ and 1 ♂ from Monroe County, Tellico River (35°19'N, 84°10'W), 5 Jun 1990, by IM Smith, IMS900079 • 2 ♂ from Monroe County, Turkey Creek, beside Forest Route 35, northeast of road from Route 165 to Miller Chapel Church, 5 Jul 1990, by IM Smith, IMS900078 • 2 ♀ and 4 ♂ from Monroe County, Tellico River (35°20'27"N, 84°11'31"W), 12 Sep 2009, by IM Smith, IMS090111 • Virginia, USA: 1 ♀ from Scott County, beside Route 58/421,) .9 kilometers east of Route 709, North Fork of Hoiston River (36°39'N, 82°28'W), 7 Jul 1990, by IM Smith, IMS900080.

Type deposition

Holotype (♀), allotype (♂), and most paratypes (19 ♀; 16 ♂) deposited in the CNC; other paratypes (5 ♀; 5 ♂) deposited in the ACUA.

Diagnosis

Torrenticola gorti specimens from Tellico River system in Monroe County (Tennessee) can be differentiated from all other Torrenticola by the distinctively dark coloration with a red spot dorsally. Other color morphs are similar to species with similar dorsal patterning, such as the Rusetria “4-Plate” group (T. dunni, T. glomerabilis, T. kittatinniana, T. pollani, T. rufoalba and T. shubini), Neoanomala Group (T. interiorensis and T. neoanomala), and T. bondi, T. elongata, T. reduncarostra, T. erectirostra, T. robisoni, T. irapalpa, T. racupalpa, T. skvarlai, and T. arktonyx. They can be differentiated from Rusetria 4-Plates and T. skvarlai by having distinct hind coxal margins. T. gorti can be differentiated from T. erectirostra, T. robisoni, and T. reduncarostra by having a straight, anteriorly-directed rostrum (upturned in others). T. gorti can be differentiated from T. arktonyx by having an unmodified dorsal plate (T. arktonyx has distinctive longitudinal dark markings on the anterior portion of the dorsal plate that fade posteriorly). T. gorti can be differentiated from T. racupalpa and T. irapalpa by being more elongate (dorsum length/width = 1.47–1.6 in T. gorti, 1.15–1.3 in others) and tibia/femur (0.65–0.73 in T. gorti, 0.77–0.91 in others). T. gorti can be differentiated from T. elongata by having a more ovoid dorsum (length/width = 1.47–1.58 in T. gorti, 1.7–2.1 in T. elongata) and larger dorsum (length ♀ = 570–600 in T. gorti, 540–565 in T. elongata; ♂ = 500–525 in T. gorti, 450–460 in T. elongata). T. gorti can be differentiated from the Neoanomala Group by having a more elongate dorsum (length/width ♀ = 1.47–1.58 in T. gorti, 1.29–1.43 in Neoanomala Group; ♂ = 1.54–1.58 in T. gorti, 1.34–1.50 in Neoanomala Group) and having a more elongate rostrum (length/width = 3.29–3.73 in T. gorti, 2.59–2.90 in Neoanomala Group). T. gorti can be differentiated from T. bondi by having a more elongate dorsum (length/width = 1.47–1.58 in T. gorti, 1.32–1.45 in T. bondi) and having a more elongate rostrum (length/width = 3.32–3.73 in T. gorti, 2.76–3.13 in T. bondi).

Description

Female (Figure 91) (n = 5) (holotype measurements in parentheses when available) with characters of the genus with following specifications.

Dorsum — (570–600 (600) long; 380–390 (380) wide) ovoid and elongate with three distinct color morphs: 1) navy blue coloration separated into anterior and posterior portions that meet or nearly meet laterally, and with bold orange coloration in between, from Tellico River, Monroe County, Tennessee; 2) purple coloration separated into anterior and posterior portions; 3) purple coloration separated into anterior and posterior portions with a strip of bold orange medially. Anterio-medial platelets (117.5–137.5 (130) long; 52.5–57.5 (57.5) wide). Anterio-lateral platelets (162.5–172.5 (172.5) long; 55–60 (60) wide) free from dorsal plate. Dgl-4 approaching midway between muscle scars and dorsum edge (distance between Dgl-4 220–260 (260)). Dorsal plate proportions: dorsum length/width 1.47–1.58 (1.58); dorsal width/distance between Dgl-4 1.46–1.73 (1.46); anterio-medial platelet length/width 2.24–2.45 (2.26); anterio-lateral platelet length/width 2.75–3.09 (2.88); anterio-lateral/anterio-medial length 1.20–1.45 (1.33).

Gnathosoma — Subcapitulum (327.5–342.5 (337.5) long (ventral); 247.75–265 (252.5) long (dorsal); 122.5–127.5 (122.5) tall) colorless. Rostrum (137.5–142.5 (142.5) long; 37.5–42.5 (40) wide) elongate. Chelicerae 330–345 (340) long) with curved fangs (56–60 (60) long). Subcapitular proportions: ventral length/height 2.59–2.76 (2.76); rostrum length/width 3.32–3.73 (3.56). Pedipalps with tuberculate ventral extensions on femora and genua. Palpomeres: trochanter (37.5–45 (45) long); femur (117.5–123.75 (120) long); genu (62.5–67.5 (67.5) long); tibia (77.5–82.5 (80) long; 20–25 (25) wide); tarsus (17.5–20 (20) long). Palpomere proportions: femur/genu 1.78–1.90 (1.78); tibia/femur 0.65–0.68 (0.67); tibia length/width 3.20–4.13 (3.20).

Venter (Figure 57) — (695–750 (750) long; 410–494 (430) wide) with three distinct color morphs: 1) navy-blue coloration; 2) colorless; 3) purple coloration. Gnathosomal bay (142.5–172.5 (172.5) long; 65–77.5 (70) wide). Cxgl-4 subapical. Medial suture (25–37.5 (27.5) long). Genital plates (160–167.5 (167.5) long; 135–140 (137.5) wide). Additional measurements: Cx-1 (285–300 (300) long (total); 107–140 (130) long (medial)); Cx-3 (290–353 (300) wide); anterior venter (187.5–195 (187.5) long). Ventral proportions: gnathosomal bay length/width 1.90–2.46 (2.46); anterior venter/genital field length 1.12–1.22 (1.12); anterior venter length/genital field width 1.36–1.44 (1.36); anterior venter/medial suture 5.20–7.80 (6.82).

Male (Figure 92) (n = 6) (allotypic measurements in parentheses when available) with characters of the genus with following specifications.

Dorsum — (500–525 (520) long; 320–340 (330) wide) ovoid and elongate with three distinct color morphs: 1) navy blue coloration separated into anterior and posterior portions that meet or nearly meet laterally, and with bold orange coloration in between; 2) purple coloration separated into anterior and posterior portions; 3) purple coloration separated into anterior and posterior portions with a strip of bold orange medially. Anterio-medial platelets (108.75–117.5 (115) long; 47.5–57.5 (50) wide). Anterio-lateral platelets (156.25–165 (157.5) long; 52.5–57.5 (53.75) wide) free from dorsal plate. Dgl-4 approaching midway between muscle scars and dorsum edge (distance between Dgl-4 210–230 (230)). Dorsal plate proportions: dorsum length/width 1.54–1.58 (1.58); dorsal width/distance between Dgl-4 1.43–1.58 (1.43); anterio-medial platelet length/width 2.04–2.35 (2.30); anterio-lateral platelet length/width 2.78–2.98 (2.93); anterio-lateral/anterio-medial length 1.36–1.44 (1.37).

Gnathosoma — Subcapitulum (280–292.5 (282.5) long (ventral); 211–231 (215) long (dorsal); 92.5–1.5 (95) tall) colorless. Rostrum (115–127.5 (122.5) long; 33.75–37.5 (35) wide) elongate. Chelicerae (275–302 (280) long) with curved fangs (36–50 (50) long). Subcapitular proportions: ventral length/height 2.76–3.16 (2.97); rostrum length/width 3.29–3.50 (3.50). Pedipalps with tuberculate ventral extensions on femora and genua. Palpomeres: trochanter (35–37.5 (35) long); femur (87.5–105 (95) long); genu (52.5–58.75 (56.25) long); tibia (62.5–72.5 (67.5) long; 18.75–22.5 (18.75) wide); tarsus (15–17.5 (15) long). Palpomere proportions: femur/genu 1.67–1.79 (1.69); tibia/femur 0.67–0.73 (0.71); tibia length/width 3.13–3.63 (3.60).

Venter — (630–680 (640) long; 380–438 (380) wide) with three distinct color morphs: 1) navy-blue coloration; 2) colorless; 3) purple coloration. Gnathosomal bay (122.5–142.5 (142.5) long; 60–67.5 (60) wide). Cxgl-4 subapical. Medial suture (62.5–87.5 (65) long). Genital plates (125–135 (135) long; 102.5–111.25 (102.5) wide). Additional measurements: Cx-1 (255–280 (280) long (total); 100–140 (140) long (medial)); Cx-3 (285–304 (290) wide); anterior venter (215–250 (215) long). Ventral proportions: gnathosomal bay length/width 1.89–2.38 (2.38); anterior venter/genital field length 1.59–1.89 (1.59); anterior venter length/genital field width 1.96–2.27 (2.10); anterior venter/medial suture 2.86–3.48 (3.31).

Immatures unknown.

Etymology

Specific epithet (gorti) refers to Gort, the fictional giant robot of The Day the Earth Stood Still. In the 2008 film, Gort was depicted with a dark body and a single red eye that shot a destructive beam. This species is named for the resemblance that the distinctive specimens from Tennessee have to Gort’s red eye.

Distribution

Appalachians (Figure 90).

Figure 90. 

Torrenticola gorti sp. n. distribution.

Figure 91. 

Torrenticola gorti sp. n. female, Tennessee specimen depicted except for A (top): A dorsal plates, note variation in color between Tennessee specimens (bottom) and elsewhere (top) B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 92. 

Torrenticola gorti sp. n. male, Tennessee specimen depicted: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Remarks

Torrenticola gorti groups with other members of the Raptor Complex with high support and specimens of this species were less than 1% different in COI sequence from each other. In all analyses, T. elongata groups with two other species (T. elongata and T. bondi) which are 4% different from each other and have non-overlapping ranges.

T. elongata to form the Elongata Group in all analyses with high support.

Based upon overall similarity, an elongate body, and distribution, we place this species in the Elongata Identification Group.

This species hypothesis is supported by low COI divergence within the species (0–2%) and high divergence between species (3–15%), and by the morphological characters outlined in the diagnosis.

Members of this species can be highly variable in color. Some members have reddish-purple or purple dorsal coloration that is separated into anterior and posterior portions. Ventral coloration can be bold, faint, or absent. Members from Tellico River, Monroe County (Tennessee), can be readily differentiated from all other Torrenticola by being dark navy blue with a red dorsal oval.

Torrenticola hoosieri Fisher & Dowling, sp. n.

Material examined

HOLOTYPE (♀): from USA, Indiana, Wayne County, south of I-70 (39°51'13"N, 85°8'4"W), 31 Jul 2014, by MJ Skvarla, MS 14-0731-001.

