Corresponding author: Marshal Hedin (
Academic editor: Sarah Crews
This revision is based on sampling efforts over the past three decades in the southern Appalachian Mountains which have provided
Hedin M, Milne MA (2023) New species in old mountains: integrative taxonomy reveals ten new species and extensive short-range endemism in
Systematists and evolutionary biologists have long been interested in mountains. Mountains function as habitat islands, serve as refugia in the face of climatic variation, and generate ecological gradients (
The spider genus
This revision focuses more specifically, but not exclusively, on montane Appalachian
In his 1984 revision of North American nesticid spiders,
We acknowledge that the land upon which we searched for and collected specimens is the traditional and ancestral territories of the Calicuas, Cheraw, Chickasaw, Eno, Kaskaskia, Keyauwee, Lumbee, Manahoac, Miccosukee, Monacan, Moneton, Mvskoke (Muscogee), Myaamia, Occaneechi, Osage, Pee Dee, Saponi, S’atsoyaha (Yuchi), Shakori, Shawandasse Tula (Shawanwaki / Shawnee), Sissipahaw, Skaruhreh / Tuscarora, Sugaree, Tsalaguwetiyi (Cherokee), Waxhaw, Yesan (Tutelo), and Yį Įsuwą (Catawba) peoples. Most specimens used in this revision were obtained from collections made during the past 25 years by the authors, many collaborators, and prior students of the first author (see Acknowledgements). The types of all previously described taxa were loaned from the
Geographic location data were taken in the field using a global positioning system (
We identified immature specimens using the following guidelines: 1) If immatures were collected in association with adults from the same geographic location and in the same microhabitats, these specimens were attributed to the same species, reflecting a very low probability of syntopy (three locations of > 450 unique collecting events; Suppl. material
Some authors have divided species delimitation into a two-step process (
One caveat to our morphology-first approach is that we expect some morphological variation within species, and the distinction between geographic variation vs. species-level divergence is not obvious using only a qualitative approach (e.g., vs. conducting morphometrics and statistical analyses). We expected morphological variation within species because the habitats occupied by these spiders are naturally fragmented (e.g., populations found in caves, isolated mountain ranges, talus fields within mountain ranges, etc.), and the spiders themselves are dispersal-limited. For example,
This revision shows that female epigynal morphology is generally (but not always) more conserved within groups of closely related taxa. This impacted our revisionary research because adult males were not always available from all collecting locations. In these cases, our species assignments for female-only locations were less confident, and sometimes relied more heavily on geographic and/or genetic (a posteriori) evidence.
To formally test or validate morphological hypotheses using independent character evidence we gathered phylogenomic-scale
Genomic DNA was extracted from leg tissues using the DNeasy Kit (Qiagen). At least 200 ng was sent to RAPID genomics for
Using individual
A species tree was also estimated under a multispecies coalescent model using ASTRAL v. 5.7.8 (
We generated mitochondrial sequences for 218 specimens using standard polymerase chain reaction (
For specimens for which we had
Combined Sanger and
We defined species under an integrative species delimitation framework as follows: “single populations or sets of populations that share diagnostic male palpal morphologies and that are supported by nuclear phylogenomic monophyly”. This definition includes some necessary caveats. First, our nuclear sample is representative but obviously not exhaustive. We did not generate
While mitochondrial evidence was sometimes useful in
Geography also played a secondary role in species delimitation because almost all
Standard terminology used to describe male and female genitalic morphology follows
Graphical overview of terminology used to describe genitalic morphology. Tennessee, Overton Co., Obe Lee Cave,
With regards to epigynal morphology, the internal fertilization and copulatory ducts are difficult to visualize in
Because adult body size, leg lengths, and carapace / abdominal color patterns were found to be variable both within and among populations of the same species (see examples below), we generally do not comment upon this variation in the species descriptions below. Instead, intraspecific variation in male and female genitalia (if present) is emphasized.
