A revision of the minor species group in the millipede genus Nannaria Chamberlin, 1918 (Diplopoda, Polydesmida, Xystodesmidae)

Jackson C. Means; Derek A. Hennen; Paul E. Marek

doi:10.3897/zookeys.1030.62544

Research Article

A revision of the minor species group in the millipede genus Nannaria Chamberlin, 1918 (Diplopoda, Polydesmida, Xystodesmidae)

Jackson C. Means^‡, Derek A. Hennen^‡, Paul E. Marek^‡

‡ Virginia Tech, Blacksburg, United States of America

Corresponding author: Jackson C. Means ( mjacks4@vt.edu )

Academic editor: Dragan Antić

This is an open access article distributed under the terms of the CC0 Public Domain Dedication.

Citation: Means JC, Hennen DA, Marek PE (2021) A revision of the minor species group in the millipede genus Nannaria Chamberlin, 1918 (Diplopoda, Polydesmida, Xystodesmidae). ZooKeys 1030: 1-180. https://doi.org/10.3897/zookeys.1030.62544

ZooBank: urn:lsid:zoobank.org:pub:48997D3B-9B4E-4348-8F30-0C8E8672569A

Abstract

Millipedes in the family Xystodesmidae (Polydesmida) are often referred to as “colorful, flat-backed millipedes” for their bright aposematic coloration and tendency to form Müllerian mimicry rings in the Appalachian region. However, there are many species of Xystodesmidae that do not display colorful warning patterns, and instead have more cryptic appearances. Perhaps for this reason, groups such as the genus Nannaria have remained understudied, despite containing a large number of undescribed species. Before his death in 2012, R. L. Hoffman worked on a revision of the genus Nannaria, and synthesized material and drawings since 1949. Here the work is continued, inferring a molecular phylogeny of the Nannariini (Nannaria + Oenomaea pulchella), and revealing two clades within the genus. One clade is named the minor species group, and the second is the wilsoni species group. This revision, using a molecular phylogenetic framework, is the basis for descriptions of 35 new species in the minor species group. A multi-gene molecular phylogeny is used to make taxonomic changes in the taxon. Eleven putative species of Nannaria are also illustrated and discussed. Additionally, detailed collection, natural history and habitat notes, distribution maps, and a key to species of the Nannaria minor species group are provided. These items are synthesized as a basis for a revision of the genus, which hopefully will aid conservation and evolutionary investigations of this cryptic and understudied group.

Keywords

Appalachia, gonopod, micro-range endemic, short-range endemic, spatulate, twisted-claw

Introduction

The genus Nannaria Chamberlin, 1918 (Polydesmida, Xystodesmidae) inhabits the eastern United States, from western Arkansas and Missouri to just outside of Rochester, New York in the north, to the coast of Virginia and to central Mississippi in the south (Fig. 1). Despite this extensive range, the group as a whole is poorly known, and only two studies in the last 40 years have specifically addressed taxonomic changes to Nannaria (Hennen and Shelley 2015; Shelley and Smith 2018). Furthermore, genetic techniques are still rarely applied in modern diplopod systematics, and Nannaria has been included only twice in a molecular phylogeny (Marek and Bond 2006; Means et al. 2021). While these works shed light on the placement of Nannaria within the Xystodesmidae, their analyses relied on limited taxon sampling of the tribe. While the affinities of the Nannariini [Nannaria + Oenomaea pulchella (Bollman, 1889)] to the other members of the family have been shown, their taxon sampling scheme omitted most nannariine species. Here, we use six genes (16S, 28S, EF1a, rpb1, COI, and fbox) to construct a molecular phylogeny of the Nannariini, and include 139 Nannaria specimens, O. pulchella and 12 taxa in the Rhysodesmini, to resolve the phylogenetic relationships between the genera of Nannariini. Additionally, we describe 35 new species, and demonstrate that Nannaria contains two clades, one of which, the wilsoni species group, is to be revised in a subsequent publication.

Figure 1.

The geographic distribution of Nannaria Chamberlin, 1918. Red: minor species group; Purple: wilsoni species group.

History of the Nannariini and diagnostic characters

The tribe Nannariini has received little attention since being proposed in 1964 by R. L. Hoffman, and as such does not have the same nomenclatural peregrinations of some animal taxa (Hoffman 1964). The species within the Nannariini have been placed in five different genera over the years (Mimuloria Chamberlin, 1928, Fontaria Gray, 1832, Castanaria Causey, 1950, Nannaria, and Polydesmus Latreille, 1802), with the most recent change by Means et al. (2021) synonymizing Mimuloria with Nannaria for the current count of two genera within the Nannariini. The oldest of these two, Nannaria, was first proposed by Chamberlin (1918), and he included within it the previously described Fontaria tennesseensis Bollman, 1888, as well as three new species, Nannaria media Chamberlin, 1918, N. infesta Chamberlin, 1918 and N. minor Chamberlin, 1918. Two of these species, N. infesta and N. media, have recently been shown to belong to separate genera (Means et al. 2021), now Howellaria infesta (Chamberlin, 1918) and Boraria stricta (Brölemann, 1896), respectively. Chamberlin’s (1918) description of Nannaria was brief, and while he stated that he would define the genus in greater detail later, Chamberlin did not revisit the task. Additionally, neither Chamberlin (1918) nor Bollman (1888) provided illustrations of the gonopods, the external male reproductive structures traditionally used for specific and generic differentiation in millipede taxonomy, for the above four species; therefore Nannaria had remained a poorly defined genus for 31 years until Hoffman (1949) finally provided a brief diagnosis of the genus. Williams and Hefner (1928) later stated that “it is not deemed necessary to make these more technical divisions” in reference to Chamberlin’s split of Fontaria into three genera, Mimuloria, Nannaria, and Apheloria Chamberlin, 1921 (Chamberlin 1918, 1921, 1928); however, they themselves did not propose an alternative arrangement. Nor did they formalize a synonymy. Hoffman’s (1949) diagnosis merely mentioned that Nannaria have laterally positioned repugnatorial pores, gonopods with paired processes, and are “olive to black with keels [paranota, or lateral extensions of the body ring] pink”. [Note that color may confound diagnosing species in the Xystodesmidae (Marek and Bond 2006), and it is now known that species with distinctly different color patterns may be the same species or population of millipede.] Perhaps due to their presence in Oenomaea pulchella and absence in H. infesta and B. stricta, Hoffman (1964) did not list the distinct spatulate pregonopodal claws of Nannaria as a diagnostic feature of the tribe (see Fig. 2). Hoffman (1964) also omitted from his diagnosis the subcoxal sternal spines that are characteristic of the tribe, though again this may be due to the lack of sternal modifications in H. infesta and B. stricta.

Figure 2.

Scanning electron microscope image of a spatulate, twisted pregonopodal claw of a male Nannaria ericacea Hoffman, 1949.

Fifteen years later, spurred by the discovery of an unusual male xystodesmid from Georgia which conformed somatically to the female type specimens of Fontaria pulchella Bollman, 1889 collected in Tennessee, Hoffman (1964) proposed the tribe Nannariini to include Nannaria and the new genus Oenomaea Hoffman, 1964 (Fig. 3). Hoffman (1964) also synonymized the genus Mimuloria under Nannaria, a change which would later be reversed by Hennen and Shelley (2015), and again synonymized by Means et al. (2021). Hoffman (1964) provided a diagnosis of the tribe, which included characters such as the spatulate pregonopodal claws, the subcoxal sternal spines and the conical coxal projections on the fourth leg pair. The three characters which he provided to differentiate the genera Oenomaea and Nannaria are the strongly twisted pregonopodal claws of Nannaria and the acutely projecting paranotal corners and the flattened, plate-like solenomerite (tip of the acropodite) of Oenomaea (Hoffman 1964).

Figure 3.

Examples of the coloration of Oenomaea pulchella Hoffman, 1964.

Nearly a half-century later, Hennen and Shelley (2015) revived Mimuloria as a separate genus, synonymizing Nannaria depalmai (Causey 1950a) under Mimuloria castanea (McNeill, 1887) and describing two new species including one subspecies. Hennen and Shelley (2015) suggested that the Nannaria species with complex gonopods be placed within Oenomaea or in a new genus, due to their gonopods having solenomeres, the structure which carries the seminal groove after a bifurcation near the tip of the acropodite; however, they did not formally make this change. The changes that Hennen and Shelley’s (2015) publication made to the Nannariini were based entirely on morphological characters, primarily the terminal acropodite ornamentation found in some Nannaria (see Figs 22, 36, and 37). This was the major character that both Mimuloria and Castanaria were based upon when they were described by Chamberlin (1928) and Causey (1950a), and Hoffman (1964: 33) stated that “I find this character to be a mutable one, and more suitable for the distinction of a species-group than a separate genus.” Mimuloria was thereafter synonymized under Nannaria by Means et al. (2021), after their six-gene (16S, 12S, tRNAval, COI, EF1a, and 28S) molecular phylogeny found that species of Mimuloria were in multiple locations within Nannaria.

Shelley and Smith (2018) moved the family Eurymerodesmidae into the Xystodesmidae, based on a hypothesized sister group relationship with the Nannariini. Due to the priority of the Eurymerodesmini, the Nannariini were reduced to a subtribe, the Nannariina. This change was made in the absence of molecular evidence, and based solely on the presence of stout setae along the inner margin of the gonopods. Based on molecular phylogenetics, Means et al. (2021) found that the Eurymerodesmini are members of the Xystodesmidae, and with the tribe Euryurini, are sister to the Nannariini; the tribe Nannariini was therefore revived as a separate tribe distinct from the Eurymerodesmini and Euryurini.

A considerable breadth of gonopodal variation exists amongst populations of Nannaria and has been a great challenge to the solely morphology-based taxonomy of the group. Hoffman (1964, 1999) and Hennen and Shelley (2015) have remarked that there exists a wide variety of gonopodal forms in Nannaria, which either indicates a highly variable massive genus or multiple genera. The most obvious divide is between those with simple, stick-like gonopods (Fig. 4A–C, the minor species group after the generotype N. minor) and those with complex, twisting, and often adorned gonopods (Fig. 4D–F, the wilsoni species group after N. wilsoni Hoffman, 1949, a species found around Blacksburg, Virginia). While Hoffman (1964) commented on the variety of gonopodal forms in the Nannaria on multiple occasions, even going so far as to suggest that there may exist up to 200 species within the genus, he never pointed out this divide between simple and complex gonopods in his publications or his personal notes. Here we present a six-gene molecular phylogeny of the Nannariini which reveals multiple clades within the minor species group, and 35 new species.

Figure 4.

Examples of the minor species group (A–C) and wilsoni species group (D–F) gonopods, anterior view, setae removed for clarity A Nannaria minor Chamberlin, 1918 B N. terricola Williams & Hefner, 1928 C N. scholastica sp. nov. D N. austricola Hoffman, 1950 E N. shenandoa Hoffman, 1949 F N. ericacea Hoffman, 1949.

Materials and methods

Specimen collection and preservation

Collection sites were selected based on a combination of factors, including: availability of suitable habitat, location of type localities, information from R. L. Hoffman’s notes on Nannaria, and natural history collections from the Virginia Museum of Natural History and the North Carolina State Museum—repositories with large holdings of nannariine specimens. Collection methods followed that of Means et al. (2015). Briefly, adult and juvenile Nannaria were collected by hand with the use of a millipede rake or gardening claw, often by removing leaf litter and digging 2–3 cm beneath the soil. For the 12 species of Rhysodesmini used in this study, collection methods were similar, though adults were often found under leaf litter, above the soil.

Adult and juvenile millipedes were brought back to the lab for identification, photography, DNA and RNA collection, preservation, and, in the case of juveniles, housed in terraria until maturation. Live millipedes were photographed with a Canon EOS 6D digital camera with a 50 mm macro lens to record color and habitus morphology. Legs from rings 8–18 were removed using forceps from the left side of adults and placed in either RNAlater (Qiagen, Hilden, Germany) for the preservation of DNA and RNA, or 100% EtOH for the preservation of DNA. Preserved legs were stored at -80 °C for archival storage. Specimens sans legs were given a unique specimen code, stored in 70% isopropanol and deposited in the Virginia Tech Insect Collection (VTEC; https://collection.ento.vt.edu). Type material deposition location is noted in each species description under Materials Examined, and includes the VTEC, the Field Museum of Natural History (FMNH), and the Virginia Museum of Natural History (VMNH). Institutions which provided specimen loans are as follows: Florida State Collection of Arthropods (FSCA), North Carolina Museum of Natural Sciences (NCSM), Museum fur Naturkunde, Berlin (MFN), Smithsonian National Museum of Natural History (NMNH), and the VMNH. Institution abbreviations mentioned in the text include The Academy of Natural Sciences of Philadelphia (ANSP) and the United States National Museum (USNM; now NMNH, see above). The abbreviation “leg.” (legit = ‘has collected’) follows the name(s) of the collector(s) of the specimen.