PARATYPES (1 ♀; 5 ♂): Indiana: 1 ♂ (ALLOTYPE) from Wayne County, south of I-70 (39°51'13"N, 85°8'4"W), 31 Jul 2014, by MJ Skvarla, MS 14-0731-001 • 1 ♀ and 4 ♂ from Wayne County, south of I-70 (39°51'13"N, 85°8'4"W), 31 Jul 2014, by MJ Skvarla, MS 14-0731-001.

Type deposition

Holotype (♀), allotype (♂), and some paratypes (3 ♂) deposited in the CNC; other paratypes (1 ♀; 2 ♂) deposited in the ACUA.

Diagnosis

Torrenticola hoosieri are similar to other members of the Tricolor Complex (T. bittikoferae, T. larvata, T. pearsoni, T. olliei, T. sierrensis, T. tricolor, T. trimaculata, and T. unimaculata, T. cardia, T. kringi, T. dimorpha, and T. mohawk) in having a short, conical rostrum. T. hoosieri can be differentiated from all other Tricolor Complex, and nearly all other Torrenticola, by lacking pedipalp ventral extensions on femora and genua. Additionally, T. hoosieri can be differentiated from all other Tricolor Complex by having more elongate pedipalp tibiae (3.67–4.33 in T. hoosieri, 2.65–3.55 in others) and by being colorless (rarely with diffuse pink dorsal coloration), except T. bittikoferae, T. pearsoni, and T. dimorpha.

Description

Female (Figure 94) (n = 2) (holotype measurements in parentheses when available) with characters of the genus with following specifications.

Dorsum — (650–700 (650) long; 450–470 (450) wide) ellipsoid and colorless, occasionally with pink coloration without a distinct pattern. Anterio-medial platelets (115–125 (115) long; 55–60 (55) wide). Anterio-lateral platelets (177.5–180 (177.5) long; 67.5–67.5 (67.5) wide) free from dorsal plate. Dgl-4 much closer to the edge of the dorsum than to the muscle scars (distance between Dgl-4 345–375 (345)). Dorsal plate proportions: dorsum length/width 1.44–1.49 (1.44); dorsal width/distance between Dgl-4 1.25–1.30 (1.30); anterio-medial platelet length/width 2.08–2.09 (2.09); anterio-lateral platelet length/width 2.63–2.67 (2.63); anterio-lateral/anterio-medial length 1.44–1.54 (1.54).

Gnathosoma — Subcapitulum (277.5–300 (277.5) long (ventral); 202.5–221.23 (202.5) long (dorsal); 127.5–138.75 (127.5) tall) colorless. Rostrum (120–132.5 (120) long; 52.5–55 (52.5) wide) short and conical. Chelicerae (260–291 (260) long) with curved fangs (75–77 (75) long). Subcapitular proportions: ventral length/height 2.16–2.18 (2.18); rostrum length/width 2.29–2.41 (2.29). Pedipalps without extensions on femora and genua. Palpomeres: trochanter (50–55 (50) long); femur (130–137.5 (130) long); genu (72.5–80 (72.5) long); tibia (102.5–110 (102.5) long; 27.5–30 (27.5) wide); tarsus (25–27.5 (27.5) long). Palpomere proportions: femur/genu 1.72–1.79 (1.79); tibia/femur 0.79–0.80 (0.79); tibia length/width 3.67–3.73 (3.73).

Venter — (790–800 (800) long; 480–551 (480) wide) colorless. Gnathosomal bay (122.5–130 (130) long; 85–105 (85) wide). Cxgl-4 subapical. Medial suture (30–30 (30) long). Genital plates (182.5–188.75 (188.75) long; 150–152.5 (150) wide). Additional measurements: Cx-1 (257–260 (260) long (total); 120–129 (120) long (medial)); Cx-3 (330–390 (330) wide); anterior venter (162.5–167.5 (162.5) long). Ventral proportions: gnathosomal bay length/width 1.17–1.53 (1.53); anterior venter/genital field length 0.86–0.92 (0.86); anterior venter length/genital field width 1.08–1.10 (1.08); anterior venter/medial suture 5.42–5.58 (5.42).

Male (Figure 95) (n = 5) (allotypic measurements in parentheses when available) with characters of the genus with following specifications.

Dorsum — (580–640 (640) long; 390–410 (400) wide) ellipsoid and colorless. Anterio-medial platelets (110–117.5 (115) long; 55–60 (60) wide). Anterio-lateral platelets (167.5–175 (175) long; 65–72.5 (70) wide) free from dorsal plate. Dgl-4 much closer to the edge of the dorsum than to the muscle scars (distance between Dgl-4 320–360 (325)). Dorsal plate proportions: dorsum length/width 1.43–1.60 (1.60); dorsal width/distance between Dgl-4 1.14–1.23 (1.23); anterio-medial platelet length/width 1.91–2.05 (1.92); anterio-lateral platelet length/width 2.31–2.58 (2.50); anterio-lateral/anterio-medial length 1.43–1.59 (1.52).

Gnathosoma — Subcapitulum (245–280 (280) long (ventral); 185–202.5 (200) long (dorsal); 107.5–120 (120) tall) colorless. Rostrum (107.5–110 (110) long; 42.5–45 (45) wide). Chelicerae (245–260 (260) long) with curved fangs (65–70 (67.5) long). Subcapitular proportions: ventral length/height 2.23–2.36 (2.33); rostrum length/width 2.44–2.59 (2.44). Pedipalps without extensions on femora and genua. Palpomeres: trochanter (37.5–47.5 (45) long); femur (117.5–125 (125) long); genu (70–75 (75) long); tibia (97.5–100 (100) long; 22.5–25 (25) wide); tarsus (22.5–27.5 (27.5) long). Palpomere proportions: femur/genu 1.67–1.73 (1.67); tibia/femur 0.79–0.83 (0.80); tibia length/width 3.90–4.33 (4.00).

Venter — (670–740 (740) long; 450–495 (460) wide) colorless. Gnathosomal bay (120–130 (125) long; 80–85 (85) wide). Cxgl-4 subapical. Medial suture (102.5–122.5 (117.5) long). Genital plates (140–150 (147.5) long; 102.5–110 (105) wide). Additional measurements: Cx-1 (240–260 (260) long (total); 120–150 (130) long (medial)); Cx-3 (330–350 (345) wide); anterior venter (237.5–270 (270) long). Ventral proportions: gnathosomal bay length/width 1.47–1.59 (1.47); anterior venter/genital field length 1.70–1.84 (1.83); anterior venter length/genital field width 2.21–2.57 (2.57); anterior venter/medial suture 2.14–2.51 (2.30).

Immatures unknown.

Etymology

Specific epithet (hoosieri) refers to Hoosier, the English demonym for a person from Indiana, the type locality.

Distribution

Known only from Wayne County, Indiana (Figure 93).

Figure 93. 

Torrenticola hoosieri sp. n. distribution.

Figure 94. 

Torrenticola hoosieri sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum, note damaged rostrum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 95. 

Torrenticola hoosieri sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Remarks

Torrenticola hoosieri group with other members of the Tricolor Complex with high support and all specimens are less than 1% different in COI sequence from each other. In the combined analysis, T. hoosieri groups with other members of the Tricolor Complex from eastern North America that are not colorful, T. projector and T. pearsoni, and are greater than 4% different in COI from these species. This species is placed in the Tricolor Identification Group.

This species hypothesis is supported by low COI divergence within the species (0–2%), high divergence between species (3–15%), and the morphological characters outlined in the diagnosis.

Torrenticola indistincta (Marshall, 1929)

Atractides indistinctus Marshall, 1929: 317.

T. indistincta: Mitchell 1954: 40 • Viets 1956: 253 • Crowell 1960: 35, 37 • Crowell 1961: 330 • Habeeb 1967: 3 • Weaver 1967: 223 • Conroy 1968: 28 • Young 1969: 373-386 • Modlin and Gannon 1973: 219, 221 • Conroy and Scudder 1975: 307.

Material examined

LECTOTYPE (1 ♀): from USA, Wisconsin, Green Lake County, Green Lake, Aug 1921, by C Juday, RM210013.

PARALECTOTYPE (1 ♂):from USA, Wisconsin, Green Lake County, Green Lake, Aug 1921, by C Juday, RM210013.

OTHER MATERIAL (5 ♀; 4 ♂): Manitoba, Canada: 1 ♂ from Fidler Lake; Station 7., (57°11'11"N, 96°56'56"W), 20 June 1977, by Freshwater Institute, IMS770231 • 1 ♀ from Northern Indian Lake; Station 9., (56°47'47"N, 98°56'56"W), 20 June 1977, by Freshwater Institute, IMS770224 • 1 ♂ from Northern Indian Lake; Station 1., (56°47'47"N, 98°56'56"W), 20 June 1977, by Freshwater Institute, IMS770232 • 1 ♀ from North Pine River near Pine River, 29 May 1981, by P Schefter, E Fuller, ROM810578 • 1 ♀ and 1 ♂ from Southern Indian Lake, (57°10'10"N, 98°29'29"W), 1 July 1977, by Freshwater Institute, IMS770234 • 1 ♀ and 1 ♂ from Southern Indian Lake, (56°47'47"N, 98°56'56"W), 27 July 1977, by Freshwater Institute., IMS770215 • 1 ♀ from Southern Indian Lake, (57°10'10"N, 98°29'29"W), 5 September 1978, by Freshwater Institute., IMS780049

Type deposition

Types (1 ♀, 1 ♂); and most other material (3 ♀; 2 ♂) deposited in the CNC; other paratypes (2 ♀; 2 ♂) deposited in the ACUA.

Diagnosis

Torrenticola indistincta are similar to other members of the Rusetria “Eastern 2-Plates” group (T. biscutella, T. caerulea, T. delicatexa, T. malarkeyorum, T. pendula, T. sellersorum, T. tysoni, T. ululata, T. whitneyae, T. microbiscutella, and T. feminellai) in having anterio-lateral platelets fused to the dorsal plate, and being distributed in the east. T. indistincta can be differentiated from other Eastern 2-Plates by having faint coloration separated into anterior and posterior portions connected medially. Female T. indistincta can be differentiated from female T. caerulea by having a larger genital field (length = 185–225 in T. indistincta, 155–165 in T. caerulea; width = 185–205 in T. indistincta, 120–145 in T. caerulea). Male T. indistincta can be differentiated from male T. caerulea by having a larger dorsum (length = 480–645 in T. indistincta, 405–460 in T. caerulea; width = 315–470 in T. indistincta, 260–305 in T. caerulea). T. indistincta can be differentiated from T. microbiscutella by having a less elongate dorsum (length/width = 1.21–1.52 in T. indistincta; 1.63–1.75 in T. microbiscutella).

Re-description

Female (Figure 97) (n = 5) (lectotypic measurements in parentheses when available) with characters of the genus with following specifications.