Holotype and paratype specimens have been deposited at the Bohart Museum of Entomology (
The following character abbreviations are used in our species descriptions:
Ink drawings of male and female genitalia were made by Nadine Dupérré. A digital camera attached to a stereomicroscope was used to capture images, which were then enlarged and printed. A tracing of this printed image was detailed and shadowed with repeated reference to the specimen under a microscope. Epigyna were removed and cleared with lactic acid prior to illustration. The left palp of male spiders was illustrated in all cases.
Specimens were digitally imaged at
The total morphological sample considered is summarized in Suppl. material
Based on the examination of male and female morphology we hypothesized the following new species
For species delimitation of
We gathered original
Concatenated maximum likelihood and coalescent-based ASTRAL analyses, with very different analytical assumptions, largely agree on overall tree structure (Figs
Mitochondrial outgroup relationships are as recovered in the
For the nine new species discovered by examination of patterns of male palpal morphology (see above), six are strongly supported (validated) by nuclear gene tree monophyly and associated support metrics (ML bootstrap, concordance factors, ASTRAL quartet scores, and ASTRAL local posterior probabilities; see Fig.
Summary species tree (from
The nuclear phylogenomic data also mostly strongly supported previously described species. We note that none of these prior hypotheses have ever been tested (validated) using independent data, as is true for almost all described spider species (see
Patterns of genitalic variation across populations within what we considered as single species are summarized in the Taxonomy section for each species. These patterns of variation are based on much larger sample sizes than previously considered, so most are newly reported here.
Below we briefly describe patterns of niche conservatism in Appalachian
Contrasting with patterns of intraspecific genetic divergence, the Appalachian
We hypothesize that niche conservatism, and perhaps interactions with competing
The Appalachian
The taxonomy presented below is structured to follow phylogenomic results, including species groups (treating the
Map of all type localities. Type locality for
Both
Males within this species group possess palps with a forked tegular apophysis, including a larger, darkened distal tegular apophysis that extends underneath the median apophysis and a pointed basal tegular apophysis (Figs
Species in this species group are distributed in caves on the Cumberland Plateau, and southwards to suitable surface microhabitats at Talladega Mountain in east-central Alabama (Fig.
Distribution of the
Morphological diagnosis as in
This highly troglomorphic taxon is only known from a few caves on the western margin of the Cumberland Plateau in Overton County, north-central Tennessee (Fig.
Morphological diagnosis as in
This troglomorphic taxon is known only from three geographically adjacent caves on Pigeon Mountain in Walker County, Georgia (Fig.
Easily distinguished from other members of the
No significant genitalic variation was noted in the material examined.
No significant genitalic variation was noted in the material examined.
Known only from Rainbow Cave, located near Pocket Creek, a tributary to the Little Sequatchie River (Fig.
The specific name is a matronym in honor of N. K. Jemisin whose ‘Broken Earth’ book series features a subterranean colony, including scientists who study caves.
The only
No significant male or female genitalic variation was noted in the material examined.
All known records are from high elevation habitats (most above 600 m, but as low as 430 m) on Talladega Mountain, east-central Alabama (Fig.
This species has been collected in dark, cool, relatively moist near-surface habitats. For example, field notes from 1992 collections north of Bald Rock indicate that spiders were collected from “below bluffs on a steep hillside”, in “talus with a heavy leaf litter cover”, where spiders were “most abundant under large rocks close to the surface”. This situation compares favorably with the original “heavy talus of ravine” collections made by A.F. Archer, and the 1995 Hedin and Dellinger collections, most of which were made in north-facing talus, although at least one collection was from southwest-facing talus. This species is perhaps not as uncommon as previously believed and (in the late 1990s) was found consistently in suitable microhabitats.
We view Gertsch’s drawing of the male conductor (fig. 129) as inaccurate (compare to Fig.