Primer design

The transcriptomes of Nannaria hokie Means, Hennen & Marek, 2021 (minor species group) and Nannaria ericacea (wilsoni species group) sequenced by Means et al. (2021) were used for gene exploration and primer development as described by Means et al. (2021). Twenty-seven gene region candidates (Suppl. material 1) were blasted (Altschul et al. 1990) against both transcriptomes and eleven were chosen for primer design and testing based on the following criteria: presence in both Nannaria transcriptomes, gene region length > 400 base pairs, and > 3% gene region variation between transcriptomes.

DNA extraction and phylogenetics

DNA was extracted and purified using a Qiagen DNeasy tissue kit and stored at -20 °C. For Nannaria, we used 4–6 legs per specimen and eluted in 50 ul of Qiagen DNeasy Buffer AE due to their small size (average 25 mm in length); while for larger bodied Xystodesmidae, such as the outgroup Pachydesmus Cook, 1895, we used three legs and 100 µl of Buffer AE. For legs stored in 100% EtOH, these were air-dried at room temperature (22 °C) prior to DNA extraction with the same protocol above. Six gene fragments were amplified for each specimen: cytochrome c oxidase subunit I gene (COI), elongation factor-1 alpha gene (EF1a), 28S ribosomal RNA gene (28S), large ribosomal RNA gene (16S), F-box protein (fbox), and DNA-directed RNA polymerase II subunit RPB1 (RPB1). Amplifications were cleaned, quantified, and then sequenced using an Applied Biosystems 3730 DNA Analyzer.

We used the programs phred and phrap within the Mesquite (Version 3.5) module Chromaseq (Version 1.31) for base-calling, generation of sequence contigs and sequence trimming (Maddison and Maddison 2010; Ewing et al. 1998). Individual genes were aligned with five iterations each in PRANK (Version 140603) and with the default HKY model with empirical base frequencies, kappa=2, the -F option and iterative guide trees (Löytynoja and Goldman 2005). Aligned sequences were then concatenated in Mesquite, partitioned by gene, codon site (for protein-coding genes), and intron/exon boundaries (for EF1a and RPB1). The concatenated matrix was then exported to PartitionFinder 2 (Version 2.1.1) as .phy and .cfg files for determination of best-fit models of nucleotide site substitution for each partition (Table 1). Partition groupings were determined using the ‘greedy’ scheme and a Bayesian Information Criterion (BIC) model selection method (Lanfear et al. 2012). For the phylogenetic analysis, the concatenated matrix was exported from Mesquite for MrBayes (Ronquist et al. 2005), and the best-fit partition block from PartionFinder 2 was included in the nexus file (Lanfear et al. 2012). In MrBayes (Version 3.2.6), we ran two simultaneous pairs of hot and cold MCMC chains, with a 25% burn-in, for 210 million generations. Sampling occurred every 100 generations until the standard deviation of split frequencies (SDSF) reached < 0.01, indicating that the chains had converged on a set of consensus topologies (Ronquist et al. 2005). The MrBayes commands “sumt” with “contype=allcompat” were used to summarize the posterior distribution of trees into a consensus tree while displaying frequencies of all bipartitions. Individual gene trees were estimated in MrBayes to assess separate gene histories. Nucleotide base composition and frequency, as well as sequence heterogeneity, were assessed in IQ-TREE 2 (Version 2.0.4, Minh et al. 2020), excluding non-Nannariini taxa. Sequence heterogeneity was measured using χ² tests of stationarity, with the null hypothesis of homogeneity (Table 2).

Morphological character scoring, illustrations, and distribution mapping

For each species, gonopods were dissected under a Leica M125 stereomicroscope (Leica Microsystems, Wetzlar, Germany), and photographed using a Canon 6D camera with a 65 mm MP-E macro lens mounted on a Passport II Portable Digital Imaging System (Visionary Digital, Charlottesville, VA). Gonopods were photographed at three angles (anterior, posterior, and medial) and photographs were taken at 4× zoom, every 0.08 mm, for an average of 8–15 images per angle. These images were then focus stacked to create a composite photograph using the program Helicon Focus (Helicon, Kharkiv, Ukraine). Gonopod photographs were then traced in Adobe Illustrator CC 2018–2021 using the pen tool (Adobe, San Jose, CA). Specimens were then scored for gonopodal, cyphopodal and somatic characters in a 48-character matrix adapted from Marek and Bond (2006) for Nannaria (Suppl. material 6). All measurements were recorded in millimeters and the following six areas were measured according to Marek (2010): body length (BL), collum width (CW), intergenal width (IW), interantennal socket width (ISW), body ring 11 width (B11W), and body ring 11 height (B11H). Latitude and longitude coordinates were obtained for literature records by georeferencing collection data in Google Earth (Google LLC, Mountain View, CA). Distribution maps were generated using SimpleMappr (Shorthouse 2010) and edited in Adobe Illustrator CC 2018–2021. Distributions represented by a single collection locality are denoted as “N/A” below.

Species delimitation

For the delimitation of species, we used a combination of morphological and molecular criteria as implemented in Marek (2010) and Means et al. (2021). Briefly, a representative specimen from a sampled population needed to display overt morphological dissimilarity from previously known species, and/or be placed in the concatenated molecular phylogeny in an un-hypothesized location, to be considered an undescribed species. There exists a paucity of somatic features which vary between Nannaria species and populations, however gonopodal variation between species can be pronounced. Differences in gonopod morphology have dictated the establishment of a majority of species in the Diplopoda, and we are confident that a combination of gonopodal variation and molecular evidence is reliable criteria for species delimitation. We accepted paraphyletic species where multiple populations of a single species were paraphyletic with respect to another species and possessed overt gonopodal differences. Specimens which displayed gonopod morphology that was sufficiently different from known species of Nannaria, but for which we did not have genetic material, were designated as incertae sedis. While we include illustrations of incertae sedis specimens to aid in future taxonomic work on Nannaria, we refrain from elevating these indeterminate specimens to species. For this revision we use the higher classification for the Xystodesmidae and Nannaria as outlined in Means et al. (2021).

Figure 5.

Nannaria cingulata sp. nov. telopodite, posterior view. Abbreviations: ap = acropodite; as = acropodite swelling; at = acropodite tip; bz = basal zone; cg = cingulum; cx = coxa; dz = distal zone; gp = gap between pp & ac; lb = basal zone lateral bulge; pp = prefemoral process; ps = prefemoral spine; sc = seminal canal; tlf = tip lateral flange; tmf = tip medial flange.

Morphological terminology

Morphological terms used to describe gonopods in Nannaria are as follows:

The coxa, defined as the basal segment of the gonopod [gonopod = a leg which has been modified into a structure for sperm transfer (Fig. 5, cx)].
The gonopodal telopodite, defined as the area distal to the coxa (Fig. 5).
The basal zone lateral bulge, defined as the lateral edge of the base of the gonopod which is occasionally pronounced in size (Fig. 5, lb)
The gonopod basal zone, defined as the area between the coxa and the acropodite (Fig. 5, bz).
The prefemoral spine, a new term, defined as the process arising at the base of the prefemoral process, often as a sharp spine, but occasionally reduced to a ridge, bulge, or completely absent (Fig. 5, ps).
The seminal canal, the linear trough that carries sperm during copulation, indicated in illustrations as a dashed line (Fig. 5, sc).
The prefemoral process, defined as the smaller of the two branches of the telopodite, and which does not carry the seminal canal (Fig. 5, pp).
The cingulum, defined as a fold or linear depression, found only in Nannaria cingulata sp. nov. (Fig. 5, cg).
The acropodite, which carries the prostatic groove (Fig. 5, ap), a distal subregion of the telopodite that does not include the prefemur.
The ‘gap’ between the prefemoral process and the acropodite (Fig. 5, gp).
The acropodite swelling, which is defined as present or absent, is a swelling of the region distal to the basal zone (Fig. 5, as).
The distal zone, defined as the area distal to the major inflection point of the gonopod (Fig. 5, dz).
The tip lateral flange, a flange which is occasionally present on the upper portion of the acropodite tip (Fig. 5, tlf).
The tip medial flange, a flange which is occasionally present on the underside of the acropodite tip (Fig. 5, tmf).
The acropodite tip, defined as the terminal portion of the acropodite (Fig. 5, at).

Results

Specimen collection

In total, 935 Nannaria specimens were collected between 2014 and 2018 throughout the eastern United States. Areas of suitable habitat included broad leaved forests and hemlock groves, typically near a body of water or riparian area, and Rhododendron coves (Fig. 6). Immature Nannaria appeared to display a preference for molting underneath Rhododendron, while adult Nannaria were readily found near the edges of Rhododendron groves, but rarely within the actual grove. Additionally, sloped hillsides near streams with dark, loamy soil and sparse root mats were often productive sites to encounter Nannaria specimens. A .kmz file of Nannaria minor species group distributions for use in programs such as Google Earth (Alphabet Inc., Mountain View, California) is provided as Suppl. material 8.

Figure 6.

Examples of Nannaria Chamberlin, 1918 habitat A broadleaved forest and Rhododendron cove interface B riparian corridor C steep mesic hillside.

Primer design

Of the 27 candidate gene regions from the Nannaria transcriptomes, eleven were selected for primer design, and ultimately three newly developed gene regions were then used in phylogenetic analyses, based on amplification success (> 90%) and sequencing success (> 90% contig rate): large subunit ribosomal RNA gene (16S), fbox domain (fbox), RNA polymerase II largest subunit (RPB1). These three gene regions were then combined with the three other gene regions standardly used in millipede molecular phylogenetics: COI, 28S, and EF1a (see below).

Table 1.

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Partitions used in MrBayes phylogenetic analysis of molecular data. PIC = Parsimony Informative Characters.

Partition	Gene region	Best-fit models	PIC
1	16S	GTR+I+G	368
2	CO1 (3^rd CP)	GTR+I+G	154
3	CO1 (1^st CP)	GTR+I+G	34
4	CO1 (2^nd CP)	GTR+G	5
5	fbox (1^st CP), 28S	GTR+I+G	121
6	fbox (3^rd CP), EF1a (3^rd CP)	GTR+G	63
7	EF1a (1^st & 2^nd CPs), RPB1 (1^st & 2^nd CPs)	K80 +I+G	40
8	EF1a (intron)	HKY+G	69
9	fbox (2^nd CP)	JC+I	10
10	RPB1 (intron & 3^rd CP)	HKY+I+G	128

Sequence alignment and phylogenetic inference

The concatenated matrix contained six genes, 152 taxa, and a total length of 5,354 bp, as follows: 16S (1–1016), COI (1017–1553), 28S (1554–3178), EF1a (3179–3879), fbox (3880–4298), and RPB1 (4299–5354; for a list of taxa and NCBI accession numbers for specific gene regions see Suppl. material 2). The six genes were divided in PartitionFinder into ten partitions (Table 1). Of the 5,354 characters, 3,787 were constant, 992 were parsimony-informative, and 575 characters were variable and parsimony-uninformative. Observed mean base pair composition for the 5,354 characters was A = 0.235, C = 0.197, G = 0.263, T = 0.303, and for each gene fragment as given in Table 2. For each gene region, nucleotide frequency was homogeneous across taxa (P < 0.05, df = 3), with no taxa failing the χ² test of stationarity (Table 2). Mean uncorrected percent difference of COI sequences between Nannaria species in the minor group was 9.8% (maximum: 14%, minimum: 0.0%, standard deviation: 1.8%). In the MrBayes analysis, likelihood values converged, as indicated by the average standard deviation of split frequencies reaching < 0.01, after 210 million generations, and one quarter of the generated trees were then discarded as burnin. A topology was generated from the six-gene concatenated matrix, with 94 of the 151 nodes having posterior probabilities ≥ 0.95. Due to the lack of possible gonopod-based identification, 33 female specimens were pruned from the minor species group tree post-analysis, and a tree with only minor species group specimens that are identifiable and diagnosable is presented in Fig. 7. To view the unpruned tree including outgroups, the wilsoni group and minor group female specimens, see Suppl. material 3.

Table 2.

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Nucleotide base frequencies and average gap/ambiguity (%) by gene.

Gene region	A	C	G	T	Av. % Gap/Ambiguity
16S	0.27	0.07	0.23	0.42	30.72
COI	0.19	0.14	0.25	0.42	0.74
EF1a	0.26	0.23	0.25	0.26	24.48
28S	0.15	0.29	0.36	0.20	7.96
fbox	0.23	0.27	0.26	0.23	0.01
Rpb1	0.31	0.18	0.23	0.29	6.94

Figure 7.