Dorsum — (590–880 (640) long; 460–720 (485) wide) ovoid with faint coloration separated into anterior and posterior portions and connected medially. Anterio-medial platelets (125–195 (125) long; 45–77.5 (47.5) wide). Anterio-lateral platelets (172.5–235 (172.5) long; 75–115 (80) wide) fused to dorsal plate. Dgl-4 much closer to the edge of the dorsum than to the muscle scars (distance between Dgl-4 340–485 (340)). Dorsal plate proportions: dorsum length/width 1.21–1.32 (1.32); dorsal width/distance between Dgl-4 1.32–1.48 (1.43); anterio-medial platelet length/width 2.35–2.94 (2.63); anterio-lateral platelet length/width 2.02–2.33 (2.16); anterio-lateral/anterio-medial length 1.19–1.38 (1.38).

Gnathosoma — Subcapitulum (300–395 (322.5) long (ventral); 225–295 (237.5) long (dorsal); 145–195 (150) tall) tall and colorless. Rostrum (120–160 (125) long; 45–62.5 (47.5) wide). Chelicerae (300–405 (335) long) with curved fangs (60–80 (62.5) long). Subcapitular proportions: ventral length/height 1.97–2.15 (2.15); rostrum length/width 2.50–2.78 (2.63). Pedipalps with tuberculate ventral extensions on femora and genua. Palpomeres: trochanter (45–65 (47.5) long); femur (112.5–160 (117.5) long); genu (62.5–90 (62.5) long); tibia (87.5–122.5 (87.5) long; 25–31.25 (25) wide); tarsus (17.5–30 (17.5) long). Palpomere proportions: femur/genu 1.67–1.88 (1.88); tibia/femur 0.74–0.80 (0.74); tibia length/width 3.50–3.92 (3.50).

Venter — (650–995 (800) long; 535–880 (565) wide) colorless. Gnathosomal bay (175–237.5 (175) long; 90–130 (97.5) wide). Cxgl-4 subapical. Medial suture absent. Genital plates (185–225 (205) long; 185–205 (190) wide). Additional measurements: Cx-1 (295–385 (305) long (total); 120–155 (135) long (medial)); Cx-3 (345–505 (345) wide); anterior venter (125–167.5 (135) long). Ventral proportions: gnathosomal bay length/width 1.79–2.02 (1.79); anterior venter/genital field length 0.66–0.83 (0.66); anterior venter length/genital field width 0.66–0.83 (0.71).

Male (Figure 98) (n = 5) (lectotypic measurements in parentheses when available) with characters of the genus with following specifications.

Dorsum — (480–645 (480) long; 315–470 (315) wide) ovoid with faint coloration separated into anterior and posterior portions and connected medially. Anterio-medial platelets (102.5–147.5 (102.5) long; 35–55 (35) wide). Anterio-lateral platelets (137.5–180 (150) long; 55–82.5 (55) wide) fused to dorsal plate. Dgl-4 much closer to the edge of the dorsum than to the muscle scars (distance between Dgl-4 235–350 (235)). Dorsal plate proportions: dorsum length/width 1.37–1.52 (1.52); dorsal width/distance between Dgl-4 1.31–1.34 (1.34); anterio-medial platelet length/width 2.33–3.05 (2.93); anterio-lateral platelet length/width 2.12–2.73 (2.73); anterio-lateral/anterio-medial length 1.19–1.46 (1.46).

Gnathosoma — Subcapitulum (238.75–317.5 (238.75) long (ventral); 167.5–232.5 (167.5) long (dorsal); 102.5–137.5 (102.5) tall) tall and colorless. Rostrum (85–125 (85) long; 32.5–50 (32.5) wide). Chelicerae (232.5–320 (232.5) long) with curved fangs (47.5–65 (47.5) long). Subcapitular proportions: ventral length/height 2.22–2.33 (2.33); rostrum length/width 2.50–2.86 (2.62). Pedipalps with tuberculate ventral extensions on femora and genua. Palpomeres: trochanter (37.5–52.5 (37.5) long); femur (86.25–117.5 (86.25) long); genu (50–77.5 (50) long); tibia (70–100 (70) long; 22.5–28.75 (22.5) wide); tarsus (17.5–22.5 (17.5) long). Palpomere proportions: femur/genu 1.52–1.73 (1.73); tibia/femur 0.80–0.85 (0.81); tibia length/width 3.11–3.48 (3.11).

Venter — (570–780 (570) long; 370–510 (370) wide) colorless. Gnathosomal bay (127.5–180 (127.5) long; 65–100 (65) wide). Cxgl-4 subapical. Medial suture (65–75 (75) long). Genital plates (117.5–165 (117.5) long; 112.5–150 (112.5) wide). Additional measurements: Cx-1 (235–315 (235) long (total); 107.5–150 (107.5) long (medial)); Cx-3 (250–395 (277.5) wide); anterior venter (190–235 (190) long). Ventral proportions: gnathosomal bay length/width 1.80–1.96 (1.96); anterior venter/genital field length 1.38–1.68 (1.62); anterior venter length/genital field width 1.53–1.69 (1.69); anterior venter/medial suture 2.53–3.54 (2.53).

Immatures unknown.

Etymology

Marshall (1929) named the specific epithet (indistincta) in reference to the Rusetria Complex character of fused (“indistinct”) lateral platelets.

Distribution

Midwest and into Manitoba (Figure 96). Young (1969) reported this species from Colorado, but this likely represents T. mulleni rather than T. indistincta.

Figure 96. 

Torrenticola indistincta distribution. Red dots indicate material examined. Blue diamonds indicate previously published reports. Green “X” indicates previously published report (Young 1969) that requires verification of identification.

Figure 97. 

Torrenticola indistincta female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 98. 

Torrenticola indistincta male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Remarks

Unfortunately, we were unable to acquire fresh material of Torrenticola indistincta and therefore this species was not included in our phylogenetic analyses. However, we were able to examine type material and additional material preserved in GAW. The fusion of the lateral platelets to the dorsal plate clearly places this species among the Rusetria Complex and its distribution is consistent with placement within the Eastern 2-Plate Identification Group.

Although Young (1969) reported this species in Colorado, we suspect that record represents the superficially similar T. mulleni based upon distribution.

Torrenticola interiorensis Fisher & Dowling, sp. n.

Material examined

HOLOTYPE (♂): from USA, Arkansas, Montgomery County, Caddo Gap, access track off Manfred Road, 0.3 km west of Route 8, 29 Jul 2011, by IM Smith, IMS110037.

PARATYPES (7 ♀; 11 ♂): Arkansas, USA: 1 ♀ (ALLOTYPE) from Montgomery County, Caddo Gap, access track off Manfred Road, 0.3 km west of Route 8, 29 Jul 2011, by IM Smith, IMS110037 • 2 ♂ from Montgomery County, Caddo Gap, access track off Manfred Road, 0.3 km west of Route 8, 29 Jul 2011, by IM Smith, IMS110037 • 1 ♀ and 1 ♂ from Newton County, Ozark-St. Francis National Forest, Little Buffalo River, 11 Jul 2012, by TD Edwards, TDE 12-0711-004 • Missouri, USA: 3 ♀ and 5 ♂ from Crawford County, Huzzah Creek, Red Bluff campground, off Road V, east of Davisville, 23 Jul 2011, by IM Smith, IMS110029 • Oklahoma, USA: 2 ♀ and 3 ♂ from Pushmataha County, Walnut Creek, beside Route 271, south of Albion, 1 Jul 1987, by IM Smith, IMS870063A.

Type deposition

Holotype (♂), allotype (♀), and some paratypes (5 ♀; 5 ♂) deposited in the CNC; other paratypes (2 ♀; 5 ♂) deposited in the ACUA.

Diagnosis

Torrenticola interiorensis are similar to species with similar dorsal patterning, such as the Rusetria 4-Plate Group (T. dunni, T. glomerabilis, T. kittatinniana, T. pollani, T. rufoalba and T. shubini), Elongata Group (T. gorti, T. elongata, and T. reduncarostra), and T. bondi, T. erectirostra, T. robisoni, T. irapalpa, T. neoanomala, T. racupalpa, T. skvarlai, and T. arktonyx. They can be differentiated from Rusetria 4-Plates and T. skvarlai by having distinct hind coxal margins. T. interiorensis can be differentiated from T. erectirostra, T. robisoni and T. reduncarostra by having a straight, anteriorly-directed rostrum (upturned in T. erectirostra and T. reduncarostra). T. interiorensis can be differentiated from T. arktonyx by having an unmodified dorsal plate (T. arktonyx has distinctive longitudinal dark markings on the anterior portion of the dorsal plate). T. interiorensis can be differentiated from T. racupalpa by having less elongate pedipalpal tibiae (length/width = 3.76–4.22 in T. interiorensis; 4.44–5.50 in T. racupalpa) and less elongate rostrum (length/width = 2.63–2.88 in T. interiorensis; 3.56–3.88 in T. racupalpa). T. interiorensis can be differentiated from T. irapalpa by having Dgl-4 closer to the dorsal edge (dorsal width/distance between Dgl-4 ♀ = 1.48–1.61 in T. interiorensis, 1.81–2.09 in T. irapalpa; ♂ = 1.42–1.45 in T. interiorensis, 1.58–1.86 in T. irapalpa) and more ovoid dorsum (length/width ♀ = 1.29–1.38 in T. interiorensis, 1.20–1.28 in T. irapalpa; ♂ = 1.34–1.47 in T. interiorensis, 1.26–1.30 in T. irapalpa). T. interiorensis can be differentiated from Elongata Group by being slightly more ovoid (dorsum length/width ♀ = 1.29–1.38 in T. interiorensis, 1.45–2.08 in Elongata Group; ♂ = 1.34–1.47 in T. interiorensis, 1.51–1.7 in Elongata Group) and having a stockier rostrum (length/width = 2.63–2.88 in T. interiorensis, 3.24–4.00 in Elongata Group). T. interiorensis can be differentiated from T. bondi by having a longer medial suture (♀ = 25–30 in T. interiorensis, 10–15 in T. bondi; ♂ = 75–83 in T. interiorensis, 55–70 in T. bondi), anterior venter/genital field width (♀ = 1.31–1.38 in T. interiorensis, 1.15–1.25 in T. bondi; ♂ = 2.09–2.27 in T. interiorensis, 1.95–2.05 in T. bondi), and Dgl-4 closer to edge of dorsum (dorsum width/distance between Dgl-4 ♀ = 1.48–1.61 in T. interiorensis, 1.63–1.72 in T. bondi; ♂ = 1.42–1.45 in T. interiorensis, 1.48–1.62 in T. bondi). Female T. interiorensis can be differentiated from female T. neoanomala by having stockier anterio-lateral platelets (length/width = 2.62–2.67 in T. interiorensis, 2.86–3.09 in T. neoanomala). Male T. interiorensis can be differentiated from male T. neoanomala by having a shorter anterior venter (220–240 in T. interiorensis, 267.5–290 in T. neoanomala) and a shorter genital field (132–138 in T. interiorensis, 145–160 in T. neoanomala).

Description

Male (Figure 100) (n = 5) (holotype measurements in parentheses when available) with characters of the genus with following specifications.