Morphological diagnosis as summarized in
The Hurricane Maze Cave specimen is similar to specimens from the type locality, possessing a short and wide epigynum with a posteriorly flaring median septum and banana-shaped spermathecae with narrow bases, lying just lateral to fovea but inside of the sclerotized epigynal outline (Fig.
Previously known only from the type locality in southeastern Tennessee (
This species group is recovered as monophyletic with both nuclear (Figs
Male and female genital morphology suggests common ancestry for this complex of nine species, also defined as a species group by Gertsch (originally not including
Comparative ♂ palps of
The
Distribution of
Morphologically very similar to geographically parapatric
Minimal palpal variation was observed for males from three sample locations, the dorsal paracymbial process in a single Rocky Bluff male being slightly wider and shorter (Fig.
Originally recorded from three locations (
Strong phylogeographic structuring is observed in the mitochondrial data with a well-supported subclade found east of the Pigeon River (FieTop, Hebo Mtn, Rocky Bluff, etc.; Fig.
As an example of natural history, one male and 12 females were collected from rocky void spaces in a moist, rocky ravine near Rocky Bluff campground (
This species is strongly supported as sister to remaining members of the
As discussed above, this species is morphologically most similar to geographically adjacent
Males from different sample locations vary in the presence / absence of a small basal projection of the dorsal process (Fig.
Females from different sample locations vary in the symmetry of the interior epigynal plates (Fig.
Found in rocky microhabitats from the rugged mountains of the eastern Great Smoky Mountains National Park, and adjacent eastern and southern locations (Fig.
Along the Maddron Bald (along Indian Camp Creek) and the Low Gap to Mt. Cammerer trails (
The species epithet (
This species is strongly supported as sister to remaining members of the
Strongly supported by both mitochondrial and
This species exhibits interesting variation in somatic morphology. Specimens from Pond Cave are pale and long-legged with reduced eye pigmentation, specimens from Burton’s Cave and Alley Cave are pale and long-legged but with well-developed eyes, and specimens from the surface Brumley Creek population exhibit an “epigean” habitus, with dark abdomens, shorter legs, and well-developed eyes (Fig.
Known only from a small area in southwestern Virginia, in the upper Clinch River drainage basin (Fig.
The variation in degree of troglomorphy within this single species suggests that this character suite (eye development, pigmentation, leg length, etc.) can evolve relatively rapidly, as seen in other cave spider taxa (e.g.,
Strongly supported by both mitochondrial and
No noteworthy variation in male or female genitalia was found across sample locations.
Known only from a small area of the Appalachian Valley and Ridge in southwestern Virginia and adjacent eastern Tennessee (Fig.
Specimens from
The shape of the tegular apophysis varies slightly across sample locations. Specimens from most locations (similar to type material from Indian Cave) possess a tegular apophysis with a broad, L-shaped base and an acute tip (see Gertsch, 1984: fig. 58), whereas other specimens have a narrower base with a gradually tapering tip (Fig.
Known from both limestone caves (both shallow and deeper situations) and dark, cool, relatively moist near-surface habitats (e.g., rock piles, shallow cliff caves). Most known populations are from caves in the upper-central Appalachian Valley and Ridge, with a few peripheral montane surface populations (e.g., Surry County, NC; Carter County, TN; Raleigh County, WV). The southeastern Surry and Carter County populations appear disjunct, separated from the remainder of the species’ range by regions occupied by other taxa in the species group (Fig.
A close morphological and genetic relative of
The female specimen from Starve Rock Cave has an epigynum very similar to specimens from the type locality.
This troglomorphic taxon was previously known only from the type locality (Grassy Creek Cave), but is now known from two nearby caves in east-central Tennessee (Fig.
Sister to
Males may be differentiated from other members of this species group by the combination of palps with a paracymbium with a wide, broad ventral process, the paradistal paracymbial process broad and triangular, a median apophysis that is a thin rectangle with an anterior sclerotized point, and a broad, singularly-pointed tegular apophysis that extends to ~ half the length of the median apophysis (Fig.