Molecular phylogeny of the minor species group. Asterisks indicate a posterior probability of > 0.95. Colored boxes and arrows indicate clades: black arrow, Nannaria scholastica sp. nov.; sky blue box, N. domestica clade; orange box, N. fowleri clade; light yellow box, N. paupertas clade; green box, N. mcelroyorum clade; burnt orange arrow, N. hardeni sp. nov.; brown box, N. serpens clade; turquoise box, N. castanea clade; lime green box, N. laminata clade; dark pink box, N. tasskelsoae clade; camo green box, N. tennesseensis clade; yellow box, N. ignis clade; blue box, N. ohionis clade; light pink box, N. blackmountainensis clade; forest green box, N. terricola clade; purple box, N. ambulatrix clade; red box, N. minor clade.

Taxonomic notes

The Nannariini was recovered as a monophyletic group (posterior probability, pp = 1), with Oenomaea pulchella sister to Nannaria (pp = 1; Suppl. material 3). Eurymerodesmidae was not recovered as sister to the Nannariini, however in light of recent genetic findings using a much greater sample of Eurymerodesmidae and Xystodesmidae, this result is likely due to low taxon sampling of other members of the subfamily here (Means et al. 2021). The genus Nannaria is split between two clades, the wilsoni and minor species groups (pp = 1), a relationship that is also recovered in five of the six individual gene trees (Suppl. material 4). The wilsoni species group contains many hypothesized undescribed species that will be described in the second part of the revision of Nannaria. The minor group species was made up of 18 divergent clades representing new species (Fig. 7), including the taxa Nannaria scholastica sp. nov. and Nannaria hardeni sp. nov., and a clade consisting entirely of indeterminate female specimens shown in Suppl. material 3 (Murphy clade, eggplant purple box). The minor group clades possess a variety of support values (average pp = 0.64; Figs 7, 113; Suppl. material 3). Three of the 45 species were paraphyletic (Nannaria ignis sp. nov., Nannaria fracta sp. nov., and Nannaria laminata Hoffman, 1949); however, we opted to recognize these taxa due to divergent gonopodal morphology and meeting our species delimitation criteria.

Color

Of 171 Nannaria specimens recorded for color, 148 (86.5%) had orange, pink, or red paranota with a dark brown or black background (as in Fig. 89B), while only 23 (13.5%) had white paranota (as in Fig. 89A). Twenty-three (15.5%) of the 148 orange/pink/red Nannaria also had metatergal stripes (as in Fig. 97).

Nannaria minor species group diagnoses

The following are diagnoses of 54 species based on our species delimitation criteria (see Materials and Methods). Diagnoses are provided as paragraphs that include a suite of characters that differentiate the species from the most morphologically similar minor group species and the sympatric or geographically closest known wilsoni group species. Descriptions are provided as a scored morphological matrix for rapid comparisons of species (Suppl. material 5 and 6). Distribution statuses are provided based on three categories, wide-range endemic (WRE): distribution area > 10,000 km²; short-range endemic (SRE): distribution area < 10,000 km² and > 1,000 km²; and micro-range endemic (MRE): distribution area < 1,000 km² (Harvey 2002; Harvey et al. 2011; Means and Marek 2017). Overall, Nannaria species in the minor group have restricted distributions, with 91% falling into the SRE or MRE categories. Species descriptions are grouped by clade, and clades are organized as they appear in the molecular phylogeny (Fig. 7).

Nannaria Taxonomy

Class Diplopoda de Blainville in Gervais, 1844

Infraclass Helminthomorpha Pocock, 1887

Order Polydesmida Leach, 1815

Family Xystodesmidae Cook, 1895

Subfamily Rhysodesminae Brolemann, 1916

Tribe Nannariini Hoffman, 1964

Genus Nannaria Chamberlin, 1918

Vernacular name: “The Twisted-Claw Millipedes”

Type species

Nannaria minor Chamberlin, 1918, by original designation.

Other taxa included

61 species.

Diagnosis

Members of the genus Nannaria are distinct from other genera of the subfamily Rhysodesminae, except the monotypic genus Oenomaea, by the twisted spatulate shape of the pregonopodal claws in males (Fig. 2) vs. bisinuately curved shape in other genera of Rhysodesminae. Nannaria vary from the contribal Oenomaea by the more rounded paranota in Nannaria (vs. more acute and hook-like in Oenomaea) and the unique gonopodal characters of Oenomaea, as discussed in Hoffman (1964).

Short description

Small-bodied (17.3–38.5 mm BL, 2.9–6.0 mm B11W) ‘flat-backed’ xystodesmid millipedes. Color: paranota red, orange, white, or (rarely) yellow—rarely connected by concolorus metatergal stripes. Background drab, ranging from pale tan to deep black. Male exoskeletal characters: sterna with paramedian lobes between 4^th pair of legs; caudal margins of 8–18 sterna modified into subcoxal spines. Pregonopodal tarsal claws spatulate [(Fig. 2)—note: the twisted spatulate shape of the pregonopodal claws in males is a unique autapomorphy and uniformly present in the tribe Nannariini (Oenomaea pulchella and Nannaria species)]. Postgonopodal tarsal claws bisinuate. Male gonopodal characters: gonopods variable, often crossing, when viewed in situ; co-planar with coxae, lacking torsion, generally simple in comparison to other Rhysodesminae—though some species display elaborate curvature and ornamentation. Gonopods never bulky as in some Apheloriini. Prefemoral process present. Prostatic groove originating at cannula, terminating at acropodite tip. Acropodite, when viewed ventrally, c- or r-shaped. Gonopodal aperture lacking modified sternum. Gonocoxae connected by membrane and muscle.

Distribution

Nannaria can be found in much of the eastern United States (Fig. 1): western Arkansas east to just outside Philadelphia (Pennslyvania), north to western New York, and south to central Mississippi. The southeastern edge of the distribution follows the Appalachian Mountains closely, though specimens from central North Carolina and Mississippi suggest that populations may eventually be found south of the Appalachians in states such as Texas and Florida. Likewise, scattered populations may exist in central and southern Illinois, as seen in neighboring Indiana and Missouri.

Etymology

Derived from the Greek word nannos, for dwarf, referring to the comparatively small size of this genus within the Xystodesmidae.