Dorsum — (510–545 (510) long; 350–405 (360) wide) ovoid with purple coloration separated into anterior and posterior portions and orange medially. Anterio-medial platelets (115–122.5 (115) long; 45–50 (45) wide). Anterio-lateral platelets (150–167.5 (150) long; 50–52.5 (50) wide) free from dorsal plate. Dgl-4 closer to the edge of the dorsum than to the muscle scars (distance between Dgl-4 245–285 (250)). Dorsal plate proportions: dorsum length/width 1.34–1.47 (1.42); dorsal width/distance between Dgl-4 1.42–1.45 (1.44); anterio-medial platelet length/width 2.45–2.72 (2.56); anterio-lateral platelet length/width 3.00–3.19 (3.00); anterio-lateral/anterio-medial length 1.30–1.37 (1.30).

Gnathosoma — Subcapitulum (265–282.5 (265) long (ventral); 200–212 (200) long (dorsal); 105–112.5 (105) tall) colorless. Rostrum (112.5–115 (112.5) long; 40–40 (40) wide). Chelicerae (260–282 (260) long) with curved fangs (45–58 (45) long). Subcapitular proportions: ventral length/height 2.51–2.60 (2.52); rostrum length/width 2.81–2.88 (2.81). Pedipalps with tuberculate ventral extensions on femora and genua. Palpomeres: trochanter (32.5–37.5 (32.5) long); femur (97.5–105 (97.5) long); genu (55–62.5 (55) long); tibia (77.5–81.25 (77.5) long; 20–21.25 (20) wide); tarsus (17.5–18.75 (17.5) long). Palpomere proportions: femur/genu 1.68–1.78 (1.77); tibia/femur 0.77–0.79 (0.79); tibia length/width 3.76–3.88 (3.88).

Venter — (608–660 (640) long; 424–480 (480) wide) mostly colorless with faint purple in areas surrounding coxae. Gnathosomal bay (102.5–137.5 (110) long; 65–85 (75) wide). Cxgl-4 subapical. Medial suture (75–82.5 (75) long). Genital plates (132.5–137.5 (132.5) long; 100–107.5 (100) wide). Additional measurements: Cx-1 (231–270 (260) long (total); 123.25–134 (130) long (medial)); Cx-3 (300–338 (315) wide); anterior venter (220–240 (220) long). Ventral proportions: gnathosomal bay length/width 1.37–2.12 (1.47); anterior venter/genital field length 1.60–1.80 (1.66); anterior venter length/genital field width 2.09–2.27 (2.20); anterior venter/medial suture 2.73–2.98 (2.93).

Female (Figure 101) (n =5) (allotypic measurements in parentheses when available) with characters of the genus with following specifications.

Dorsum — (550–620 (550) long; 410–480 (415) wide) ovoid with purple coloration separated into anterior and posterior portions and orange coloration. Anterio-medial platelets (123.75–142.5 (123.75) long; 50–60 (50) wide). Anterio-lateral platelets (160–187.5 (160) long; 60–70 (60) wide) free from dorsal plate. Dgl-4 closer to the edge of the dorsum than to the muscle scars (distance between Dgl-4 255–305 (280)). Dorsal plate proportions: dorsum length/width 1.29–1.38 (1.33); dorsal width/distance between Dgl-4 1.48–1.61 (1.48); anterio-medial platelet length/width 2.36–2.52 (2.48); anterio-lateral platelet length/width 2.62–2.68 (2.67); anterio-lateral/anterio-medial length 1.21–1.33 (1.29).

Gnathosoma — Subcapitulum (307.5–325 (310) long (ventral); 220–240 (232.5) long (dorsal); 120–130 (120) tall) colorless. Rostrum (122.5–140 (125) long; 45–50 (45) wide). Chelicerae (300–340 (310) long) with curved fangs (50–65 (55) long). Subcapitular proportions: ventral length/height 2.44–2.58 (2.58); rostrum length/width 2.63–2.83 (2.78). Pedipalps with tuberculate ventral extensions on femora and genua. Palpomeres: trochanter (40–45 (40) long); femur (112.5–127.5 (115) long); genu (65–70 (65) long); tibia (85–95 (85) long; 21.25–22.5 (21.25) wide); tarsus (17.5–22.5 (20) long). Palpomere proportions: femur/genu 1.73–1.82 (1.77); tibia/femur 0.74–0.76 (0.74); tibia length/width 3.78–4.22 (4.00).

Venter — (680–780 (680) long; 470–580 (510) wide) mostly colorless with purple in areas surrounding coxae. Gnathosomal bay (112.5–152.5 (145) long; 70–100 (90) wide). Cxgl-4 subapical. Medial suture (25–30 (30) long). Genital plates (155–175 (155) long; 137.5–152.5 (145) wide). Additional measurements: Cx-1 (242–310 (280) long (total); 125–160 (140) long (medial)); Cx-3 (320–390 (350) wide); anterior venter (190–205 (190) long). Ventral proportions: gnathosomal bay length/width 1.22–2.07 (1.61); anterior venter/genital field length 1.17–1.23 (1.23); anterior venter length/genital field width 1.31–1.38 (1.31); anterior venter/medial suture 6.33–8.20 (6.33).

Immatures unknown.

Etymology

Specific epithet (interiorensis) refers to the Interior Highlands, where this species was found within both major regions (Ozarks and Ouachitas), but not found outside these regions, which suggests it is endemic to the region.

Distribution

Interior Highlands (Ozarks and Ouachitas), likely endemic (Figure 99).

Figure 99. 

Torrenticola interiorensis sp. n. distribution.

Figure 100. 

Torrenticola interiorensis sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 101. 

Torrenticola interiorensis sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Remarks

Torrenticola interiorensis groups with other members of the Raptor Complex with high support and specimens are less than 2% different in COI sequence from each other. In the combined analysis, T. interiorensis groups with the superficially similar T. neoanomala, and specimens from these species are greater than 9% different in COI sequence from each other. Based upon this relationship and their similarity, we place these species in the Neoanomala Identification Group. The Neoanomala Group shares a phylogenetic affinity for members of the similar-looking Erectirostra Group.

This species hypothesis is supported by low COI divergence within the species (0–2%) and high divergence between species (3–15%), and by the morphological characters outlined in the diagnosis.

Torrenticola irapalpa Fisher & Dowling, sp. n.

Material examined

HOLOTYPE (♀): from New Brunswick, Canada, York County, Stanley, Nashwaak River, Stanley Municipal Park, 19 Jun 2012, by IM Smith, IMS120031, DNA 2956.

PARATYPES (39 ♀; 26 ♂): Arkansas, USA: 1 ♀ from Newton County, Buffalo National River, Cecil Creek (36°5'15.72"N, 93°13'23.28"W), 13 Jun 2012, by TD Edwards, TDE 12-0613-010 • 1 ♂ from Newton County, Ozark-St. Francis National Forest, Little Buffalo River, 11 Jun 2012, by TD Edwards, TDE 12-0711-004 • 2 ♀ from Polk County, beside Forest Route 38, north of Shady Lake Recreation Area, East Saline Creek (34°22'23.39"N, 94°1'51.22"W), 30 Jul 2011, by IM Smith, IMS110041 • Kentucky, USA: 2 ♀ and 1 ♂ from McCreary County, White Oak Junction, Rock Creek, beside Forest Route 556, 2.3 kilometers south of Route 1363, 8 Jul 1990, by IM Smith, IMS900082A & IMS900082B • Illinois, USA: 3 ♀ and 1 ♂ from Kankakee County, Kankakee River State Park, beside Route 102, Rock Creek (41°12'N, 88°0'W), 9 Sep 1991, by IM Smith, IMS910027A • Indiana, USA: 1 ♀ from Wayne County, south of intersection of Interstate 70 and Route 1 (39°51'13"N, 85°8'4"W), 31 Jul 2014, by MJ Skvarla, MS 14-0731-001 • Missouri, USA: 3 ♀ Wayne County, beside Road 143 near Sam A Baker State Park, 8 Jul 1960, by DR Cook, DRC600010 • New Brunswick, Canada: 2 ♀ and 1 ♂ from Charlotte County, Rollingdam, Digdeguash River, beside Highway 770 at covered bridge, 30 Jun 1989, by IM Smith, IMS890053 • 1 ♀ and 1 ♂ from Charlotte County, Digdeguash River, beside Sorrel Ridge Road, west of Whittier Road, 10 Jun 2012, by IM Smith, IMS120015 • 2 ♀ and 1 ♂ from Charlotte County, Rollingham, Digdegaush River, beside Highway 770, 3 Oct 2011, by IM Smith, IMS110118 • 1 ♂ (ALLOTYPE) from York County, Stanley, Nashwaak River, Stanley Municipal Park, 19 Jun 2012, by IM Smith, IMS120031, DNA 2957 • Ohio, USA: 1 ♂ from Montgomery County, Engelwood Metro Park (39°52'58"N, 84°17'33"W), 31 Jul 2014, by MJ Skvarla, MS 14-0731-002 • 1 ♀ and 2 ♂ from Pickaway County, beside Scioto-Darby Road, just north of Darby, Big Darby Creek (40°2'N, 83°15'W), 30 Jun 1997, by IM Smith & M MacKenzie, IMS970016 • Ontario, Canada: 1 ♀ and 1 ♂ from Grey County, Durham, Saugeen River, beside County Road 27 near Durham Conservation Area, 9 Jun 1989, by IM Smith, IMS890028A • Pennsylvania, USA: 1 ♀ from Bedford County, Chaneysville, Sweet Root Picnic Area beside Route 326, 18 Jul 1990, by IM Smith, IMS900105 • 1 ♀ from Huntingdon County, Alan Seeger Natural Area, Stone Creek, beside road from McAlevys Fort to Route 322, 19 Jul 1990, by IM Smith, IMS900107 • 1 ♂ from Tioga County, Straight Run, beside Straight Run Road, 2.1 kilometers north of Route 6 between Ansonia & Wellsboro, 20 Jul1990, by IM Smith, IMS900111 • Saskatchewan, Canada: 2 ♀ and 2 ♂ from Torch River beside Highway 106 at access road, 30 kilometers north of Highway 55 at Smeaton, 30 Jul 1988, by IM Smith, IMS880054A • 2 ♀ and 2 ♂ from Torch River at end of forest access road, 10 kilometers north of Highway 55 at Love, 30 Jul 1988, by IM Smith, IMS880056 • Texas, USA: 2 ♀ from Bandera County, Vanderpool, beside Route 187, Sabinal River (29°48'10"N, 99°34'30"W), 2 May 2009, by IM Smith, IMS090007 • 1 ♀ and 1 ♂ from Kinney County, Brackettville, beside Route 90, 12.1 kilometers west of Route 131, Pinto Creek (29°20'6"N, 100°32'5"W), 28 Sep 1994, by IM Smith, IMS940025 • 1 ♀ from Uvalde County, Garner State Park, Frio River (29°35'22"N, 99°44'12"W), 28 May 1998, by IM Smith, IMS980027A • 1 ♀ and 1 ♂ from Val Verde County, Bakers Crossing Campground off Route 163, Devils River (29°58'N, 101°9'W), 5 Oct 1999, by IM Smith, IMS990061A • 2 ♀ from Val Verde County, Dolan Falls Preserve, Devils River (29°53'12"N, 100°59'37"W), 24 May 2011, by IM Smith, IMS110013 • 4 ♀ and 4 ♂ from Val Verde County, Dolan Falls Preserve, Snake Spring (29°53'43"N, 100°58'58"W), 25 May 2011, by IM Smith, IMS110015 • Virginia, USA: 1 ♀ and 1 ♂ from Bath County, Jackson River, beside Forest Route 1843 (continuation of Route 623), 3.5 kilometers south of Route 220, 16 Jul 1990, by IM Smith, IMS900100 • 2 ♀ and 2 ♂ from Scott County, North Fork Holston River, beside Route 58/421, 0.9 kilometers east of Route 709 at Hiltons, 7 Jul 1990, by IM Smith, IMS900080 • West Virginia, USA: 1 ♀ and 1 ♂ from Pendleton County, South Branch of the Potomac River, beside Route 28/55, 20.8 kilometers southwest of Route 42, 17 Jul 1990, by IM Smith, IMS900104.