In males from different sample locations the distal tip of the tegular apophysis varies in shape from blunt (e.g., Grandfather Mtn, Edgemont Rd) to more fingerlike (e.g., N Linville Caverns, China Creek, Elk River Cave, etc.). This variation does not obviously follow geographic or phylogeographic (see below) lines. Females from different sample locations are relatively conservative in epigynal morphology (Fig.
Fig.
Previously known only from caves, but quite common and abundant in suitable near-surface habitats. Mostly from the uplands between the Linville and Grandfather Mountains of western North Carolina, northeast of the Asheville Basin (Fig.
Strong phylogeographic structuring is observed in the mitochondrial data, with a well-supported subclade found east of the Linville Gorge (China Creek, Green Mountain, Elk River Cave, Rockhouse Creek, etc.; Fig.
This species is supported as sister to
Males are diagnosed from closely-related
There is minor variation in the depth of the distally bifurcate tegular apophysis (e.g., slightly deeper in neighboring Pigeonroost Creek and Rock Creek Recreation Areas specimens), but in general populations are conspicuously homogeneous despite a large and fragmented geographic distribution (Fig.
Most sample locations are from limestone caves in the central part of the upper Tennessee River valley, near Knoxville, Tennessee, and extending northeast and southwest from there (Fig.
We identified spiders from Sensabaugh Saltpeter Cave (3 imm) and ‘Cave by Clinch River’ (one ♀) as
As discussed directly below
Because neither female morphology nor mitochondrial placement can strictly distinguish
Male
Adult males from multiple collection events, including the lower elevation Henson Creek specimens, all closely approximate the holotype male. The distal portion of the tegular apophysis for the male from upper Roan Valley (
The epigyna of females from multiple locations closely approximate the paratype female.
Restricted to Roan Mountain and immediate vicinity at elevations near or above 1800 meters, except for the Henson Creek location (~ 900 meters) on the southeastern flanks of Roan Mountain (Fig.
We have collected comprehensively in this region, finding the sister species
Named after the highlands of Roan Mountain along the North Carolina / Tennessee border.
While male morphological evidence clearly supports this species as distinct in a “morphology first” framework (unique forked base of tegular apophysis), the
Mitochondrial data fail to support
Overall, this taxonomic situation illustrates patterns of nuclear vs. mitochondrial vs. morphological discordance as also found elsewhere in Appalachian
A species group strongly supported by nuclear phylogenomics (Figs
Comparative ♂♀ genitalia of
Species of the
Distribution of
Males may be distinguished from other members of the species group by the palp with a uniquely shaped tegular apophysis (except in comparison to
Notable variation exists in the shape of the dorsal process of the paracymbium, which is sometimes narrow and finger-like (
Epigyna vary across sample locations in the length of the projection of the median septum, the shape of the epigynal pockets (though generally spherical), the width of epigynal pocket lateral hoods, and the length of the spermathecae (Fig.
Previously known only from two locations but now known to be reasonably widespread in the Great Balsam and Pisgah Mountains southwest of Asheville North Carolina, west of the Asheville Basin (Fig.
No obvious phylogeographic trends are apparent in the mitochondrial data, with geographically separate locations seemingly less genetically divergent than in other similarly widespread taxa (Fig.
Male paracymbium with three medial processes that lie between the ventral and dorsal processes, including ventromedial, distomedial, and dorsomedial processes (Fig.
In the northern Newberry Creek population the male distomedial process is reduced (but present as low spikes), and the base of the dorsal paracymbial processes is wider then narrows to a forked tip (Fig.
Previously known only from fissure caves, including those summarized by
In comparison to its sister species
Extensive population-level variation is seen in the male palps across relatively short geographic distances in this species. This includes variation in the shape of the shoe-shaped tegular apophysis and the sclerotized extension, the presence and shape of the paracymbial ventromedial and distomedial processes, and the shape of the dorsal paracymbial process (Fig.