Key to the minor species group of Nannaria Chamberlin, 1918

1	Gonopods fairly straight, parallel for most of length, rarely crossing when viewed in situ (Fig. 8A)	2
–	Gonopods curved, often crossing when viewed in situ (Fig. 8B)	10
2 (1)	Prefemoral basal spine fused with prefemoral process for entire length (Fig. 86B). Ohio	*N. terricola* (Figs 86, 87)
–	Prefemoral basal spine separate from prefemoral process for part of length (Fig. 88A)	3
3 (2)	Prefemoral basal spine large, pronounced (Fig. 57A)	4
–	Prefemoral basal spine small, reduced (Fig. 88C)	5
4 (3)	Prefemoral basal spine pointed, sharp (Fig. 80C). Acropodite with small, shelf-like medial flange just before tip (Fig. 80C, red arrow). Telopodite basal zone with pronounced medial swelling (Fig. 80A, red circle). Virginia – Kentucky border	N. fracta sp. nov. (Figs 80, 81)
–	Prefemoral basal spine pointed, blunt (Fig. 57A). Distal zone much reduced, rounded, with small, lobed lateral flange (Fig. 57A, red triangle). Prefemur with large, laminate prefemoral process, curving medially, crossing acropodite and expanding before tip. Knox Co., Tennessee	N. monsdomia sp. nov. (Figs 57, 58)
5 (4)	Prefemoral proccess arising from prefemoral spine, dorsomedially (Fig. 88A). Distal zone short, directed dorsomedially, with large, triangular lateral flange (Fig. 88A, red arrow). Telopodite basal zone large, subequal to length of acropodite. Southwestern Virginia	N. ambulatrix sp. nov. (Figs 88, 89)
–	Prefemoral process arising from prefemur (Fig. 54A)	6
6 (5)	Prefemoral process sinuous, serpentine (Fig. 69A, B). Distal zone short, directed dorsomedially, with large, triangular lateral flange (Fig. 69A, red arrow). Virginia – Tennessee border	N. suprema sp. nov. (Figs 69, 70)
–	Prefemoral process variable, though never sinuous, serpentine	7
7 (6)	Prefemoral process laminate (Fig. 54A). Distal zone much reduced, with rounded, simple tip. Acropodite with medial swelling. Acropodite appearing as straight line in medial view. Knox Co., Tennessee	*N. equalis* (Fig. 54)
–	Prefemoral process acicular (Fig. 100A)	8
8 (7)	Gonopod basal zone with slight lateral bulge (Fig. 100A). Acropodite tip with prominent lobed lateral flange (Fig. 100A, red arrow). Acropodite swollen medially before apex (Fig. 100A, red triangle). Greene Co., Tennessee	N. hippopotamus sp. nov. (Figs 100, 101)
–	Gonopod basal zone without lateral bulge (Fig. 85A)	9
9 (8)	Gonopod basal zone height, when viewed anteriorly, subequal to half length of acropodite. Acropodite straight. Distal zone short, rectangular in composition when viewed anteriorly, bent medially at 90° angle with slight cephalically-directed upturn at terminal edge (Fig. 82A, red arrow). Southwestern Virginia, southern West Virginia	N. solenas sp. nov. (Figs 82–84)
–	Gonopod basal zone height, when viewed anteriorly, greater than half length of acropodite. Acropodite straight, curving medially at nearly 90° angle at apex. Acropodite tip with small, rounded lateral flange (Fig. 85C, red arrow). Prefemur with straight, acicular prefemoral process and reduced prefemoral spine (Fig. 85C, red triangle). Wise Co. Virginia	N. spruilli sp. nov. (Fig. 85)
10 (1)	Prefemoral basal spine absent (Fig. 71B)	11
–	Prefemoral basal spine present (Fig. 90B)	16
11 (10)	Prefemoral basal spine fused with prefemoral process for entire length (Fig. 38A). Acropodite long, curving medially. Distal zone with medial and lateral flanges at 90° angle to solenomere (Fig. 38B, red arrow). Acropodite with small medial flange near apex (Fig. 38A, red triangle). Missouri	*N. missouriensis* (Figs 38, 39)
–	Prefemoral basal spine separate from prefemoral process for some part of length (Fig. 88A)	12
12 (11)	Prefemoral process thin, acicular (Fig. 110A). Distal zone short, simple, curving smoothly posteriorly, with a crochet hook-like appearance (Fig. 110A, red triangle). Prefemoral spine lacking. Pennsylvania	*N. oblonga* (Fig. 110)
–	Prefemoral process variable, though never thin, acicular	13
13 (12)	Prefemoral process laminate (Fig. 54A)	14
–	Prefemoral process simple, curving (Fig. 71A). Acropodite tip with small, triangular lateral flange (Fig. 71A, red arrow). Telopodite basal zone with slight lateral bulge (Fig. 71A, red triangle). Gap between prefemoral process and acropodite greater than width of acropodite basal zone. Pulaski Co. Kentucky	N. alpina sp. nov. (Figs 71, 72)
14 (13)	Prefemoral process curving medially, when viewed anteriorly (Fig. 20A). Distal zone curving dorsally, with tip directed caudally (Fig. 20B). Acropodite tip with lateral flange (Fig. 20A, red arrow). Acropodite with slight twist and swelling at midpoint (Fig. 20C, red triangle). Mercer Co., West Virginia	N. castra sp. nov. (Figs 20, 21)
–	Prefemoral process variable, though never curving medially, when viewed anteriorly	15
15 (14)	Prefemoral process straight, when viewed anteriorly (Fig. 108A). Putnam Co., Tennessee	*N. rhysodesmoides* (Fig. 108)
–	Prefemoral process curves laterally, when viewed anteriorly (Fig. 9A). Prefemoral process with slight constriction at base (Fig. 9C, red triangle). Prefemoral spine lacking. Distal zone twisted dorsolaterally (Fig. 9). Tip directed dorsolaterally, flattened, with enlarged acuminate flange on inner margin (Fig. 9B, red arrow). Rockbridge Co., Virginia	N. scholastica sp. nov. (Figs 9, 10)
16 (10)	Prefemoral basal spine fused with prefemoral process for entire length (Fig. 38A)	17
–	Prefemoral basal spine separate from prefemoral process for some part of length (Fig. 88A)	23
17 (16)	Prefemoral basal spine reduced to a rounded bulge (Fig. 92C)	18
–	Prefemoral basal spine rectangular, shelf-like (Fig. 52C)	19
18 (17)	Prefemoral process tip directed medially (Fig. 92A). Prefemoral process long thin, bent 90° and directed medially at half-way point (Fig. 92A, red arrow). Prefemoral spine reduced and fused with prefemoral process for entire length (Fig. 92C, red triangle). Bland and Tazewell Cos., Virginia, Randolph Co. West Virginia	N. botrydium sp. nov. (Figs 92, 93)
–	Prefemoral process tip directed dorsally (Fig. 63B). Prefemoral spine reduced to slight swelling at base of prefemoral process (Fig. 63C, red triangle). Acropodite tip rounded with small, rounded lateral flange (Fig. 63A, red arrow). Acropodite with medial swelling before apex. Bland Co., Virginia	N. tenuis sp. nov. (Figs 63, 64)
19 (17)	Prefemoral process simple, curving medially (Fig. 52A). Prefemoral tip bent dorsally (Fig. 52B). Prefemoral spine fused to prefemoral process throughout length, forming ridge along base of prefemoral process (Fig. 52C, red triangle). Acropodite distal zone with large, lobed lateral flange (Fig. 52A, red arrow). Southern West Virginia, Southwestern Virginia, Northeastern Tennessee	N. breweri sp. nov. (Figs 52, 53)
–	Prefemoral process variable, though never curving simple, medially	20
20 (19)	Prefemoral process acicular (Fig. 59A)	21
–	Prefemoral process variable, though never acicular	22
21 (20)	Gonopod basal zone height, when viewed anteriorly, subequal to half length of acropodite (Fig. 73A). Acropodite curving dorsomedially with pronounced medial swelling (Fig. 73A). Acropodite tip with large, hooked lateral flange (Fig. 73A, red arrow). Prefemoral spine reduced and fused to prefemoral process, forming small ridge (Fig. 73B, red triangle). Cumberland Co., Tennessee, East to confluence of Tennessee, Kentucky, North Carolina and Virginia	N. blackmountainensis sp. nov. (Figs 73, 74)
–	Gonopod basal zone height, when viewed anteriorly, greater than half length of acropodite (Fig. 59A). Acropodite distal zone heavily reduced, with small, triangular lateral flange (Fig. 59B, red arrow). Prefemoral spine fused to prefemoral process throughout length (Fig. 59A, red triangle). Eastern Tennessee	*N. tennesseensis* (Figs 59, 60)
22 (20)	Prefemoral process laminate (Fig. 67A). Prefemoral process curving dorsomedially (Fig. 67B). Prefemoral spine fused with prefemoral process, reduced to shelf-like ridge at base of prefemoral process (Fig. 67C, red triangle). Acropodite distal zone curving medially, with tip directed caudally (Fig. 67A, red arrow). Mercer Co., West Virginia	N. sheari sp. nov. (Figs 67, 68)
–	Prefemoral process sinuous (Fig. 13A). Prefemoral spine reduced and fused with prefemoral process, forming basal ridge (Fig. 13C, red triangle). Acropodite semi-circular, curving dorsomedially throughout, giving appearance of slight swelling just basal to tip in posterior view (Fig. 13C, red arrow). Carroll Co., Virginia, Surry Co., North Carolina	N. komela sp. nov. (Figs 13, 14)
23 (16)	Prefemoral basal spine large, pronounced (Fig. 57A)	24
–	Prefemoral basal spine small, reduced (Fig. 88C)	43
24 (23)	Prefemoral basal spine rectangular, shelf-like (Fig. 40A). Acropodite long and curving medially before apex. Acropodite tip with small, triangular lateral flange (Fig. 40A, red arrow). Telopodite basal zone height ca. 1/2 length of acropodite. Roanoke Co., Virginia	N. stellapolis sp. nov. (Figs 40, 41)
–	Prefemoral basal spine variable, though never rectangular, shelf-like	25
25 (24)	Prefemoral basal spine pointed, blunt (Fig. 57A)	26
–	Prefemoral basal spine variable, though never pointed, blunt	28
26 (25)	Prefemoral process arising from prefemur (110A). Prefemoral process straight, laminate, separated widely from projected, blunt prefemoral spine (Fig. 94A, red triangle). Acropodite distal zone short, directed dorsomedially, with large, lobed lateral flange (Fig. 94A, red arrow). Sullivan Co. Tennessee	N. tsuga sp. nov. (Figs 94, 95)
–	Prefemoral process arising dorsomedially from prefemoral spine (Fig. 102A)	27
27 (26)	Prefemoral process stout (Fig. 102A). Acropodite tip blunt, with small, triangular lateral flange (Fig. 102A, red arrow). Acropodite swollen before apex (Fig. 102A, red triangle). Telopodite basal zone height > 1/3 length of acropodite. Lee Co. Virginia	N. honeytreetrailensis sp. nov. (Figs 102, 103)
–	Prefemoral process thin, sinuous (Fig. 112A). Meigs Co. Tennessee	*N. sigmoidea* (Fig. 112)
28 (25)	Prefemoral process arising from the top of the prefemoral spine (Fig. 32A)	29
–	Prefemoral process variable, though never arising from the top of the prefemoral spine	32
29 (28)	Prefemoral process sinuous (Fig. 112A). Acropodite apex with distinct constriction (Fig. 96A, red arrow). Tip expanded distally, with small lobed lateral and medial flanges. Acropodite with expanded medial flange (Fig. 96A, red triangle). Morgan Co. Tennessee	N. cryomaia sp. nov. (Figs 96, 97)
–	Prefemoral process straight, acicular (Fig. 32A)	30
30 (29)	Prefemoral process straight, when viewed anteriorly, arising from top of prefemoral spine (Fig. 75A). Prefemoral spine large, projecting, acicular. Acropodite curving medially before apex (Fig. 65). Southeastern Ohio, northwestern West Virginia	*N. ohionis* (Figs 65, 66)
–	Prefemoral process curves medially, when viewed anteriorly (Fig. 32A)	31
31 (30)	Gonopod basal zone height, when viewed anteriorly, subequal to half-length of acropodite (Fig. 73A). Acropodite long, thin, curving medially. Distal zone with medial flange at 135° to solenomere, and lateral flange at 90° angle to solenomere (Fig. 32B, red triangle). Prefemoral process small, thin, curving medially, arising from top of projected, stout prefemoral spine (Fig. 32A, red arrow). Mississippi north to Indiana, west to Arkansas and Missouri	*N. castanea* (Figs 32, 33)
–	Gonopod basal zone height, when viewed anteriorly, less than half length of acropodite (Fig. 35A). Acropodite long and curving ventromedially. Distal zone bent ventroposteriorly, with laminate flange encircling tip forming a hood-like structure around dorso-posteriorly projected, laminate solenomere — laminate flange partially obscuring solenomere when viewed laterally (Fig. 35A, red arrow). Acropodite with small medial flange near apex (Fig. 35C, red triangle). Prefemoral process small, thin, curving medially, arising from top of projected acuminate prefemoral spine. Southwestern Virginia	*N. hokie* (Figs 35–37)
32 (28)	Prefemoral process arising from prefemur (Fig. 98A)	33
–	Prefemoral process arising dorsomedially from prefemoral spine (Fig. 102A)	36
33 (32)	Prefemoral process simple, curving medially (Fig. 98A). Prefemoral spine prominent, claw-like, curving cephalically. Acropodite tip blunt, with small, triangular lateral and medial flanges (Fig. 98A, red arrows). Acropodite swollen before apex (Fig. 98A, red triangle). Greene Co., Tennessee	N. daptria sp. nov. (Figs 98, 99)
–	Prefemoral process variable, though never simple, curving medially	34
34 (33)	Prefemoral process sinuous, bending ventrally before curving cephalolaterally, arising dorsomedially from large, sharp prefemoral spine (Fig. 61B). Distal zone of acropodite curving dorsomedially (Fig. 61B, red arrow). Tip of acropodite with small, rounded lateral flange (Fig. 61A, red triangle). Bland and Wythe Cos., Virginia	N. ignis sp. nov. (Figs 61, 62)
–	Prefemoral process variable, though never sinuous, bending ventrally	35
35 (34)	Prefemoral process curves medially, when viewed anteriorly (Fig. 109A). Prefemoral spine acicular and paralleling prefemoral process (Fig. 109B, red arrow). Acropodite tip simple, blunt. Central North Carolina	*N. conservata* (Fig. 109)
–	Prefemoral process straight, acicular (Fig. 28A). Prefemoral spine sharp, tooth-like, widely separated from prefemoral process (Fig. 28A, red arrow). Acropodite curving medially before apex, distal zone reduced. Henry and Pittsylvania Cos., Virginia	N. hardeni sp. nov. (Figs 28, 29)
36 (32)	Prefemoral process stout, acicular, arising dorsomedially from projected cephalically-curving prefemoral spine (Fig. 90A). Acropodite distal zone curving dorsomedially, tip rounded, directed caudally with small lateral flange (Fig. 90A). Acropodite with slight swelling on inner margin (Fig. 90B, red triangle) and dimple on outer margin (Fig. 90A, red arrow). Bland Co., Virginia	N. asta sp. nov. (Figs 90, 91)
–	Prefemoral process variable, though never stout, acicular	37
37 (36)	Prefemoral process sinuous (Fig. 24A)	38
–	Prefemoral process simple, curving in a single arc (Fig. 104A)	42
38 (37)	Prefemoral process curves medially, when viewed anteriorly (Fig. 26A)	39
–	Prefemoral process curves laterally, when viewed anteriorly (Fig. 61A)	41
39 (38)	Prefemoral process tip directed ventrally (Fig. 26B)	40
–	Prefemoral process tip directed cephalically (Fig. 111). Prefemoral process long, thin, curving medially, arising from sharp projected prefemoral spine (Fig. 111A, red arrow). Acropodite simple, curving medially throughout with extremely short, blunt distal zone. Rutherford Co., North Carolina	*N. rutherfordensis* (Fig. 111)
40 (39)	Basal zone width, when viewed anteriorly, subequal to space between acropodite and prefemoral process at greatest divide (Fig. 30A). Acropodite gradually tapering towards tip, with small, lobed medial flange (Fig. 26A, red triangle). Tip directed medially, rounded and simple. Prefemoral process thin, sinuous, acicular. Prefemoral spine small, medially directed, with secondary hump proximal to acropodite base (Fig. 26C, red arrow). Western West Virginia, northeastern Kentucky	N. mcelroyorum sp. nov. (Figs 26, 27)
–	Basal zone width, when viewed anteriorly, wider than space between acropodite and prefemoral process at greatest divide (Fig. 24A). Acropodite curving medially before bending ventromedially at a nearly 90° angle (Fig. 22B, red triangle). Distal zone short, bent dorsomedially at 90° angle. Acropodite shaft swollen before apex, with medial flange, and sinuous region. Prefemoral process curving ventromedially, coplanar (when viewed medially) with acropodite. Prefemoral spine prominent, acicular. Montgomery Co., Virginia	N. paupertas sp. nov. (Figs 22, 23)
41 (38)	Prefemoral process tip directed ventrally (Fig. 24A). Prefemoral process arising from large, pronounced prefemoral spine (Fig. 24B, red arrow). Acropodite linear, bent medially forming 130° angle with telopodite basal zone. Tip simple, rectangular, bent medially. Carter Co., Kentucky	N. caverna sp. nov. (Figs 24, 25)
–	Prefemoral process tip directed cephalically (Fig. 30A). Prefemoral spine enlarged, cephalically directed. Acropodite with slightly swollen medial area (Fig. 30C, red triangle), and small, lobed medial flange (Fig. 30A, red arrow) before apex. Tip directed medially, blunt and simple. Distal zone greatly reduced. Southwestern Virginia, northeastern North Carolina	N. serpens sp. nov. (Figs 30, 31)
42 (37)	Prefemoral process tip directed medially (Fig. 104A). Prefemoral process paralleling curve of acropodite, arising dorsolaterally from pronounced curving prefemoral spine. Acropodite apex with distinct constriction (Fig. 104A, red arrow), tip sharp, with acuminate, triangular lateral flange. Acropodite with expanded, laminate medial flange (Fig. 104A, red triangle). Campbell Co., Tennessee	N. kassoni sp. nov. (Figs 104, 105)
–	Prefemoral process tip directed ventrally (Fig. 42A). Prefemoral process arising from enlarged prefemoral spine. Acropodite distal zone short, directed medially with small triangular lateral flange (Fig. 42A, red triangle). Floyd and Pulaski Cos., Virginia	N. stellaradix sp. nov. (Figs 42, 43)
43 (23)	Prefemoral basal spine pointed, sharp (Fig. 45C)	44
–	Prefemoral basal spine variable, though never pointed, sharp	52
44 (43)	Prefemoral process arising from prefemur (Fig. 45A)	45
–	Prefemoral process arising dorsomedially from prefemoral spine (Fig. 15A)	50
45 (44)	Prefemoral process sinuous (Fig. 45B)	46
–	Prefemoral process variable, though never sinuous	47
46 (45)	Prefemoral process curves laterally when viewed anteriorly (Fig. 45A). Acropodite shaft swollen before apex, with cingulum (Fig. 45A, red rectangle). Prefemoral spine greatly reduced to small, tooth-like projection (Fig. 45C, red oval). Acropodite distal zone short, bent dorsomedially at 45° angle. Tip blunt with small, lobed lateral flange (Fig. 45C, red arrow). Telopodite basal zone with lateral bulge (Fig. 45A, red triangle). West Virginia panhandle and eastern Virginia	N. cingulata sp. nov. (Figs 45–47)
–	Prefemoral process fairly straight when viewed anteriorly (Fig. 19A), arising from medial side of shelf-like prefemoral spine (Fig. 19C, red arrow). Acropodite simple, without cingulum or modifications. Distal zone short, simple, at nearly 90° angle to acropodite. Border region between central West Virginia and Virginia	*N. simplex* (Fig. 19)
47 (45)	Prefemoral process acicular (Fig. 34A)	48
–	Prefemoral process laminate and sinuous (Fig. 48A). Prefemoral spine reduced to small, acuminate projection (Fig. 48C, red arrow). Acropodite semi-circular, curving dorsomedially with abrupt 90° curve after apex. Acropodite tip simple, directed posteriorly, terminating in sharp, claw-like point. Central to southwestern Virginia, southeastern West Virginia	*N. laminata* (Figs 48, 49)
48 (47)	Prefemoral process straight, when viewed anteriorly, arising from small, blunt prefemoral spine (Fig. 34A). Acropodite distal zone with medial and lateral flanges at 90° angle to solenomere (Fig. 34A, red arrow). Acropodite with small medial flange near apex (Fig. 34A, red triangle). Newton Co., Arkansas	*N. davidcauseyi* (Fig. 34)
–	Prefemoral process otherwise	49
49 (48)	Prefemoral process curving medially, when viewed anteriorly (Fig. 55A). Prefemoral spine reduced to thorn-like structure, arising from prefemoral process, directed ventrally (Fig. 55B, red triangle). Acropodite distal zone short, rounded. Floyd Co., Georgia	N. fritzae sp. nov. (Figs 55, 56)
–	Prefemoral process curves laterally, when viewed anteriorly (Fig. 106A). Prefemoral spine small, curving cephalically (Fig. 106B, red triangle). Acropodite distal zone extremely short, blunt. Northern border of Tennessee and North Carolina, and Loudon Co., Tennessee	*N. minor* (Figs 106, 107)
50 (44)	Prefemoral process sinuous, arising from base of stout, tooth-like prefemoral spine, paralleling medial curve of acropodite (Fig. 15A). Acropodite distal zone short, curving medially forming 130° angle with acropodite (Fig. 15). Tip and distal zone simple, rectangular, blunt. Northeastern Virginia and eastern West Virginia north to southwestern New York	*N. fowleri* (Figs 15, 16)
–	Prefemoral process otherwise	51
51 (50)	Prefemoral process stout, curving laterally (Fig. 17A). Prefemoral spine stout, tooth-like (Fig. 17C, red arrow). Alleghany and Rockbridge Cos., Virginia	N. piccolia sp. nov. (Figs 17, 18)
–	Prefemoral process simple, curving medially (Fig. 50A). Prefemoral process arising dorsally from prefemoral spine (Fig. 50A, red arrow), paralleling medial curve of acropodite. Prefemoral spine large, curving cephalically. Eastern West Virginia	N. tasskelsoae sp. nov. (Figs 50, 51)
52 (43)	Prefemoral basal spine reduced to rounded bulge (Fig. 78C). Prefemoral process dorsomedially curving. Acropodite tip with prominent triangular lateral flange curving abruptly at 90° angle towards tip (Fig. 78A, red triangle). Acropodite with laminate medial flange just proximal to tip. Davidson and Marshall Cos. Tennessee	*N. dilatata* (Figs 78, 79)
–	Prefemoral basal spine rectangular in conformation, shelf-like (Fig. 75C)	53
53 (52)	Prefemoral process long, acicular (Fig. 75A). Prefemoral spine pronounced, sharp, partially fused to prefemoral process forming a ridge (Fig. 75C, red triangle). Acropodite acicular, bending abruptly medially at 90° angle at tip. Distal zone quadrate. Acropodite with small, shelf-like medial flange just before tip (Fig. 17C, red arrow). Telopodite basal zone with slight medial swelling (Fig. 75A, red circle). Harlan and Leslie Cos., Kentucky	N. bobmareki sp. nov. (Figs 75, 76)
–	Prefemoral process large, laminate and sinuous (Fig. 11A). Prefemoral spine reduced to small, rectangular shelf. Acropodite semi-circular, gently curving dorsomedially throughout. Acropodite tip with small, triangular lateral flange (Fig. 11A, red triangle). Tip terminating in sharp, caudally-directed point. Watauga Co., North Carolina	*N. domestica* (Figs 11, 12)