Type deposition

Holotype (♀), allotype (♂), and most paratypes (34 ♀; 20 ♂) deposited in the CNC; other paratypes (5 ♀; 5 ♂) deposited in ACUA.

Diagnosis

Torrenticola irapalpa are similar to other members of the Raptor Group (T. gnoma, T. longitibia, T. mjolniri, T. elusiva, T. racupalpa, T. raptor, T. danielleae, T. daemon, and T. ivyae) in having round bodies; Dgl-4 close to muscles scars; long, thin subcapitular rostra; and long, thin pedipalp tibiae. T. irapalpa can be differentiated from T. longitibia, T. mjolniri, T. elusiva, T. racupalpa, and T. ivyae by having Dgl-4 closer to the dorsal edge (dorsal width/distance between Dgl-4 = 1.81–2.09 in T. irapalpa, 2.19–2.77 in others) and a less elongate rostrum (length/width = 2.66–3.39 in T. irapalpa; 3.56–4.32 in others). T. irapalpa can be differentiated from T. gnoma by having Dgl-4 closer to the dorsal edge (dorsal width/distance between Dgl-4 ♀ = 1.81–2.09 in T. irapalpa, 2.65–3.29 in T. gnoma; ♂ = 1.58–1.86 in T. irapalpa, 2.06–2.73 in T. gnoma) and by dorsal coloration and pattern. T. irapalpa can be differentiated from T. raptor by having less elongate tibiae (length/width ♀ = 4.09–5.67 in T. irapalpa, 6.00–7.54 in T. raptor; ♂ = 4.25–4.75 in T. irapalpa, 5.29–5.63 in T. raptor); less elongate subcapitulum (ventral length/height = 2.52–2.90 in T. irapalpa, 2.98–3.27 in T. raptor); and by dorsal pattern. T. irapalpa can be differentiated from T. danielleae by having Dgl-4 further from the dorsal edge (dorsal width/distance between Dgl-4 = ♀ = 1.81–2.09 in T. irapalpa, 1.57–1.70 in T. danielleae; ♂ = 1.58–1.86 in T. irapalpa, 1.42–1.52 in T. danielleae); a less elongate rostrum (length/width = 2.66–3.39 in T. irapalpa, 3.43–3.75 in T. danielleae); and by dorsal coloration and pattern. Female T. irapalpa can be differentiated from female T. daemon by having Dgl-4 further from the dorsal edge (dorsal width/distance between Dgl-4 ♀ = 1.81–2.09 in T. irapalpa, 1.59–1.67 in T. daemon) and a less elongate gnathosomal bay (length/width ♀ = 1.35–1.86 in T. irapalpa, 1.95–2.42 in T. daemon). Male T. irapalpa can be differentiated from male T. daemon by dorsal coloration and pattern.

Description

Female (Figure 103) (n = 9) (holotype measurements in parentheses when available) with characters of the genus with following specifications.

Dorsum — (575–710 (610) long; 465–580 (475) wide) circular with navy blue to bluish-purple coloration separated into anterior and posterior portions with bold orange medially. Anterio-medial platelets (132.5–160 (137.5) long; 52.5–67.5 (62.5) wide). Anterio-lateral platelets (155–197.5 (177.5) long; 67.5–87.5 (77.5) wide) free from dorsal plate. Dgl-4 midway between muscle scars and dorsum edge (distance between Dgl-4 465–580 (475)). Dorsal plate proportions: dorsum length/width 1.20–1.28 (1.28); dorsal width/distance between Dgl-4 1.81–2.09 (1.90); anterio-medial platelet length/width 2.12–2.52 (2.20); anterio-lateral platelet length/width 2.17–2.39 (2.29); anterio-lateral/anterio-medial length 1.17–1.42 (1.29).

Gnathosoma — Subcapitulum (300–360 (340) long (ventral); 219–270 (270) long (dorsal); 125–157.5 (145) tall) with faint navy blue to bluish-purple coloration, sometimes colorless. Rostrum (126.25–152.5 (142.5) long; 55–76 (65) wide) elongate. Chelicerae (245–360 (360) long) with curved fangs (55–75 (65) long). Subcapitular proportions: ventral length/height 2.29–2.57 (2.34); rostrum length/width 2.66–3.39 (3.17). Pedipalps elongate (especially tibiae) with long tuberculate ventral extensions on femora and genua. Palpomeres: trochanter (45–52.5 (45) long); femur (115–132.5 (132.5) long); genu (65–80 (72.5) long); tibia (100–117.5 (102.5) long; 17.5–20 (20) wide); tarsus (17.5–20 (20) long). Palpomere proportions: femur/genu 1.66–1.89 (1.83); tibia/femur 0.77–0.89 (0.77); tibia length/width 4.09–5.67 (4.82).

Venter — (690–870 (770) long; 508–743 (550) wide) with navy blue to bluish purple coloration. Gnathosomal bay (132.5–165 (165) long; 87.5–120 (90) wide). Cxgl-4 far from apex. Medial suture (10–22.5 (15) long). Genital plates (155–175 (175) long; 140–162.5 (150) wide). Additional measurements: Cx-1 (266–334 (310) long (total); 103–156 (140) long (medial)); Cx-3 (332–487 (375) wide); anterior venter (156–197.5 (182.5) long). Ventral proportions: gnathosomal bay length/width 1.35–1.86 (1.83); anterior venter/genital field length 0.99–1.18 (1.04); anterior venter length/genital field width 1.06–1.27 (1.22); anterior venter/medial suture 8.56–15.63 (12.17).

Male (Figure 104) (n = 8) (allotypic measurements in parentheses when available) with characters of the genus with following specifications.

Dorsum — (520–620 (560) long; 400–490 (435) wide) circular with navy blue to bluish-purple coloration separated into anterior and posterior portions with bold orange medially. Anterio-medial platelets (115–150 (140) long; 55–67.5 (62.5) wide). Anterio-lateral platelets (160–188.75 (187.5) long; 62.5–77.5 (75) wide) free from dorsal plate. Dgl-4 midway between muscle scars and dorsum edge (distance between Dgl-4 220–310 (250)). Dorsal plate proportions: dorsum length/width 1.26–1.30 (1.29); dorsal width/distance between Dgl-41.58–1.86 (1.74); anterio-medial platelet length/width 2.00–2.48 (2.24); anterio-lateral platelet length/width 2.34–2.60 (2.50); anterio-lateral/anterio-medial length 1.22–1.41 (1.34).

Gnathosoma — Subcapitulum (275–340 (310) long (ventral); 207–240 (240) long (dorsal); 105–132.5 (107.5) tall) with faint navy blue to bluish-purple coloration, sometimes colorless. Rostrum (112.5–137.5 (135) long; 40–47.5 (40) wide) elongate. Chelicerae (252–305 (305) long) with curved fangs (51–60 (55) long). Subcapitular proportions: ventral length/height 2.52–2.90 (2.88); rostrum length/width 2.81–3.38 (3.38). Pedipalps elongate (especially tibiae) with long tuberculate ventral extensions on femora and genua. Palpomeres: trochanter (35–50 (42.5) long); femur (107.5–123.75 (115) long); genu (60–70 (67.5) long); tibia (87.5–108.75 (95) long; 20–25 (20) wide); tarsus (18.75–20 (20) long). Palpomere proportions: femur/genu 1.70–1.83 (1.70); tibia/femur 0.81–0.91 (0.83); tibia length/width 4.25–4.75 (4.75).

Venter — (630–705 (690) long; 490–540 (500) wide) with navy blue to bluish purple coloration. Gnathosomal bay (135–177.5 (146.25) long; 75–95 (75) wide). Cxgl-4 far from apex. Medial suture (50–75 (75) long). Genital plates (127.5–158.75 (143.75) long; 100–137.5 (122.5) wide). Additional measurements: Cx-1 (260–290 (290) long (total); 137–155 (155) long (medial)); Cx-3 (323–360 (345) wide); anterior venter (215–255 (245) long). Ventral proportions: gnathosomal bay length/width 1.42–2.06 (1.95); anterior venter/genital field length 1.51–1.75 (1.70); anterior venter length/genital field width 1.78–2.20 (2.00); anterior venter/medial suture 3.27–4.40 (3.27).

Immatures unknown.

Etymology

Specific epithet (irapalpa) refers to the pedipalps of this species, which resemble ferocious weapons of wrath (ira, L. fury, rage, wrath; palpus, L. hand, feeler).

Distribution

Primarily eastern (Figure 102), but note that T. irapalpa is one of few Torrenticola found in the Great Plains, at least in the south (Texas) and north (Saskatchewan).

Figure 102. 

Torrenticola irapalpa sp. n. distribution.

Figure 103. 

Torrenticola irapalpa sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 104. 

Torrenticola irapalpa sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Remarks

Torrenticola irapalpa groups with other members of the Raptor Complex with high support and specimens of this species are less than 2% different in COI sequence from each other. In the combined analysis, T. irapalpa groups with T. gnoma with high support, but the position of this clade was not recovered. These species are greater than 9% from each other. Based upon overall similarity, distribution, and phylogenetic position, this species is placed within the Raptor Identification Group.

This species hypothesis is supported by low COI divergence within the species (0–2%) and high divergence between species (3–15%), and by the morphological characters outlined in the diagnosis.

Torrenticola ivyae Fisher & Dowling, sp. n.

Material examined

HOLOTYPE (♀): from USA, Florida, Polk County, beside Rt. 471 at junction of Pasco, Polk and Sumter County lines, (28°19'19"N, 82°4'4"W), 24 April 1992, by IM Smith, IMS920003

PARATYPES (5 ♀; 5 ♂): Florida, USA: 2 ♀ from Hillsborough County, beside Rt. 41/301 near Hillsborough River State Park, south of Zephyrhills, (28°10'10"N, 82°12'12"W), 24 April 1992, by IM Smith, IMS920001 • 2 ♀ and 3 ♂ from Pasco County, beside Rt. 39 just north of Crystal Springs, (28°11'11"N, 82°10'10"W), 27 April 1992, by IM Smith, IMS920011 • 1 ♂ (ALLOTYPE) from Polk County, beside Rt. 471 at junction of Pasco, Polk and Sumter County lines, (28°19'19"N, 82°4'4"W), 24 April 1992, by IM Smith, IMS920003 • 1 ♀ and 1 ♂ from Polk County, beside Rucks Dairy Road 1.3 km south of Lake Arbuckle Road, (27°42'42"N, 81°26'26"W), 25 April 1992, by IM Smith, IMS920004

Type deposition

Holotype (♀), allotype (♂), and some paratypes (3 ♀; 2 ♂) deposited in the CNC; other paratypes (2 ♀; 2 ♂) deposited in the ACUA.