Variation exists in the shape of the lateral epigynal pockets (ventral view), but the overall vulval pocket morphology, spermathecal shape, and parallel epigynal plates is fairly conserved across populations (Fig.
Populations have been collected from the larger Bald Mountains area along the North Carolina / Tennessee border, southwest of Erwin, Tennessee (Fig.
This species is named to recognize and honor Dr. Alan Templeton, Charles Rebstock Professor Emeritus of Biology, Washington University. A brilliant evolutionary, speciation, and conservation biologist, with a deep love for all biodiversity. PhD dissertation advisor of MH, honored here for his inspiration and support during the first author’s formative years as an evolutionary biologist.
Two strongly supported geographic subclades are recovered with mitochondrial data (Fig.
Male palps differ in many ways from other members of the species group (including closest relatives), with a forked base of the tegulum, a narrow, curved tegular apophysis, a beak-like basal process of the median apophysis, and a translucent dorsal paracymbial process with a relatively wide base (Fig.
Male variation was observed in the shape of the median apophysis lateral process, the paracymbial ventral process, and the proximal fork of the tegulum (Fig.
Previously known only from the type location (Mt. Mitchell), corresponding to the highest uplands east of the Mississippi River in North America, above 2000 meters in elevation. Our new records indicate that this species is more widespread in the Black Mountains (both to the north and southeast), and we include here new records from west of the Blacks, in the Great Craggy Mountains (Fig.
Most collections have resulted in a relatively modest number of specimens taken. For example, at an apparently pristine boulderfield along the South Toe River (
Strongly supported as a clade by
See
This species shows surprisingly little genitalic variation despite a relatively large geographic distribution (e.g., compare ♂ Fig.
Previously known only from the type locality (Cedar Creek Cave), this species is a fairly widespread surface-dwelling species (Fig.
Surface collections are mostly from shaded boulderfields, with field notes suggesting spiders to be “fairly common” under rocks in void spaces. Montreat specimens were found in dark cracks and crevices of a man-made rock wall within 3 meters of a stream.
As discussed below, possibly synonymous with
Strongly supported as a clade by
Gertsch cites the type data for this “
The type female is clearly a representative of the
A possible region to search for
The male palp is like that of
The palp of the paratype male is similar to the holotype.
Adult females from the type locality vary in body size and in carapace and abdomen color (dark vs. light) but share a similar epigynum.
Known only from the type locality from along the Cane River, a tributary of the Nolichucky River. Adjacent collections have thus far only resulted in the collection of non-sister
Specimens from the type collection were found to be relatively common in void spaces beneath rocks in a small shaded boulderfield in roadside forest, at approximately 700 meters in elevation.
Named after the Cane River, a small north-flowing river found only in Yancey County, North Carolina.
Morphologically very similar to
This small species group is strongly supported as monophyletic on both concatenated and coalescent phylogenomic trees (Figs
Consistent with phylogenomic data, each species in this group is morphologically distinctive, easily separated by diagnostic features of both male and female genital morphology (Fig.
Comparative ♂♀ genitalia of
We do not identify diagnostic morphological features for the entire species group, as many aspects of both male and female morphology occur elsewhere in the combined lineages sister to the
Each of the species in this species group occupies a relatively small geographic distribution in three disjunct pockets of the far western Blue Ridge (Fig.
Distribution of the
Males are easily distinguished from other members of the species group by the unique shape of the median apophysis, the shape of the tegular apophysis and tegular keel, the shape of the dorsal paracymbial process, and possession of a thorn-shaped distomedial paracymbial process (Fig.
Males from different geographic locations show very minor variation in the width (at base) of the dorsal paracymbial process and depth of indentation between dorsal and distal processes (Fig.