Figure 8.

Two general types of gonopods in the minor species group of Nannaria Chamberlin, 1918 A Nannaria ambulatrix sp. nov. (non-type ♂, VTEC, MPE03794), fairly straight gonopods which do not cross in situ B N. hokie Means, Hennen & Marek, 2021 (paratype ♂, VMNH, MPE00886), more highly curved gonopods which do cross in situ.

minor species group diagnosis

Species in the minor species group differ from species in the wilsoni species group based on the presence of the following characters (best viewed with ≥ 20× magnification): acropodite often highly setaceous throughout, not with setation ending at midpoint as in the wilsoni species group. Acropodite anterior bend lacking twist as seen in the wilsoni species group. Prefemoral process often with prefemoral spine, lacking in the wilsoni species group. Without cyphopod receptacle modification and expanded 2^nd coxae of females, as is present in the wilsoni species group.

Species in the minor species group may be confused in field with Gyalostethus monticolens (Chamberlin, 1951), Boraria Chamberlin, 1943, Howellaria Hoffman, 1950, and Idaloria Marek, Means & Hennen, 2021, due to similarity in body length (average 27 mm) and relatively simple acropodites. However, the minor species group differs based on the presence of the following characters (best viewed with ≥ 20× magnification): Males with spatulate pregonopodal claws (Fig. 2), not bisinuate or uncinate. Acropodite typically thin, long and continually curving; if straight, lacking S-shaped acropodite tip as in G. monticolens. Acropodite tip rarely directed cephalically. With triangular sternal projections, not lacking as in Gyalostethus Hoffman, 1965. Color in life without yellow paranotal spots as in Boraria stricta, or tan/brown as in G. monticolens.

scholastica clade

Components. Nannaria scholastica sp. nov. Based on the molecular phylogeny (Fig. 7), Nannaria scholastica sp. nov. is sister to the remaining species of the minor species group. In combination with morphological characters detailed in the below diagnosis, N. scholastica sp. nov. is a member of its own monotypic clade (Fig. 113). The basal placement of this species is not entirely surprising, as it has simple gonopods, yet lacks a prefemoral spine, therefore only partially conforming with the rest of the minor group. The geographic isolation of this species is worrisome for its conservation. Hoffman had only one specimen (a male) in the VMNH collection, from the campus of Washington and Lee University (WLU). JCM and DAH discovered an abundant and seemingly healthy population during a visit to WLU, however repeated collection trips by the authors, Hoffman, and collaborators in the surrounding mountains have revealed no additional populations. This species may be a relict that is in threat of extinction should the small forest that it now occupies be destroyed or altered.

Distribution. The campus of WLU (Fig. 114, black dot).

Nannaria scholastica sp. nov.

http://zoobank.org/97C73812-E51C-4BCB-9D97-9F145A01590A

Figs 9, 10 Vernacular name: “The Scholarly Twisted-Claw Millipede”

Material examined

Holotype: United States – Virginia • ♂; Rockbridge County, Lexington, Washington and Lee University campus, hillside beside community gardens; 37.7956°N, -79.4427°W; elev. 330 m; 14 Nov. 2017; hand collected; J. Means, D. Hennen leg.; VTEC MPE03485.

Paratypes: United States – Virginia • 4 ♂♂; same collection data as holotype; VTEC MPE03486, 3498, 3500, 3517 • 3 ♂♂; same collection data as holotype; VMNH MPE03516, 3518, 3625 • 1 ♀; same colletion data as holotype; VTEC MPE03499 • 1 ♂; Rockbridge County, W & L campus Lexington; 37.7956°N, -79.4427°W; 20 Nov. 1948; R. Hoffman leg.; VMNH NAN0306. For detailed collection data see Suppl. material 7.

Diagnosis

Adult males of Nannaria scholastica sp. nov. are distinct from other Nannaria and the nearby N. shenandoa, based on the following combination of characters: Gonopods. Gonopodal acropodite gently curving ventromedially before apex, not straight as in N. terricola. Distal zone twisted dorsolaterally (Fig. 9). Tip directed dorsolaterally, flattened, with enlarged acuminate flange on inner margin (Fig. 9B, red arrow), not with folds, grooves, flanges as N. shenandoa. Telopodite lacking swollen basal zone of N. terricola and heavily reduced basal zone of N. shenandoa. Prefemur with curved, saber-like prefemoral process, not straight as in N. terricola. Prefemoral process (when viewed medially) not crossing the acropodite as in N. shenandoa. Prefemoral process slightly constricted at base (Fig. 9C, red triangle), one-half length of acropodite. Prefemur lacking prefemoral spine. Color. Tergites with white paranotal spots (Fig. 10). Dark brown background. Dorsum of collum smooth with white margin.

Figure 9.

Nannaria scholastica sp. nov. paratype ♂ (VMNH, NAN0306) left gonopod A anterior view B medial view; red arrow indicates medial flange C posterior view; red triangle indicates slight prefemoral process constriction. Scale bar: 0.5 mm.

Figure 10.

Nannaria scholastica sp. nov. holotype ♂ (VTEC, MPE03485) coloration. Scale bar: 4.0 mm.

Measurements

♂ holotype (VTEC, MPE03485): BL = 32.5, CW = 4.2, IW = 2.1, ISW = 0.9, B11W = 5.4, B11H = 3.2; ♀ paratype (VTEC, MPE03499): BL = 34.2, CW = 3.9, IW = 2.5, ISW = 1.0, B11W = 5.4, B11H = 3.8.

Variation

No known variation.

Distribution

Known only from the type locality (Virginia: Rockbridge County, Suppl. material 7; Figs 114, 126). Distribution area: N/A; status: MRE.

Ecology

Individuals of N. scholastica sp. nov. were collected from a mesic mixed pine-deciduous forest composed of maple, tuliptree, black cherry, beech, and white pine. They were found under leaf litter on top of the soil. Nannaria scholastica sp. nov. was found along with an undescribed species of Nannaria in the wilsoni species group; however, the two species had ostensibly stratified themselves in two longitudinal strips, with the wilsoni species at the top of the hill, and N. scholastica sp. nov. ca. 10 m lower down on the slope. Similar specific elevational spacing has been observed in N. serpens sp. nov. and N. stellaradix sp. nov.; however, in this case the minor group N. serpens sp. nov. and N. stellaradix sp. nov. were found from the higher elevation areas of the hill, while the undescribed wilsoni group species were found at the bottom of the hill.

Etymology

This species is named for its discovery on the campus of an academic institution, Washington and Lee University. The specific name scholastica is Latin for academic, and is a feminized adjective.

Type locality

United States, Virginia, Rockbridge County, Lexington, Washington and Lee University campus hillside beside community gardens, 37.7956°N, -79.4427°W.

domestica clade

Components. Nannaria domestica Shelley, 1975, N. komela sp. nov., and a female from the I-77 NC Welcome Center. Members of the domestica clade share a curving, semi-circular acropodite and laminate prefemoral process. Surprisingly, N. domestica and N. komela sp. nov. are not closely related to N. laminata, despite N. laminata also having circular gonopods with laminate prefemoral processes. Due to the simplicity of these forms, however, it is likely that similarity between the N. domestica clade and N. laminata is due to convergence, rather than a shared evolutionary history. Based on the molecular phylogeny (Fig. 7) the female from the I-77 NC Welcome Center would appear to be the same species as N. komela sp. nov.