Diagnosis

Torrenticola ivyae are similar to other members of the Raptor Group (T. gnoma, T. irapalpa, T. longitibia, T. mjolniri, T. elusiva, T. racupalpa, T. raptor, T. danielleae, and T. daemon) in having round bodies; Dgl-4 close to muscles scars; long, thin subcapitular rostra; and long, thin pedipalp tibiae. T. ivyae can be differentiated from T. gnoma, T. elusiva, T. danielleae, and T. daemon by having a more elongate rostrum (length/width ♀ = 4.00–4.15 in T. ivyae, 2.74-3.75 in others; ♂ = 3.85–4.08 in T. ivyae, 2.56–3.57 in others) and more elongate pedipalpal tibiae (length/width ♀ = 5.07–5.64 in T. ivyae, 4.05–5.00 in others, ♂ = 4.75–5.20 in T. ivyae, 3.88–4.44 in others). T. ivyae can be differentiated from T. irapalpa, T. raptor, T. danielleae, and T. daemon by having Dgl-4 closer to the muscle scars (dorsum width/distance between Dgl-4 = 2.20–2.75 in T. ivyae, 1.42–2.09 in others). T. ivyae can be differentiated from T. longitibia (known only from males) by femur/genu (1.83–1.88 in T. ivyae, 2.1–2.17 in T. longitibia) and having less elongate tibiae (length/width = 4.75–5.20 in T. ivyae, 5.50–5.50 in T. longitibia). Female T. ivyae can be differentiated from female T. racupalpa by having a more elongate rostrum (length/width = 4.00–4.15 in T. ivyae, 3.56–3.82 in T. racupalpa) and more elongate pedipalpal tibiae (length/width = 5.07–5.64 in T. ivyae, 4.44–5.00 in T. racupalpa). Male T. ivyae can be differentiated from male T. racupalpa by having a longer anterior venter (♂ 220–230 in T. ivyae, 200–205 in T. racupalpa) and a shorter genital field (♂ 142–148 in T. ivyae, 160–165 in T. racupalpa). Female T. ivyae can be differentiated from female T. mjolniri by having a smaller dorsum (length ♀ = 550–590 in T. ivyae, 605–640 in T. mjolniri; width ♀ = 460–500 in T. ivyae, 510–545 in T. mjolniri) and a shorter anterior venter (♀ 155–170 in T. ivyae, 180–195 in T. mjolniri). Male T. ivyae can be differentiated from male T. mjolniri by having a less elongate subcapitulum (ventral length/width = 2.57–2.75 in T. ivyae, 2.82–3.00 in T. mjolniri). Additionally, T. ivyae can be differentiated from T. mjolniri by being found in Florida (T. ivyae is known from the northeast).

Description

Female (Figure 106) (n = 5) (holotype measurements in parentheses when available) with characters of the genus with following specifications.

Dorsum — (550–590 (580) long; 460–500 (495) wide) circular with reddish-purple coloration anteriorly and posteriorly connected medially. Anterio-medial platelets (120–135 (120) long; 60–62.5 (62.5) wide). Anterio-lateral platelets (165–180 (165) long; 65–75 (75) wide) free from dorsal plate. Dgl-4 much closer to the muscle scars than to the edge of the dorsum (distance between Dgl-4 180–195 (180)). Dorsal plate proportions: dorsum length/width 1.16–1.20 (1.17); dorsal width/distance between Dgl-4 2.42–2.75 (2.75); anterio-medial platelet length/width 1.92–2.25 (1.92); anterio-lateral platelet length/width 2.20–2.62 (2.20); anterio-lateral/anterio-medial length 1.26–1.38 (1.38).

Gnathosoma — Subcapitulum (300–325 (325) long (ventral); 232.5–257.5 (247.5) long (dorsal); 115–127.5 (120) tall) colorless. Rostrum (130–145 (145) long; 32.5–35 (35) wide) elongate. Chelicerae (315–340 (335) long) with curved fangs (55–60 (60) long). Subcapitular proportions: ventral length/height 2.55–2.71 (2.71); rostrum length/width 4.00–4.15 (4.14). Pedipalps elongate with tuberculate ventral extensions on femora and genua. Palpomeres: trochanter (42.5–46.25 (45) long); femur (117.5–125 (125) long); genu (65–67.5 (65) long); tibia (91.25–100 (95) long; 17.5–18.75 (18.75) wide); tarsus (15–17.5 (17.5) long). Palpomere proportions: femur/genu 1.81–1.92 (1.92); tibia/femur 0.76–0.83 (0.76); tibia length/width 5.07–5.64 (5.07).

Venter — (630–755 (750) long; 490–540 (540) wide) with reddish-purple coloration. Gnathosomal bay (150–175 (165) long; 67.5–75 (72.5) wide). Cxgl-4 far from apex. Medial suture (25–25 (25) long). Genital plates (150–165 (157.5) long; 142.5–150 (150) wide). Additional measurements: Cx-1 (280–300 (300) long (total); 120–145 (145) long (medial)); Cx-3 (320–335 (325) wide); anterior venter (155–170 (170) long). Ventral proportions: gnathosomal bay length/width 2.00–2.44 (2.28); anterior venter/genital field length 0.94–1.13 (1.08); anterior venter length/genital field width 1.03–1.19 (1.13); anterior venter/medial suture 6.20–6.80 (6.80).

Male (Figure 107) (n = 4) (allotypic measurements in parentheses when available) with characters of the genus with following specifications.

Dorsum — (530–550 (550) long; 390–450 (420) wide) circular with reddish-purple coloration anteriorly and posteriorly connected medially. Anterio-medial platelets (125–135 (130) long; 55–62.5 (57.5) wide). Anterio-lateral platelets (160–190 (160) long; 60–75 (60) wide) free from dorsal plate. Dgl-4 much closer to the muscle scars than to the edge of the dorsum (distance between Dgl-4 165–205 (180)). Dorsal plate proportions: dorsum length/width 1.19–1.36 (1.31); dorsal width/distance between Dgl-4 2.20–2.39 (2.33); anterio-medial platelet length/width 2.00–2.45 (2.26); anterio-lateral platelet length/width 2.27–2.92 (2.67); anterio-lateral/anterio-medial length 1.23–1.46 (1.23).

Gnathosoma — Subcapitulum (295–302.5 (300) long (ventral); 220–230 (220) long (dorsal); 110–115 (110) tall) colorless. Rostrum (125–132.5 (130) long; 32.5–32.5 (32.5) wide) elongate. Chelicerae (295–305 (295) long) with curved fangs (45–50 (50) long). Subcapitular proportions: ventral length/height 2.57–2.75 (2.73); rostrum length/width 3.85–4.08 (4.00). Pedipalps elongate with tuberculate ventral extensions on femora and genua. Palpomeres: trochanter (40–42.5 (40) long); femur (110–112.5 (110) long); genu (60–60 (60) long); tibia (90–97.5 (90) long; 17.5–20 (17.5) wide); tarsus (15–16.25 (16.25) long). Palpomere proportions: femur/genu 1.83–1.88 (1.83); tibia/femur 0.82–0.87 (0.82); tibia length/width 4.75–5.20 (5.14).

Venter — (625–700 (700) long; 470–500 (470) wide) with reddish-purple coloration. Gnathosomal bay (140–150 (142.5) long; 62.5–72.5 (65) wide). Cxgl-4 far from apex. Medial suture (60–80 (75) long). Genital plates (142.5–147.5 (145) long; 112.5–125 (120) wide). Additional measurements: Cx-1 (280–290 (280) long (total); 125–145 (145) long (medial)); Cx-3 (310–330 (310) wide); anterior venter (220–230 (230) long). Ventral proportions: gnathosomal bay length/width 1.93–2.36 (2.19); anterior venter/genital field length 1.52–1.59 (1.59); anterior venter length/genital field width 1.76–2.00 (1.92); anterior venter/medial suture 2.75–3.67 (3.07).

Immatures unknown.

Etymology

Specific epithet (ivyae) named in honor of Ivy Fisher, our (JRF and DMF) beautiful daughter, who was born in Florida, the type locality.

Distribution

Florida (Figure 105).

Figure 105. 

Torrenticola ivyae sp. n. distribution.

Figure 106. 

Torrenticola ivyae sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 107. 

Torrenticola ivyae sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Remarks

Unfortunately, we were unable to acquire fresh material of Torrenticola ivyae and therefore this species is not included in our phylogenetic analyses. However, we were able to examine morphology with material preserved in GAW. The overall similarity, elongate subcapitular rostra, elongate pedipalpal tibiae, and Dgl-4 close to the muscle scars, are consistent with placing this species in the Raptor Complex and Raptor Identification Group.

Torrenticola karambita Fisher & Dowling, sp. n.

Material examined

HOLOTYPE (♀): from USA, Tennessee, Sevier County, Great Smokey Mountains National Park, Sugarlands Nature Trail (35°40'47"N, 83°31'51"W), 10 Sep 2010, by IM Smith, IMS100125, DNA 1758.

PARATYPES (1 ♀; 3 ♂): 1 ♂ (ALLOTYPE) from Sevier County, Great Smokey Mountains National Park, Sugarlands Nature Trail (35°40'47"N, 83°31'51"W), 10 Sep 2010, by IM Smith, IMS100125, DNA 1846 • 1 ♀ and 2 ♂ from Sevier County, Great Smokey Mountains National Park, Sugarlands Nature Trail (35°40'47"N, 83°31'51"W), 10 Sep 2010, by IM Smith, IMS100125

Type deposition

Holotype (♀), allotype (♂), and some paratypes (1 ♂) deposited in the CNC; other paratypes (1 ♀; 1 ♂) deposited in the ACUA.

Diagnosis

Torrenticola karambita is similar to other members of the Erectirostra Group (T. erectirostra and T. robisoni) in having an upturned rostrum that is wide when viewed ventrally. T. karambita can be differentiated from T. erectirostra and T. robisoni by lacking coloration (T. erectirostra has purplish dorsal coloration) and a slightly stockier rostrum (length/width ♀ = 1.57–1.62 in T. karambita, 1.72–2.09 in others; ♂ = 1.6–1.95 in T. karambita, 2.0–2.2 in T. erectirostra).

Description

Female (Figure 109) (n = 2) (holotype measurements in parentheses when available) with characters of the genus with following specifications.

Dorsum — (690–725 (690) long; 490–500 (490) wide) ovoid and colorless. Anterio-medial platelets (145–162.5 (145) long; 65–75 (65) wide). Anterio-lateral platelets (217.5–220 (217.5) long; 80–82.5 (82.5) wide) free from dorsal plate. Dgl-4 much closer to the edge of the dorsum than to the muscle scars (distance between Dgl-4 340–360 (340)). Dorsal plate proportions: dorsum length/width 1.41–1.45 (1.41); dorsal width/distance between Dgl-4 1.39–1.44 (1.44); anterio-medial platelet length/width 2.17–2.23 (2.23); anterio-lateral platelet length/width 2.64–2.75 (2.64); anterio-lateral/anterio-medial length 1.35–1.50 (1.50).