Females from different geographic locations show very minor variation in the shape of the anterior internal sclerotized epigynal lobes (Fig.
Most populations are from the southwestern flanks of the Snowbird Mountains of western North Carolina (Fig.
At the type locality of Tipton Creek,
Named after Dr. Jason Bond, Professor and Schlinger Chair of Insect Systematics at the University of California Davis. Jason was born in the southern Appalachians, schooled in the mountains of western North Carolina, and perhaps sometimes paddled in the Snowbird Mountains. Jason has been a longtime close friend and arachnological colleague of MH and is for him forever a source of scientific (and life) inspiration.
The immature specimens from Tipton Creek are here attributed to
The diagnosis of
Minor variation is observed in the height and width of the paired epigynal plates across geographic locations (Fig.
This troglomorphic species is only known from caves in karst windows along the northwestern edge of Great Smoky Mountains National Park (Cades Cove, Tuckaleechee Cove; Fig.
Several male features distinguish
Males from different locations varied slightly in the shape of the basal fork of the tegular apophysis (Fig.
Epigynal structure fairly uniform across collecting locations (Fig.
Most populations are from the Chunky Gal, Tusquitee, and Valley River Mountains of western North Carolina (Fig.
At Fires Creek (
Collection records suggest that this species is less common in the Chunky Gal Mountains than in the more westerly Tusquitee and Valley River Mountains.
This species is named to recognize and honor Michael Lowder, faculty member at Stanly Community College, native North Carolinian, fan of western North Carolina, and collector of many Appalachian
The extent of mitochondrial divergence observed in this taxon over a small geographic region (including only Chunky Gal, Tusquitee, and Valley River Mountains) is notable (Fig.
This group includes the sister species
The unique morphology of each species is discussed below. We do not attempt to identify diagnostic morphological features for this small species group.
The diagnosis of
The shape of the basal tegular fork varies notably across cave locations. One male from Salt River Cave (
Known from possibly hundreds of caves in northwest Alabama and south-central Tennessee (Fig.
Based on consideration of morphology
Closely related to
The Hugden Branch Cave female specimen, representing the second known location for this species, is troglomorphic with an epigynum that closely matches females from the type locality.
This troglomorphic species is known from two nearby caves from a single mountain in southeastern Tennessee, near Chattanooga (Fig.
This small species group is strongly supported as monophyletic on both concatenated and coalescent phylogenomic trees (Figs
The unique morphology of each species is discussed below; we otherwise do not attempt to identify diagnostic morphological features for this small species group.
This group includes the geographically widespread
Distribution of
Male palp with a distinctive elongate conductor, with a tip that lacks the strong distal fold found in other Appalachian taxa. Strongly concave median apophysis with medial point, tegular apophysis with a shallow fork, basal branch just a small lobe (Fig.
The paradistal dorsal process of the paracymbium varies in shape from very low and inconspicuous (Fig.
Minor variation was observed in the shape of the epigynal median septum (sometimes with a median bulge, then narrowing distally, viewed ventrally, Fig.
This species has the largest known geographic distribution of any Appalachian
The southern Pitchfork Cave population is highly disjunct from all other more northerly records; this is possibly an artifact of insufficient collecting effort on the eastern edge of the Cumberland Plateau in east-central Tennessee (Fig.
This species is known from both caves (both deeper and twilight situations) and dark, relatively moist near-surface habitats (mostly void spaces in rock piles). As noted above,
This species is not recovered as monophyletic on mitochondrial gene trees but is instead fragmented into three separate clades (Fig.
Nearly eyeless, long-legged taxon. Male palp most similar to that of
Known only from a handful of limestone caves from three adjacent counties in northwest Georgia (Fig.
Morphological diagnosis as in
This troglomorphic taxon is currently known from only two caves in northwestern Georgia (Fig.
Phylogenomic structure indicates three subclades within this larger group, including a distinctive
The mitochondrial data do not support the overall
Male genital morphology suggests common ancestry for this complex of eight species (Fig.