Distribution. the domestica clade is known from a small area in southwestern Virginia and northwestern North Carolina (Fig. 114).

Nannaria domestica Shelley, 1975

Figs 11, 12 Vernacular name: “The Domestic Twisted-Claw Millipede”

Nannaria domestica Shelley, 1975: 186, figs 10–12. Hoffman 1999: 366. Shelley 2000: 196. Marek et al. 2014: 36. Means et al. 2021: S69.

Material examined

United States – North Carolina • 1 ♂; Watauga County, Cliff Dwellers Inn, side of road, slope; 36.1386°N, -81.6694°W; elev. 1018 m; 25 Apr. 2015; hand collected; J. Means leg.; VTEC MPE00305 • 6 ♀♀; same collection data as preceding; VTEC MPE00303, 310–313, 315 • 1 ♂; Watauga County, Blowing Rock, Goforth Rd., 0.5 N US 321; 36.1293°N, -81.6619°W; 11 Oct. 1975; R. Shelley leg.; NCSM NAN0510; 3 ♂♂, 1 ♀; same collection data as preceding; 8 Sep. 1973; NCSM NAN0525. For detailed collection data see Suppl. material 7.

Diagnosis

Adult males of N. domestica are distinct from other Nannaria and the nearby N. blackmountainensis sp. nov. based on the following combination of characters: Gonopods. Gonopodal acropodite semi-circular, gently curving dorsomedially throughout, not with abrupt 90° angle after apex as in N. laminata or pre-apex swelling as in N. blackmountainensis sp. nov. Acropodite tip with small, triangular lateral flange (Fig. 11A, red triangle), not hooked as in N. blackmountainensis sp. nov. and not simple, lacking flange as in N. laminata. Tip terminating in sharp, caudally-directed point, not blunt, rectangular as in N. blackmountainensis sp. nov. Height of telopodite basal zone ca. 1/4 length of acropodite, not > 1/3 length of acropodite as in N. laminata and ca. 1/2 length as in N. blackmountainensis sp. nov. Prefemoral process large, laminate and serpentine with prefemoral spine reduced to small, rectangular shelf, not fused with prefemoral process as in N. blackmountainensis sp. nov. Color. Tergites with orange paranotal spots (Fig. 12). Dark brown background. Dorsum of collum smooth with orange margin.

Figure 11.

Nannaria domestica ♂ (VTEC, MPE00305) left gonopod A anterior view; red triangle indicates lateral flange B medial view C posterior view. Scale bar: 0.5 mm.

Figure 12.

Nannaria domestica ♂ (VTEC, MPE00305) coloration. Scale bar: 4.0 mm.

Measurements

♂ (VTEC, MPE00305): BL = 28.6, CW = 3.3, IW = 1.8, ISW = 0.8, B11W = 4.2, B11H = 2.7; ♀ (VTEC, MPE00312): BL = 31.9, CW = 3.9, IW = 1.8, ISW = 0.9, B11W = 4.9, B11H = 3.5.

Variation

No known variation

Distribution

Known only from Blowing Rock, Watauga County, North Carolina (Suppl. material 7; Fig. 126). Distribution area: N/A; status: MRE.

Ecology

Individuals of N. domestica have been collected from under leaf litter in mesic broadleaved forests composed of oak, maple, and rhododendron.

Etymology

Shelley (1975) gives no etymology for this name, however it is reasonable to assume that the name refers to the type specimen being collected from a residence.

Type locality

United States, North Carolina, Watauga County, Blowing Rock, from the yard of residence on Goforth Road, 0.5 miles north U.S. highway 321.

Notes

In the original paper, Shelley (1975: 186) examined and designated the holotype male (NCSM 1007) collected by himself on October 16, 1971.

Nannaria komela sp. nov.

http://zoobank.org/DB1B3BA2-0240-4EF7-B387-AEB0C7AC62E1

Figs 13, 14 Vernacular name: “The Lambsburg Twisted-Claw Millipede”

Material examined

Holotype: United States – Virginia • ♂; Carroll County, Lambsburg, Lambsburg Rd., Hawks State Forest, hillside above Turkey Creek; 36.6062°N, -80.7720°W; elev. 767 m; 7 Oct. 2017; hand collected; J. Means, D. Hennen leg.; VTEC MPE03523.

Paratypes: United States – Virginia • 1 ♂; same collection data as holotype; VTEC MPE03536 • 1 ♂; same collection data as holotype; VMNH MPE03537 • 1 ♀; same collection data as holotype; VTEC MPE03699 • 1 ♀; same collection data as holotype; VMNH MPE03700 • 1 ♂; Carroll County, crest of Blue Ridge at end of VA. Hy. 716, ca. 3 miles W of Lambsburg; 36.5852°N, -80.8155°W; 3 June 1986; R. Hoffman, R. Highton leg.; VMNH NAN0295.

Other material

United States – North Carolina • 1 ♀; Surry County, gully behind North Carolina Welcome Center; 36.5605°N, -80.7469°W; elev. 421 m; 24 June 2017; hand collected; J. Means leg.; VTEC MPE02891 • 1 ♂; Surry County, 8NW Dobson, 1338, 2^nd bridge, 0.5 S 1421; 36.4719°N, -80.8355°W; 11 Aug. 1975; R. Shelley, J. Clamp leg.; NCSM NAN0506 • 4 ♂♀; Surry County, along 1328, 1.8 mi from int. of 1325 on hillside in dry stream bed, 11.2 mi W Dobson; 36.4181°N, -80.9283°W; 7 Sep. 1978; W. Jones leg.; NCSM NAN0512; SCAU – Virginia • 1 ♂; Russell County, 4200’ Beartown Mtn.; 36.9360°N, -81.8859°W; 16 Oct. 1976; C. Rushin leg.; VMNH NAN0141. For detailed collection data see Suppl. material 7.

Diagnosis

Adult males of Nannaria komela sp. nov. are distinct from other Nannaria and the nearby N. wilsoni, based on the following combination of characters: Gonopods. Gonopodal acropodite semi-circular, curving dorsomedially throughout, with appearance of slight swelling before apex in posterior view (Fig. 13C, red arrow), not with corkscrew before apex as in N. wilsoni. Tip simple, rounded, curving dorsally. Tip not sharp, with small triangular lateral flange as in N. domestica, or with laminate expansion and tooth-like lateral flange as in N. wilsoni. Height of telopodite basal zone reduced, ca. 1/4 length of acropodite, not ca. 1/6 as in N. wilsoni. Prefemoral process serpentine, curving ventrally. Prefemoral process not laminate as in N. domestica, or long, curving medially as in N. wilsoni. Prefemoral spine reduced and fused with prefemoral process, forming basal ridge (Fig. 13C, red triangle), not separate as in N. domestica. Color. Tergites with orange paranotal spots (Fig. 14). Black background. Dorsum of collum smooth with orange margin.

Figure 13.

Nannaria komela sp. nov. paratype ♂ (VMNH, NAN0295) left gonopod A anterior view B medial view C posterior view; red arrow indicates dorsomedial bend giving appearance of medial swelling; red triangle indicates prefemoral spine fused with prefemoral process. Scale bar: 0.5 mm.

Figure 14.

Nannaria komela sp. nov. holotype ♂ (VTEC, MPE03523) coloration. Scale bar: 4.0 mm.

Measurements

♂ holotype (VTEC, MPE03523): BL = 28.1, CW = 3.6, IW = 2.2, ISW = 0.8, B11W = 4.6, B11H = 3.0; ♀ paratype (VTEC, MPE03699): BL = 33.3, CW = 3.8, IW = 2.0, ISW = 1.0, B11W = 5.0, B11H = 3.3.

Variation

The specimen from Beartown Mountain (VMNH NAN0141) has a slightly less serpentine prefemoral process, and a more pronounced prefemoral spine.

Distribution

Known from southwestern Virginia and northwestern North Carolina (Virginia: Carroll and Russell counties; North Carolina: Surry County, Suppl. material 7; Fig. 126). Distribution area: 70 km²; status: MRE.

Ecology

Individuals of Nannaria komela sp. nov. have been collected from mesic deciduous forests dominated by oak, maple, tuliptree, and rhododendron. They are often found under 1–2 cm moist soil on hillsides.

Etymology

The specific epithet is an arbitrary combination of letters from the Greek kome, meaning ‘village,’ and melon, meaning ‘sheep,’ which refers to the town of Lambsburg, Virginia nearby to the type locality. It is to be treated as a noun in apposition.

Type locality

United States, Virginia, Carroll County, Lambsburg, Lambsburg Rd., Hawks State Forest, hillside above Turkey Creek, 36.6062°N, -80.7720°W.

fowleri clade

Components. Nannaria fowleri Chamberlin, 1947, N. piccolia sp. nov., N. simplex Hoffman, 1949, and a female specimen from Mill Gap, VA. Members of the fowleri clade have simple gonopods that have a gradually curving acropodite with the tip bent at a 90° to the main branch. Hoffman labeled the specimens he collected around Longdale Mines, Virginia, as topotypes of N. simplex, however, after inspecting the N. simplex type material we do not agree that these forms are representatives of the same species. Nannaria simplex has a long, thin prefemoral process, while N. piccolia sp. nov. specimens have short, stout, claw-like prefemoral processes (Fig. 17). Unfortunately, while JCM visited the type locality of N. simplex (McGraw’s Gap, Virginia) three times, he was unable to recover specimens of N. simplex to test this relationship with molecular data (instead only finding an undescribed species in the wilsoni group, despite McGraw’s Gap being the type locality for both N. simplex and N. ericacea). However, due to the close geographic proximity and the shared gonopodal characters listed above, N. simplex is ostensibly within the fowleri clade.

Distribution. the fowleri clade extends from western Virginia, to eastern West Virginia, to western Maryland (The Allegheny Mountains), up through central Pennsylvania and into eastern New York (Fig. 114).

Nannaria fowleri Chamberlin, 1947

Figs 15, 16 Vernacular name: “Fowler’s Twisted-Claw Millipede”

Nannaria fowleri Chamberlin, 1947: 29, fig. 14. Chamberlin and Hoffman 1958: 40. Shelley 1988: 1656. Hoffman 1999: 366. Shelley 2002: 1872. Marek et al. 2014: 37. Means et al. 2021: S70.

Nannaria cayugae Chamberlin, 1949: 4, fig. 3. Chamberlin and Hoffman 1958: 40. Shelley 1988: 1656.