Gnathosoma — Subcapitulum (325–350 (325) long (ventral); 229–254 (229) long (dorsal); 135–135 (135) tall) colorless. Rostrum (110–117.5 (110) long; 70–72.5 (70) wide) wide and unturned with dentation. Chelicerae (320–329 (321) long) with curved fangs (40–52 (41) long). Subcapitular proportions: ventral length/height 2.41–2.59 (2.41); rostrum length/width 1.57–1.62 (1.57). Pedipalps short and stocky (especially tibiae) with short tuberculate ventral extensions on femora and genua. Palpomeres: trochanter (52.5–55 (52.5) long); femur (101.25–105 (101.25) long); genu (65–67.5 (67.5) long); tibia (55–57.5 (55) long; 30–30 (30) wide); tarsus (17.5–17.5 (17.5) long). Palpomere proportions: femur/genu 1.50–1.62 (1.50); tibia/femur 0.54–0.55 (0.54); tibia length/width 1.83–1.92 (1.83).

Venter — (850–850 (850) long; 595–607 (607) wide) colorless. Gnathosomal bay (170–172.5 (172.5) long; 125–130 (130) wide). Cxgl-4 far from apex. Medial suture (15–17.5 (15) long). Genital plates (195–197.5 (195) long; 165–175 (165) wide). Additional measurements: Cx-1 (336–348 (336) long (total); 159–175 (174) long (medial)); Cx-3 (415–441 (415) wide); anterior venter (202.5–222.5 (202.5) long). Ventral proportions: gnathosomal bay length/width 1.33–1.36 (1.33); anterior venter/genital field length 1.04–1.13 (1.04); anterior venter length/genital field width 1.23–1.27 (1.23); anterior venter/medial suture 12.71–13.50 (13.50).

Male (Figure 110) (n = 3) (allotypic measurements in parentheses when available) with characters of the genus with following specifications.

Dorsum— (610–655 (610) long; 410–440 (420) wide) ovoid and colorless. Anterio-medial platelets (133.75–145 (133.75) long; 65–67.5 (65) wide). Anterio-lateral platelets (205–217.5 (207.5) long; 75–77.5 (75) wide) free from dorsal plate. Dgl-4 much closer to the edge of the dorsum than to the muscle scars (distance between Dgl-4 290–330 (290)). Dorsal plate proportions: dorsum length/width 1.44–1.60 (1.45); dorsal width/distance between Dgl-4 1.33–1.45 (1.45); anterio-medial platelet length/width 2.06–2.19 (2.06); anterio-lateral platelet length/width 2.65–2.81 (2.77); anterio-lateral/anterio-medial length 1.44–1.55 (1.55).

Gnathosoma — Subcapitulum (285–290 (290) long (ventral); 202–210 (202) long (dorsal); 108.75–115 (112.5) tall) colorless. Rostrum (97.5–102.5 (97.5) long; 52.5–60 (60) wide) wide and unturned with dentation. Chelicerae (268–269 (269) long) with curved fangs (43–52 (47) long). Subcapitular proportions: ventral length/height 2.48–2.67 (2.58); rostrum length/width 1.63–1.95 (1.63). Pedipalps short and stocky (especially tibiae) with short tuberculate ventral extensions on femora and genua. Palpomeres: trochanter (47.5–48.75 (47.5) long); femur (87.5–95 (95) long); genu (57.5–62.5 (62.5) long); tibia (55–57.5 (57.5) long; 27.5–27.5 (27.5) wide); tarsus (15–17.5 (17.5) long). Palpomere proportions: femur/genu 1.44–1.52 (1.52); tibia/femur 0.61–0.63 (0.61); tibia length/width 2.00–2.09 (2.09).

Venter — (760–785 (760) long; 490–521 (512) wide) colorless. Gnathosomal bay (165–175 (165) long; 105–107.5 (107.5) wide). Cxgl-4 far from apex. Medial suture (70–77.5 (77.5) long). Genital plates (156.25–167.5 (157.5) long; 120–125 (125) wide). Additional measurements: Cx-1 (284–320 (284) long (total); 115–154 (116) long (medial)); Cx-3 (366–373 (372) wide); anterior venter (247.5–248.75 (248.75) long). Ventral proportions: gnathosomal bay length/width 1.53–1.67 (1.53); anterior venter/genital field length 1.48–1.58 (1.58); anterior venter length/genital field width 1.99–2.06 (1.99); anterior venter/medial suture 3.19–3.54 (3.21).

Immatures unknown.

Etymology

Specific epithet (karambita) refers to the upturned rostrum that has dentation on both sides in females, which resembles a karambit—small, recurved knives used in the Filipino martial arts practiced by JRF. The karambit is thought to have originated with the Minangkabau people of West Sumatra based upon a similarity to a tiger’s claws.

Distribution

Known only from Great Smokey Mountains National Park, Sevier County, Tennessee (Figure 108).

Figure 108. 

Torrenticola karambita sp. n. distribution.

Figure 109. 

Torrenticola karambita sp. n. female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 110. 

Torrenticola karambita sp. n. male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Remarks

Torrenticola karambita groups with other members of the Raptor Complex with high support in all analyses and specimens are less than 2% different in COI sequence from each other. In the combined analysis, T. karambita groups with two other species that share the modified subcapitulum with upturned rostrum: T. erectirostra and T. robisoni. These species are greater than 9% different from each other. Based upon overall similarity, distribution, and phylogenetic position, these species are placed within the Erectirostra Identification Group.

This species hypothesis is supported by low COI divergence within the species (0–2%), high divergence between species (3–15%), and the morphological characters outlined in the diagnosis.

Torrenticola keesdavidsi Cramer, 1992

Torrenticola keesdavidsi Cramer, 1992: 17.

Material examined

(6 ♀; 10 ♂) . Arizona, USA: 1 ♀ and 1 ♂ from Cochise County, Chiricahua Mountains; beside Forest Road 42 near junction with Forest Road 42B, (31°55'55"N, 109°15'15"W), 16 July 1987, by IM Smith, IMS870093A • 1 ♀ and 1 ♂ from Cochise County, Chiricahua Mountains; Sycamore Campground east of Sunizona, (31°52'52"N, 109°20'20"W), 15 July 1987, by IM Smith, IMS870091 • 3 ♂ from Cochise County, Chiricahua Mountains; Cave Creek Recreation Area; John Hands Picnic Area off Forest Road 42A west of Portal, (31°53'53"N, 109°13'13"W), 15 July 1987, by IM Smith, IMS870092A • 1 ♀ and 1 ♂ from Coconino County, Oak Creek Canyon; beside Rt. 89A just north of Pine Flat Campground, (35°1'1"N, 111°44'44"W), 21 July 1987, by IM Smith, IMS870099A • 1 ♀ from Coconino County, Oak Creek Canyon; beside Rt. 89A between Bootlegger and Banjo Bill Campgrounds, (34°58'58"N, 111°45'45"W), 21 July 1987, by IM Smith, IMS870100B • 1 ♂ from Coconino County, Oak Creek Canyon; beside Rt. 89A just north of Pine Flat Campground, (35°1'1"N, 111°44'44"W), 21 July 1987, by IM Smith, IMS870099B • New Mexico, USA: 2 ♀ from Catron County, Glenwood; Whitewater Picnic Area 8 km east of Rt. 180, (33°22'22"N, 108°50'50"W), 12 July 1987, by IM Smith, IMS870084 • 2 ♂ from Catron County, beside Rt. 15, 65 km north of Rt. 180 (Silver City), (33°12'12"N, 108°13'13"W), 10 July 1987, by IM Smith, IMS870081A • 1 ♂ from Catron County, beside Rt. 15, 65 km north of Rt. 180 (Silver City), (33°12'12"N, 108°13'13"W), 10 July 1987, by IM Smith, IMS870081B.

Type deposition

Holotype (♀) and allotype (♂) deposited in coll. Cristina Cramer, Instituto de Biología, UNAM.

Diagnosis

Torrenticola keesdavidsi are similar to other members of the Rala Group (T. rala, T. lamellipalpis, T. boettgeri, T. kurtvietsi, T. dolichodactyla, and T. anoplopalpa) by being colorless, having incomplete hind coxal margins and being distributed in the southwest. T. keesdavidsi can be differentiated from all other Rala Group by having a shorter dorsum (length ♀ = 555–605 in T. keesdavidsi, 630–800 in others; ♂ = 500–590 in T. keesdavidsi, 600–780 in others) and dentate, flanged ventral extensions on the femora (others are lacking extensions, have tuberculate extensions, or flat, wide lamellate extensions). Additionally, T. keesdavidsi can be differentiated from all other Rala Group by having more elongate pedipalpal tibiae (length/width = 4.50–5.20 in T. keesdavidsi, 1.75–3.38 in others), except T. lamellipalpis (4.32–4.94).

Re-description

Female (Figure 112) (n = 5) with characters of the genus with following specifications.

Dorsum — (555–605 long; 450–495 wide) circular and colorless. Anterio-medial platelets (135–137.5 long; 50–60 wide). Anterio-lateral platelets (180–192.5 long; 70–80 wide) free from dorsal plate. Dgl-4 closer to the edge of the dorsum than to the muscle scars (distance between Dgl-4 315–335). Dorsal plate proportions: dorsum length/width 1.21–1.31; dorsal width/distance between Dgl-4 1.41–1.48; anterio-medial platelet length/width 2.25–2.70; anterio-lateral platelet length/width 2.38–2.75; anterio-lateral/anterio-medial length 1.33–1.43.

Gnathosoma — Subcapitulum (312.5–327.5 long (ventral); 222.5–240 long (dorsal); 97.5–105 tall) colorless. Rostrum (120–130 long; 35–37.5 wide). Chelicerae (295–310 long) with curved fangs (45–50 long). Subcapitular proportions: ventral length/height 3.07–3.21; rostrum length/width 3.33–3.57. Pedipalps with dentate, flanged ventral extensions on femora and genua. Palpomeres: trochanter (35–37.5 long); femur (105–110 long); genu (55–57.5 long); tibia (90–95 long; 18.75–20 wide); tarsus (15–17.5 long). Palpomere proportions: femur/genu 1.91–2.00; tibia/femur 0.84–0.88; tibia length/width 4.50–4.93.

Venter — (695–755 long; 510–560 wide) colorless. Gnathosomal bay (145–150 long; 67.5–80 wide). Cxgl-4 far from apex. Medial suture (35–55 long). Genital plates (162.5–175 long; 152.5–160 wide). Additional measurements: Cx-1 (285–295 long (total); 140–150 long (medial)); Cx-3 (320–330 wide); anterior venter (192.5–222.5 long). Ventral proportions: gnathosomal bay length/width 1.88–2.19; anterior venter/genital field length 1.18–1.29; anterior venter length/genital field width 1.26–1.44; anterior venter/medial suture 4.05–5.50.

Male (Figure 113) (n = 5) with characters of the genus with following specifications.