Comparative ♂ morphology of
Species in this group are distributed in the montane southern Blue Ridge west of the Asheville Basin, except for the geographically disjunct
The diagnosis of
In males from non-type locations the ventromedial process is more confluent with the ventral process (less displaced medially) and more elongate.
Different populations exhibit very little epigynal variation despite a large and fragmented geographic distribution. Some adult females from Holly Flats campground are approximately one-half the size of other adult females.
Previously known only from the type locality at Joyce Kilmer Memorial Forest, which now represents one of the easternmost known records for the species. The species distribution almost forms a circle in the montane uplands that surround the Ducktown lowlands (lacking the eastern edge), with disjunct Cohutta, Sosebee Cove, and Blankenship Cave populations (Fig.
As an example of natural history we include here field notes for Cohutta Wilderness (
As discussed above
Monophyletic on mitochondrial and nuclear trees, with high gene and site CF values for the latter. Phylogenomic evidence strongly supports
Sister to other members of a phylogenomic subclade including
Males from four non-type localities match topotypic males very closely (Fig.
Females from different locations share a very similar epigynal morphology (Fig.
Known only from a very small area in the upper Chattooga River and upper Whitewater River drainages (Fig.
This species is named to recognize and honor Bob Dellinger, a special naturalist from western North Carolina. Bob’s knowledge of the flora and fauna of southern Appalachia is remarkable, and he personally collected or helped to collect (with first author MH) many
Similar to regional congener
This species is known only from the type locality south of the Tennessee River in north-central Alabama (Fig.
Collections in 1992 revealed a very large spider population in Cave Spring Cave, perhaps up to 1,000 individuals. This cave is home to a protected bat colony and located in a US National Wildlife Refuge. The extraordinary size of the
Part of a near phylogenomic trichotomy with
Most similar to close phylogenomic kin
Males are only known from the type locality and all match the holotype male, except for
Females from different locations share a very similar epigynal morphology (Fig.
Known only from three parallel north-flowing drainages in the Great Smoky Mountains National Park, including the Middle Prong of the Little Pigeon River, and more easterly draining Indian Camp and Cosby Creeks.
At the type locality in 1992 spiders were “very abundant in rock crevices, low to the ground, close to the river”.
Along the Maddron Bald and Mt. Cammerer trails we collected both
Named to honor Dr. Greta Binford. Friend, arachnologist, and Past President of the American Arachnological Society (AAS), here recognized for her inspirational spider research and her leadership in making the AAS a more diverse and welcoming society. We suspect that Dr. Binford would also greatly appreciate the beauty of the habitats that this spider calls home.
Part of a near phylogenomic trichotomy with
This species was called “N novsp2” (from site 48) in
This species is included in a phylogenomic subclade with
Other than the holotype male only two other males are known, and these closely match the holotype.
Females from adjacent locations share a very similar epigynal morphology (Fig.
Known from three closely adjacent locations from near the headwaters of the West Prong of the Little Pigeon River, Great Smoky Mountains National Park, on the southwest slopes of Mt. Leconte (Fig.
1994 collections from near the Chimney Picnic Area resulting in collections of a male and eight females were from a “large talus breakdown in a south-facing cove” in rich hardwood forest.
Named to honor Wilma Dykeman (1920–2006), a writer, speaker, teacher, historian, and environmentalist who spent most of her life in western North Carolina and eastern Tennessee. Mrs. Dykeman was devoted to social justice and environmental integrity, discussing Appalachian water pollution in her classic 1955 book ‘The French Broad’, and sharing a social justice award in 1957 for her co-authored book ‘Neither Black Nor White’.