Material examined

United States – Maryland • 1 ♂; Allegany County, E of Flinstone, 1600 ft. summit of Town Hill Mtn.; 39.6903°N, -78.4067°W; 19 May 1958; L. Hubricht leg.; VMNH NAN0352 • 1 ♂; Allegany County, Cumberland, Knobby Mountain; 39.6527°N, -78.7626°W; 29 March; K. Holbrook; NCSM NAN0463; SCAU – New York • 1 ♂; Cattaraugus County, Allegany St. Pk., Anderson Trail; 42.0023°N, -78.8357°W; 10 May 1958; Muchmore leg.; VMNH NAN0347 • 1 ♂; McKean County, 8 Oct. 1959; VMNH NAN0348 • 1 ♂; Monroe County, Mendon Ponds Park; 42.9988°N, -77.5641°W; 17 Apr. 1958; Muchmore leg.; VMNH NAN0344; SCAU – Pennsylvania • 1 ♂; Centre County, Toftrees State Game Lands; 40.9680°N, -77.9154°W; 1993; T. McCoy leg.; NCSM NAN0451 • 3 ♂♂; Clinton County, Rauchtown, Ravensburg State Park, bank of Rauchtown Run, north of restrooms beside playfield; 41.1034°N, -77.2435°W; elev. 368 m; 18 Sep. 2017; hand collected; J. Means & D. Hennen leg.; VTEC MPE03071, 3094, 3906 • 10 ♀♀; same collection data as preceding; VTEC MPE03072, MPE03095–102, 3907 • 1 ♂; Franklin County, Caledonia State Park, Thaddeus Stevens Trail; 39.9079°N, -77.4777°W; elev. 287 m; 15 Sep. 2017; hand collected; J. Means, D. Hennen leg.; VTEC MPE03017 • 1 ♂; Franklin County, low woods, near creek 1.8 miles S of Caledonia; 39.7359°N, -77.4967°W; 8 June 1956; L. Hubricht leg.; VMNH NAN0345 • 1 ♂; Franklin County, road to Cowants Gap State Park; 39.9957°N, -77.9154°W; 11 Apr.1959; R. Mighton leg.; VMNH NAN0349 • 2 ♂♂; Potter County, Patterson Park, Susquehanna State Forest, 10 mi SSE of Coudersport, roadside logs; 41.6406°N, -77.9471°W; 20 Aug. 1971; W. Shear leg.; VMNH NAN0050 • 1 ♂; Potter County, Austin Dam ruins; 41.6944°N, -78.1197°W; 12 June; W. Shear leg.; VMNH NAN0342 • 1 ♂; Potter County, Denton Hill State Park, 14 mi. E of Coudersport, beech-birch-maple litter; 41.7685°N, -77.8334°W; 22 Aug. 1971; W. Shear leg.; VMNH NAN0037 • 1 ♂; Potter County, Rock Ridge Road, 2 mi from Lyman Run State Park, wet birch logs near spring; 41.6609°N, -77.8864°W; 23 Aug. 1971; W. Shear leg.; VMNH NAN0033; SCAU – Virginia • 5 ♂♂♀♀; Bath County, GWNF Long Spring Run above Little Back Creek; 38.2213°N, -79.8387°W; 24 July 1992; J. Pagels, D. Kobuszewski leg.; VMNH NAN0651 • 1 ♂; same collection data as preceding; 11 Sep. 1992; D. Kobuszewski leg.; VMNH NAN0654 • 2 ♂♂; same collection data as preceding; 17 July 1992; J. Pagels leg.; VMNH NAN0659 • 1 ♂; Bath County, “F” GWNF H’Town; 37.9993°N, -79.8315°W; 1 July 1992; J. Pagels leg.; VMNH NAN0653 • 4 ♂♂; Highland County, Locust Spring Rec. Area, 8 mi. NW of Hightown; 38.5152°N, -79.6899°W; elev. 1158 m; 23 May 1973; R. Hoffman leg.; VMNH NAN0343 • 1 ♂; same collection data as preceding; 28 Apr. 1972; R. Hoffman, Knight leg.; VMNH NAN0346 • 1 ♂; Highland County, GWNF west bank, Laurel Fork (between Bear Hollow & Newman Rd.); 38.2862°N, -79.7468°W; 17 May – 15 June 1993; VMNH NAN0658; SCAU – West Virginia • 2 ♀♀; Pendleton County, old road headed up to Spruce Knob; 38.6947°N, -79.5146°W; elev. 1010 m; 24 May 2015; hand collected; M. Kasson leg.; VTEC MPE00433, 446 • 1 ♂; Pendleton County, Spruce Knob; 38.7308°N, -79.4913°W; elev. 1112 m; 2016; C. Stauder leg.; VTEC MPE01816. For detailed collection data see Suppl. material 7.

Diagnosis

Adult males of N. fowleri are distinct from other Nannaria and the sympatric N. shenandoa, based on the following combination of characters: Gonopods. Gonopodal acropodite gradually curving medially before apex, not straight as in N. shenandoa. Distal zone short, bent medially forming 130° angle with acropodite (Fig. 15). Telopodite with basal zone > 1/4 length of prefemoral process. Tip and distal zone simple, rectangular, blunt, < 1/8 length of acropodite, not large, curving with flange as in N. shenandoa. Prefemoral process arising from base of prefemoral spine, paralleling medial curve of acropodite (Fig. 15A), not directed laterally and crossing acropodite when viewed medially as in N. shenandoa or arising from top of prefemoral spine as in N. tasskelsoae sp. nov. Prefemur with stout, tooth-like prefemoral spine (Fig. 15C, red triangle). Color. Tergites with orange paranotal spots (Fig. 16). Dark brown background. Dorsum of collum smooth with orange margin. Chamberlin (1947) described the holotype as having yellow paranota, but his description was from a preserved specimen which had likely lost its color due to its preservation in alcohol.

Figure 15.

Nannaria fowleri Chamberlin, 1947, ♂ (VTEC, MPE03017) left gonopod A anterior view B medial view C posterior view; red triangle indicates stout, tooth-like prefemoral spine. Scale bar: 0.5 mm.

Figure 16.

Nannaria fowleri Chamberlin, 1947, ♂ (VTEC, MPE03017) coloration. Scale bar: 4.0 mm.

Measurements

♂ (VTEC, MPE03017): BL = 25.1, CW = 3.7, IW = 2.3, ISW = 0.8, B11W = 4.6, B11H = 2.9; ♀ (VTEC, MPE03098): BL = 28.8, CW = 3.7, IW = 2.3, ISW = 0.8, B11W = 4.4, B11H = 3.1.

Variation

A male taken from Cumberland, Maryland (NCSM NAN0463), has a slight acropodite medial swelling, which has not been observed in other individuals of N. fowleri.

Distribution

Nannaria fowleri has a larger distribution than most minor species group Nannaria, perhaps due to post-glacial northward expansion from the eastern Appalachians (Virginia, West Virginia) north into Maryland, Pennsylvania and New York (Virginia: Bath and Highland counties; West Virginia: Pendleton and Pocahontas counties; Maryland: Allegany County; Pennsylvania: Centre, Clinton, Franklin, McKean, and Potter counties; New York: Cattaraugus and Monroe counties; Suppl. material 7; Fig. 127). Distribution area: 43,054 km²; status: WRE.

Ecology

Individuals of N. fowleri have been collected from mesic hardwood forests composed of maple, hemlock, and scattered pine, often buried 1–2 cm under dark, sandy soil.

Etymology

Chamberlin (1947) gave no etymology for the name fowleri but it is reasonable to assume that the name refers to one of the collectors of the holotype, H. W. Fowler.

Type locality

United States, Maryland, Garrett County, Jennings.

Notes

In the original description, Chamberlin (1947: 29) examined and designated the holotype male (ANSP Type no. 9951) collected in July 1907 by W. Stone and H. W. Fowler.

Nannaria piccolia sp. nov.

http://zoobank.org/5CF65D66-4448-4ADA-9C6E-086B1614C70B

Figs 17, 18 Vernacular name: “The Lichen-Loving Twisted-Claw Millipede”

Material examined

Holotype: United States – Virginia • ♂; Rockbridge County, 1.5 air miles NW of Collierstown, Lake Robertson Recreation Area, Mountain Trail, hillside near Hawks Creek stream crossing; 37.8065°N, -79.6152°W; elev. 457 m; 20 Feb. 2018; hand collected; J. Means, D. Hennen leg.; VTEC MPE03809.

Paratypes: United States – Virginia • 1 ♂; same collection data as holotype; VTEC MPE03812 • 1 ♂; same collection data as holotype; VMNH MPE03816 • 2 ♀♀; same collection data as holotype; VTEC MPE03811, 17 • 1 ♀; same collection data as holotype; VMNH MPE03818.

Other material

United States – Virginia • 1 ♂; Alleghany County, Longdale Mines; 37.8083°N, -79.6834°W; 15 Sep. 1948; R. Hoffman leg.; VMNH NAN0645 • 1 ♂; Rockbridge County, Rockbridge Alum Springs, 8 mi SW Goshen; 37.9086°N, -79.6123°W; 1 June 1970; Newman leg.; VMNH NAN0647. For detailed collection data see Suppl. material 7.

Diagnosis

Adult males of Nannaria piccolia sp. nov. are distinct from other Nannaria and the sympatric N. shenandoa, based on the following combination of characters: Gonopods. Gonopodal acropodite gradually curving medially before apex, not straight as in N. shenandoa. Distal zone short, bent medially forming 130° angle with acropodite (Fig. 17). Telopodite basal zone > 1/3 length of acropodite, not < 1/3 as in N. fowleri and N. shenandoa. Tip and distal zone simple, rectangular, blunt, < 1/8 length of acropodite, not large, curving with flange as in N. shenandoa. Prefemoral process short, laterally curved, arising dorsomedially from prefemoral spine, not long, serpentine, paralleling curve of acropodite as in N. fowleri, or crossing over acropodite as in N. shenandoa. Space between prefemoral process and acropodite wider than telopodite basal zone, not thinner as in N. fowleri. Prefemur with stout, tooth-like prefemoral spine (Fig. 17C, red arrow). Color. Tergites with either orange or white paranotal spots (Fig. 18A, B) and occasionally orange paranotal spots with orange stripes (Fig. 18 C). Brown to black background. Dorsum of collum smooth with either orange or white margin, depending on color morph.

Figure 17.

Nannaria piccolia sp. nov. holotype ♂ (VTEC, MPE03809) left gonopod A anterior view B medial view C posterior view; red arrow indicates stout, tooth-like prefemoral spine. Scale bar: 0.5 mm.

Figure 18.

Nannaria piccolia sp. nov. coloration, A holotype ♂ (VTEC, MPE03809) orange paranotal spots B paratype ♂ (VTEC, MPE03812) white paranotal spots C paratype ♀ (VTEC, MPE03811) orange paranotal spots with orange stripes. Scale bar: 4.0 mm.

Measurements

♂ holotype (VTEC, MPE03809): BL = 26.4, CW = 3.9, IW = 2.4, ISW = 0.8, B11W = 5.0, B11H = 2.9; ♀ paratype (VTEC, MPE03811): BL = 36.5, CW = 4.3, IW = 2.7, ISW = 0.9, B11W = 5.5, B11H = 4.0

Variation

No known variation.

Distribution

Known from a small area in western Virginia (Virginia: Allegany and Rockbridge counties, Suppl. material 7; Fig. 126). Distribution area: 35 km²; status: MRE.

Ecology

Individuals of Nannaria piccolia sp. nov. have been collected from mesic deciduous forests dominated by oak, maple, and hickory, typically found under leaf litter on hillsides.

Etymology

Named as a gesture of goodwill to the community of lichen specialists, in honor of the lichen species Piccolia nannaria (Tuck.) Lendemer & Beeching. The specific epithet is a noun in apposition.

Type locality

United States, Virginia, Rockbridge County, 1.5 air miles NW of Collierstown, Lake Robertson Recreation Area, Mountain Trail, hillside near Hawks Creek stream crossing, 37.8065°N, -79.6152°W.

Nannaria simplex Hoffman, 1949

Fig. 19 Vernacular name: “The Simple Gonopod Twisted-Claw Millipede”

Nannaria simplex Hoffman, 1949: 384, figs 13, 14. Chamberlin and Hoffman 1958: 42. Hoffman 1999: 368. Marek et al. 2014: 38. Means et al. 2021: S72.

Material examined

Holotype: United States – Virginia • ♂; Alleghany County, McGraw Gap, 3 miles northwest of Clifton Forge; [37.8586°N, -79.8660°W]; 19 June 1947; R. Hoffman leg.; NMNH #1807.

Other material

United States – Virginia • 2 ♂♂; Augusta County, 5 mi SW of Reddish Knob, FS 85, “mature site” pitfalls; 38.4102°N, -79.3066°W; 28 May 1988; K. Buhlmann leg.; VMNH NAN0353 • 1 ♂; Bath County, Jewel Hole Hollow, off FR 141, just E of upper storage reservoir, S of Paddy Knob; 38.2427°N, -79.7955°W; elev. 914 m; 10 June 1997; MeShean Project leg.; VMNH NAN0646 • 23 ♂♀; Rockingham County, Shenandoah Mountain, DF site off Va. 924, ca. 0.5 mi E of WVA state line, jct with FS 85 (the “8 year old stand” of Plethodon punctatus study); 38.4783°N, -79.2136°W; 17 June 1988; K. Buhlmann leg.; VMNH NAN0350; SCAU – West Virginia • 1 ♂; Pendleton County, Moyers, ca. 5 mi. SE of Franklin; 38.5154°N, -79.3621°W; 5 May 1961; D. Whitehead leg.; VMNH NAN0351. For detailed collection data see Suppl. material 7.

Diagnosis

Adult males of Nannaria simplex are distinct from other Nannaria and the sympatric wilsoni species group, N. ericacea, based on the following combination of characters: Gonopods. Gonopodal acropodite gently curving medially basal to apex, not nearly straight basal to apex as in N. ericacea. Distal zone short, simple, at nearly 90° bend to acropodite — not serpentine, with prominent lateral flange as in N. ericacea. Telopodite basal zone height ca. 1/2 length of acropodite, not < 1/3 length as in N. fowleri and N. ericacea. Prefemur with thin, somewhat serpentine prefemoral process arising from medial side of prefemoral spine, directed caudally—not paralleling acropodite as in N. fowleri, or large, crossing under acropodite distal zone as in N. ericacea. Prefemoral spine shelf-like (Fig. 19C, red arrow)—not pronounced and tooth-like as in N. fowleri, or lacking as in N. ericacea. Color. Hoffman (1949) described N. simplex as having “reddish pink” tergites with a black background. Collum completely black.

Figure 19.

Nannaria simplex Hoffman, 1949, holotype ♂ left gonopod (NMNH #1807) A anterior view B medial view C posterior view, red arrow indicates shelf-like prefemoral spine. Scale bar: 0.5 mm.