Dorsum — (500–590 long; 410–450 wide) circular and colorless. Anterio-medial platelets (120–130 long; 55–60 wide). Anterio-lateral platelets (165–190 long; 65–75 wide) free from dorsal plate. Dgl-4 closer to the edge of the dorsum than to the muscle scars (distance between Dgl-4 280–320). Dorsal plate proportions: dorsum length/width 1.21–1.33; dorsal width/distance between Dgl-4 1.40–1.48; anterio-medial platelet length/width 2.09–2.27; anterio-lateral platelet length/width 2.33–2.71; anterio-lateral/anterio-medial length 1.32–1.48.

Gnathosoma — Subcapitulum (280–310 long (ventral); 202.5–230 long (dorsal); 85–105 tall) colorless. Rostrum (107.5–125 long; 32.5–35 wide). Chelicerae (260–297.5 long) with curved fangs (40–50 long). Subcapitular proportions: ventral length/height 2.81–3.29; rostrum length/width 3.29–3.57. Pedipalps with dentate, flanged ventral extensions on femora and genua. Palpomeres: trochanter (32.5–35 long); femur (97.5–110 long); genu (50–56.25 long); tibia (87.5–97.5 long; 17.5–20 wide); tarsus (15–17.5 long). Palpomere proportions: femur/genu 1.84–1.96; tibia/femur 0.86–0.95; tibia length/width 4.81–5.20.

Venter — (630–720 long; 460–510 wide) colorless. Gnathosomal bay (130–152.5 long; 60–75 wide). Cxgl-4 far from apex. Medial suture (80–95 long). Genital plates (140–155 long; 115–125 wide). Additional measurements: Cx-1 (260–280 long (total); 130–150 long (medial)); Cx-3 (280–340 wide); anterior venter (225–260 long). Ventral proportions: gnathosomal bay length/width 1.93–2.17; anterior venter/genital field length 1.55–1.86; anterior venter length/genital field width 1.96–2.17; anterior venter/medial suture 2.61–2.89.

Immatures unknown.

Etymology

Cramer (1992) named this species in honor of Kees Davids “as an acknowledgement of his great contributions to acarology and aquatic ecology” (translated from Spanish).

Distribution

Southwest (Arizona and New Mexico), extending southward into Mexico (Figure 111).

Figure 111. 

Torrenticola keesdavidsi distribution. Blue star represents type locality (Cramer 1992); and red circles represent new records and material examined.

Figure 112. 

Torrenticola keesdavidsi female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 113. 

Torrenticola keesdavidsi male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Remarks

Unfortunately, we were unable to acquire fresh material of Torrenticola keesdavidsi and therefore this species is not included in our phylogenetic analyses. We were also unable to examine type material. However, we were able to examine new material from Arizona and New Mexico that fits well within the species hypothesis proposed by Cramer (1992). The overall appearance, incomplete hind coxal margins, distribution, and lack of color place this species in the Rala Group.

Torrenticola kittatinniana Habeeb, 1955

Torrenticola kittatinniana Habeeb, 1955: 2.

Material examined

HOLOTYPE (♂): from USA, New Jersey, Sussex County, Little Flatbrook, north of Bevans, 12 Oct 1953, by H Habeeb, HH530110.

PARATYPES (1 ♀ and 0 ♂): New Jersey, USA: 1 ♀ (ALLOTYPE) from Morris County, Brook, Brookside, 20 May 1953, by H Habeeb, HH530045.

Type deposition

Holotype (♀) and paratypes (1 ♀) deposited in the CNC.

Diagnosis

Torrenticola kittatinniana are similar to other members of the Rusetria “4-Plate” group (T. dunni, T. glomerabilis, T. pollani, T. rufoalba and T. shubini) and T. skvarlai in having anterio-lateral platelets free from the dorsal plate, dorsal coloration separated into anterior and posterior portions, and indistinct hind coxal margins. Female T. kittatinniana can be differentiated from female T. dunni by having shorter pedipalpal genua (64 in T. kittatinniana, 70–75 in T. dunni); a shorter subcapitulum (ventral length = 310 in T. kittatinniana, 330–355 in T. dunni); and stockier anterio-medial platelets (length/width = 2.83 in T. kittatinniana, 2.33–2.54 in T. dunni). Male T. kittatinniana can be differentiated from male T. dunni by having a shorter anterior venter (235 in T. kittatinniana, 277–285 in T. dunni) and thinner dorsum (340 in T. kittatinniana, 350–370 in T. dunni). T. kittatinniana can be differentiated from T. pollani by having a stockier rostrum (length/width = 2.71–3.16 in T. kittatinniana, 3.27–3.82 in T. pollani) and stockier tibiae (length/width ♀ = 3.3 in T. kittatinniana, 3.8–4.2 in T. pollani; ♂ = 2.80 in T. kittatinniana, 3.4–3.8 in T. pollani). Female T. kittatinniana can be differentiated from female T. shubini by having a more elongate rostrum (length/width = 3.16 in T. kittatinniana, 2.5–2.7 in T. shubini) and a shorter subcapitulum (125 in T. kittatinniana, 140–145 in T. shubini). Male T. kittatinniana can be differentiated from male T. shubini by having a longer dorsum (500 in T. kittatinniana, 400–465 in T. shubini) and a longer genital field (115 in T. kittatinniana, 90–108 in T. shubini). T. kittatinniana can be differentiated from T. glomerabilis by having Dgl-4 closer to the dorsal edge (dorsal width/distance between Dgl-4 = 1.2–1.4 in T. kittatinniana, 1.5–1.7 in T. glomerabilis) and stockier tibiae (length/width ♀ = 3.3 in T. kittatinniana, 4.11–4.5 in T. glomerabilis; ♂ = 2.8 in T. kittatinniana, 3.5–4.4 in T. glomerabilis). T. kittatinniana can be differentiated from T. rufoalba by having a longer dorsum (♀ = 640 in T. kittatinniana, 550 in T. rufoalba; ♂ = 500 in T. kittatinniana, 440 in T. rufoalba) and more elongate anterio-medial platelets (length/width = 2.83–2.88 in T. kittatinniana, 2.45–2.61 in T. rufoalba). Additionally, male T. kittatinniana have a longer anterior venter (235 in T. kittatinniana, 195 in T. rufoalba). T. kittatinniana can be differentiated from T. skvarlai by having a conical pedipalpal femoral tubercle, whereas T. skvarlai has a broad and flat pedipalpal femoral tubercle, and by having a longer anterior venter (♀ = 165 in T. kittatinniana, 140–152.5 in T. skvarlai; ♂ = 235 in T. kittatinniana, 177.5–205 in T. skvarlai).

Re-description

Female (Figure 115) (n = 1) (allotype only) with characters of the genus with following specifications.

Dorsum — (550 long; 400 wide) ovoid with purple coloration separated into anterior and posterior portions bordered with orange. Anterio-medial platelets (107.5 long; 41.25 wide). Anterio-lateral platelets (168.75 long; 55 wide) free from dorsal plate. Dgl-4 much closer to the edge of the dorsum than to the muscle scars (distance between Dgl-4 255). Dorsal plate proportions: dorsum length/width 1.38; dorsal width/distance between Dgl-4 1.57; anterio-medial platelet length/width 2.61; anterio-lateral platelet length/width 3.07; anterio-lateral/anterio-medial length 1.57.

Gnathosoma— Subcapitulum (310 long (ventral); 235 long (dorsal); 127.5 tall) colorless. Rostrum (130 long; 42.5 wide). Chelicerae (315 long) with curved fangs (62.5 long). Subcapitular proportions: ventral length/height 2.43; rostrum length/width 3.06. Pedipalps with tuberculate ventral extensions on femora and genua. Palpomeres: trochanter (42.5 long); femur (115 long); genu (65 long); tibia (87.5 long; 25 wide); tarsus (17.5 long). Palpomere proportions: femur/genu 1.77; tibia/femur 0.76; tibia length/width 3.50.

Venter — (640 long; 450 wide) mostly colorless with faint purple coloration in areas surrounding coxae. Gnathosomal bay (142.5 long; 92.5 wide). Cxgl-4 subapical. Medial suture (17.5 long). Genital plates (167.5 long; 155 wide). Additional measurements: Cx-1 (125 long (total); 125 long (medial)); Cx-3 (335 wide); anterior venter (155 long). Ventral proportions: gnathosomal bay length/width 1.54; anterior venter/genital field length 0.93; anterior venter length/genital field width 1.00; anterior venter/medial suture 8.86.

Male (Figure 116) (n = 1) (holotype only) with characters of the genus with following specifications.

Dorsum — (500 long; 340 wide) ovoid with purple coloration separated into anterior and posterior portions bordered with orange. Anterio-medial platelets (90 long; 31.25 wide). Anterio-lateral platelets (140 long; 47.5 wide) free from dorsal plate. Dgl-4 much closer to the edge of the dorsum than to the muscle scars (distance between Dgl-4 280). Dorsal plate proportions: dorsum length/width 1.47; dorsal width/distance between Dgl-4 1.21; anterio-medial platelet length/width 2.88; anterio-lateral platelet length/width 2.95; anterio-lateral/anterio-medial length 1.56.

Gnathosoma — Subcapitulum (237.5 long (ventral); 180 long (dorsal); 100 tall) colorless. Rostrum (95 long; 35 wide). Chelicerae (222.5 long) with curved fangs (45 long). Subcapitular proportions: ventral length/height 2.38; rostrum length/width 2.71. Pedipalps with tuberculate ventral extensions on femora and genua. Palpomeres: trochanter (37.5 long); femur (87.5 long); genu (57.5 long); tibia (70 long; 25 wide); tarsus (17.5 long). Palpomere proportions: femur/genu 1.52; tibia/femur 0.80; tibia length/width 2.80.

Venter — (600 long; 435 wide) mostly colorless with faint purple coloration in areas surrounding coxae. Gnathosomal bay (107.5 long; 72.5) wide). Cxgl-4 subapical. Medial suture (102.5 long). Genital plates (115 long; 82.5 wide). Additional measurements: Cx-1 (235 long (total); 125 long (medial)); Cx-3 (285 wide); anterior venter (235 long). Ventral proportions: gnathosomal bay length/width 1.48; anterior venter/genital field length 2.04; anterior venter length/genital field width 2.85; anterior venter/medial suture 2.29.

Immatures unknown.

Etymology

Habeeb (1955) did not specify an explanation for the specific epithet (kittatinniana). However, we suspect it refers to the type locality, which is located in the Great Kittatinny Valley, which is named for Kittatinny Mountain in northwestern New Jersey, a northern extension of the Appalachian Ridge and Valley province.

Distribution

Northern New Jersey (Figure 114).

Figure 114. 

Torrenticola kittatinniana distribution.

Figure 115. 

Torrenticola kittatinniana female: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Figure 116. 

Torrenticola kittatinniana male: A dorsal plates B venter (legs removed) C subcapitulum D pedipalp (setae not accurately depicted). Scale = 100 µm.

Remarks

Unfortunately, we were unable to acquire more specimens of Torrenticola kittatinniana and therefore this species is not included in our phylogenetic analyses. We were able to examine the type specimens.