Part of a near phylogenomic trichotomy with
This species was called “
Compared to other members of the challenging
northeastern
Males and females from both sides of the Little Tennessee River barrier (see below) share very similar genitalic morphologies (Figs
From montane habitats in southern North Carolina and northern Georgia. Populations are found both east (Cowee Mountains, including the type locality) and west (Nantahala Mountains) of the Little Tennessee River, a known dispersal barrier in other arachnid taxa (e.g.,
As an example of natural history, 1992 collections near Standing Indian Campground were made in a northwest-facing rocky ravine, where many specimens were collected “in dark ravine, wet, deep litter, rocks,
The species is obviously morphologically very similar to a disjunct
We have made extensive collections of other
See Diagnosis of
Females of
Males and females from different populations share very similar genitalic morphologies (Fig.
With a distribution similar to
As an example of natural history, specimens from Blowing Cave were collected from a cave entrance, while those from Little River were collected from beneath rockpiles directly adjacent to a stream.
One possibility is that the latter clade (Little River, Blowing Cave) is not
Male palps of
southwestern
We here discuss and distinguish
In the northeast we examined males from seven locations in addition to the type locality, noting minimal palpal variation (Fig.
northeastern
In the southwest we examined males from eighteen separate locations. All southwestern males approximated character conditions seen in northeastern males for all but one character. Males from eight locations possessed a paracymbium with the paradistal process lacking (and distomedial process moving towards the edge; Fig.
Females from southwestern populations vary slightly (Fig.
southwestern
This relatively wide-ranging montane species occurs from the northern side of the Great Smoky Mountains National Park, southwestward across the Little Tennessee River to the Yellow Creek, Cheoah, Snowbird, Nantahala, Valley River, and Tusquitee Mountains (Fig.
We hypothesize that the geographic gap north and northeast of Fontana Lake in the Great Smoky Mountains is an artifact of poor sampling, as this region is mostly roadless (Fig.
As an example of natural history, at Ball Road (
See comments above regarding the unlikely
Nuclear phylogenomic data is mostly consistent with this single species hypothesis, except for the southern disjunct Rock Creek Road population, further discussed below. Only one “
The mitochondrial evidence is similarly challenging to interpret in this complex, as mitochondrial data do not support the larger
The southern disjunct Rock Creek Road sample (Fig.
Many people helped to collect specimens, including Keith Crandall, Alan Cressler, Shahan Derkarabetian, Jonathan Mayes, Maureen McCormack, Steve Perlaky, Patti Perlaky and Dustin Wood. Collecting trips with Bob Dellinger, Steve O’Kane, Chris Phillips, Michael Lowder, Pierre Paquin, and the Appalachian crews of 2005–2007 (Dalton Hedin, Lars Hedin, Robin Keith, Jim Starrett, Steven Thomas) were particularly productive and memorable, and we owe all these people a huge thanks. Steve Perlaky from Chattanooga TN provided special assistance on several occasions, and his generosity is truly appreciated. Dr. Fred Coyle contributed many specimens based on his survey work in the Great Smoky Mountains National Park, and excellent hospitality. Special thanks are also owed to Dr. Kirk Zigler and Dr. Matt Niemiller who (along with their team members) have made many important recent collections, some by special request. We would also like to thank Wil Orndorff, Tom Malabad, and Katarina Kosič Ficco of the Virginia DCR Natural Heritage Program for their collections of specimens from Virginia. Many thanks to all landowners who graciously allowed spider collections on their property. Collections of Hedin, Dellinger, and Coyle from Great Smoky Mountains National Park were made with appropriate permits. Grants from the National Science Foundation (DDIG #9213184 to Alan Templeton and MH; DEB 1937725 to MH), Highlands Biological Station (MH), National Speleological Society (MH), the U.S. Fish and Wildlife Service (MH and Bob Dellinger), and the USDA (MH) funded this research. The late Dr. Norman Platnick (
occurrences (excel file)
occurrences (excel file)
Specimens used in molecular work
DNA records (excel file)
Input alignments, analysis log files, and output tree files
zip. archive