Measurements

♂ holotype (NMNH, #1807): BL = 29.4, CW = 3.9, IW = 2.6, ISW = 0.9, B11W = 4.5, B11H = 3.2.

Variation

No known variation.

Distribution

Known from the border of Virginia and West Virginia near the Shenandoah Mountains (Virginia: Alleghany, Augusta, Bath, and Rockingham counties; West Virginia: Pendleton County; Suppl. material 7; Fig. 127). Distribution area: 1,213 km²; status: SRE.

Ecology

Hoffman (1949) provides little ecological information, other than mentioning that the type specimen was found under hemlock bark. The authors JCM and DAH have visited the type locality, McGraw Gap, on multiple occasions and the habitat is composed of a mesic mixed hardwood and pine forest dominated by oak, maple, tuliptree, rhododendron, pine and hemlock, with an extensive fern understory.

Etymology

Hoffman (1949) gave no etymology for his choice of specific name but it is reasonable to assume that the name refers to the relatively simple form of the gonopods.

Type material

United States, Virginia, Alleghany County, McGraw Gap, 3 miles northwest of Clifton Forge; [37.8586°N, -79.8660°W].

Notes

In the original description, Hoffman (1949: 385) examined and designated the holotype male (NMNH #1807) collected on June 19, 1947 by R. Hoffman.

paupertas clade

Components. – Nannaria castra sp. nov., N. paupertas sp. nov., a female from Raleigh, North Carolina and a female from Carroll County, Virginia. Members of the paupertas clade share a medially curved prefemoral process and caudally directed acropodite tip. The division of N. castra sp. nov. and N. sheari sp. nov. (from Camp Creek Falls State Park and Brush Creek Preserve, respectively) into two separate clades was highly unexpected. Both specimens have medially curved prefemoral processes, with caudally directed acropodite tips, and the Camp Creek specimen does not share a sinuous region with N. paupertas sp. nov., a feature which we expected to unite N. paupertas sp. nov. with N. cingulata sp. nov. Furthermore, N. castra sp. nov. and N. sheari sp. nov. specimens were collected from localities only ~ 8 km apart. However, gonopods of N. paupertas sp. nov. and N. castra sp. nov. do share some morphological characteristics (see above) and therefore the similarities between the N. castra sp. nov. and N. sheari sp. nov. specimens are likely due to convergence. The development of a sinuous region in N. paupertas sp. nov. and a cingulum in N. cingulata sp. nov. is also likely attributable to convergence. The female from Raleigh, North Carolina may be N. conservata, as that is the only known species from that area. However, without a male, and in light of the complex nature of clade distributions within the minor group, it is possible that this female specimen represents an additional undescribed species. Collection of a male of this species, as well as the species from Carroll County, Virginia, will help resolve relationships within this clade.

Distribution. – the paupertas clade extends from central North Carolina, through southwestern Virginia and southern West Virginia (Fig. 114).

Nannaria castra sp. nov.

http://zoobank.org/A35BF7A1-2D72-4670-AE27-4276CD168523

Figs 20, 21 Vernacular name: “The Camp Creek Twisted-Claw Millipede”

Material examined

Holotype: United States – West Virginia • ♂; Mercer County, Camp Creek State Park; 37.5147°N, -81.1297°W; elev. 616 m; 12 Nov. 2017; hand collected; J. Means leg.; VTEC MPE03470.

Paratypes: United States – West Virginia • 2 ♀♀; same collection data as holotype; VTEC MPE03476, 3482 • 3 ♂♂; same collection data as holotype; VTEC MPE03475, 3477, 3478 • 3 ♂♂; same collection data as holotype; VMNH MPE03480, 81, 83 • 1 ♀; same collection data as holotype; VMNH MPE03479.

Other material

United States – West Virginia • 2 ♀♀; Mercer County, Camp Creek State Park, Farley Branch; 37.5082°N, -81.1370°W; elev. 657 m; 21 July 2005; hand collected; P. Marek, C. Spruill leg.; VTEC SPC000749, 750 • 1 ♂; Mercer County, Camp Creek State Park, Nash Fork Hollow; 37.5233°N, -81.1311°W; 36 Mar. 1968; W. Shear leg.; VMNH NAN0014. For detailed collection data see Suppl. material 7.

Diagnosis

Adult males of Nannaria castra sp. nov. are distinct from other Nannaria and the nearby N. aenigma Means, Hennen & Marek in Means et al. 2021, based on the following combination of characters: Gonopods. Gonopodal acropodite relatively straight before curving medially at half-way point, not continually curving as in N. paupertas sp. nov. Distal zone curving dorsally, with tip directed caudally (Fig. 20B)—not curving ventromedially with tip directed dorsomedially as in N. paupertas sp. nov., or tip directed medially as in N. aenigma. Acropodite tip with lateral flange (Fig. 20A, red arrow). Acropodite with slight twist and swelling at midpoint (Fig. 20C, red triangle), lacking medial flange of N. paupertas sp. nov., or lateral flange of N. aenigma. Telopodite basal zone thin, with slight lateral bulge, height subequal to 1/2 length of acropodite, not wide as in N. paupertas sp. nov., or > 1/6 length as in N. aenigma. Prefemur with dorsomedially curving prefemoral process (Fig. 20B), not ventromedially curving as in N. paupertas sp. nov., or straight as in N. aenigma. Prefemoral spine absent, not prominent, acicular as in N. paupertas sp. nov. Color. Tergites with orange paranotal spots (Fig. 21). Black background. Dorsum of collum smooth with orange margin.

Figure 20.

Nannaria castra sp. nov. holotype ♂ (VTEC, MPE03470) left gonopod A anterior view; red arrow indicates lateral flange B medial view C posterior view; red triangle indicates acropodite midpoint twist and swelling. Scale bar: 0.5 mm.

Figure 21.

Nannaria castra sp. nov. holotype ♂ (VTEC, MPE03470) coloration. Scale bar: 4.0 mm.

Measurements

♂ holotype (VTEC, MPE03470): BL = 28.5, CW = 3.6, IW = 1.8, ISW = 0.7, B11W = 4.2, B11H = 2.5. ♀ paratype (VTEC, MPE03476): BL = 27.8, CW = 3.6, IW = 2.0, ISW = 0.9, B11W = 4.5, B11H = 3.2.

Variation

No known variation.

Distribution

Known only from the type locality, Camp Creek State Park (West Virginia: Mercer County; Suppl. material 7; Fig. 126). Distribution area: > 1 km²; status: MRE.

Ecology

Individuals of N. castra sp. nov. were found in a mesic hardwood and rhododendron forest on top of hardpacked soil surrounding the entrance parking lot to Camp Creek State Park, and along the road leading towards the park campground. The majority of individuals were walking on top of the soil beneath leaf litter.

Etymology

This species is named for its type locality. The specific name a noun in apposition from the Latin castra for ‘camp.’

Type locality

United States, West Virginia, Mercer County, Camp Creek State Park, 37.5147°N, -81.1297°W.

Nannaria paupertas sp. nov.

http://zoobank.org/C8B741BA-38E7-4472-9364-3A0F4EB52DAB

Figs 22, 23 Vernacular name: “The Poverty Creek Twisted-Claw Millipede”

Material examined

Holotype: United States – Virginia • ♂; Montgomery County, Blacksburg, Pandapas Pond, Poverty Creek Trail near forest access rd. 708; 37.2678°N, -80.4852°W; elev. 648 m; 7 July 2014; hand collected; J. Means leg.; VTEC MPE00108.

Paratype: United States – Virginia • 1 ♀; same collection data as holotype; VMNH MPE00109.

Other material

United States – Virginia • 1 ♂; Montgomery County, Pandapas Pond, NW of Blacksburg, Poverty Creek Trail; 37.2805°N, -80.4720°W; elev. 651 m; hand collected; D. Hennen leg.; VTEC MPE02084 • 1 ♂; Montgomery County, gully on north slope of Brush Mtn. off forest service road 208; 37.2764°N, -80.4837°W; elev. 693 m; 9 May 2015; J. Means leg.; VTEC MPE00359 • 1 ♂; Montgomery County, Pandapas Pond, in bottomland below horse trail parking lot; 37.2824°N, -80.4485°W; elev. 738 m; hand collected; J. Means leg.; VTEC MPE00834 • 1 ♀; same collection data as preceding; VTEC MPE00836. For detailed collection data see Suppl. material 7.

Diagnosis

Adult males of Nannaria paupertas sp. nov. are distinct from other Nannaria and the sympatric N. ericacea, based on the following combination of characters: Gonopods. Gonopodal acropodite gently curving medially before bending ventromedially at a nearly 90° angle (Fig. 22B, red triangle), not straight as in N. ericacea or curving dorsomedially as in N. cingulata sp. nov. Distal zone short, bent dorsomedially at 90° angle, not laminate, curving medially with flanges as in N. ericacea or bent dorsomedially at 45° as in N. cingulata sp. nov. Acropodite tip directed dorsomedially, without flanges, not laminate and serpentine, directed medially as in N. ericacea. Telopodite basal zone height > 1/3 length of acropodite, not greatly reduced as in N. ericacea or with lateral bulge as in N. cingulata sp. nov. Acropodite shaft swollen before apex, with medial flange, and sinuous region, not with lateral flange as in N. ericacea or cingulum as in N. cingulata sp. nov. Prefemur with prefemoral process curving ventromedially, not ventrally as in N. cingulata sp. nov. and N. ericacea. Prefemoral process (when viewed medially) coplanar with acropodite, not crossing acropodite as in N. ericacea. Prefemoral process 2/3 length of acropodite, not subequal as in N. ericacea or 1/2 length as in N. cingulata sp. nov. Prefemoral spine prominent, acicular, not reduced as in N. cingulata sp. nov. or absent as in N. ericacea. Color. Tergites with orange paranotal spots (Fig. 23). Dark brown background. Dorsum of collum smooth with orange margin.

Figure 22.

Nannaria paupertas sp. nov. holotype ♂ (VTEC, MPE00108) left gonopod A anterior view B medial view; red triangle indicates acropodite ventromedial bend C posterior view. Scale bar: 0.5 mm.

Figure 23.

Nannaria paupertas sp. nov. holotype ♂ (VTEC, MPE00108) coloration. Scale bar: 4.0 mm.

Measurements

♂ holotype (VTEC, MPE00108): BL = 30.3, CW = 4.1, IW = 2.1, ISW = 0.9, B11W = 4.8, B11H = 3.1; ♀ paratype (VMNH, MPE00109): BL = 31.7, CW = 4.0, IW = 2.2, ISW = 1.0, B11W = 5.1, B11H = 3.6.

Variation

A dead male collected from Craig County is morphologically similar to N. paupertas sp. nov., with the exception of a prefemoral process which is straight for most of its length before bending nearly 90° ventrally, and a highly reduced prefemoral spine. Genetic sequences of a sympatric female did not support conspecificity with N. paupertas sp. nov. (Suppl. material 7), and therefore the Craig County population may represent a separate species and is not included in N. paupertas sp. nov.

Distribution

Known only from a small area around Pandapas Pond (an impoundment of Poverty Creek) in Montgomery County, Virginia (Virginia: Montgomery County, Suppl. material 7; Fig. 126). Distribution area: 2 km²; status: MRE.

Ecology

Individuals of N. paupertas sp. nov. have been collected from the side of walking trails in mesic, broadleaved forests, composed of oak, maple, ferns, and scattered pine, under deciduous leaf litter adjacent to rhododendron groves.

Etymology

This species is named for the Poverty Creek Trail by which it was first collected. The specific name is derived from the Latin paupertas, meaning poverty, and is a noun in apposition.

Type locality

United States, Virginia, Montgomery County, Blacksburg, Pandapas Pond, Poverty Creek Trail near forest access rd. 708, 37.2678°N, -80.4852°W.

Murphy clade

Components. female specimens from Mattaponi Wildlife Management Area, Virginia, the Appalachian Trail crossing of Virginia highway 621, and Dr. Brian Murphy’s property near Blacksburg, Virginia. This clade is unfortunately completely comprised of female specimens and therefore little in the way of conclusions can be drawn as to its characteristics. The specimens from Dr. Brian Murphy’s property and the AT crossing of Virginia highway 621 are likely representatives of the same undescribed species, while the specimen from Mattaponi Wildlife Management Area may represent a second undescribed species.

mcelroyorum clade

Components. Nannaria caverna sp. nov. and N. mcelroyorum sp. nov. The mcelroyorum clade is characterized by a simple, medially directed acropodite tip, and a prefemoral process arising dorsomedially from the prefemoral spine. We had expected N. mcelroyorum sp. nov. and N. caverna sp. nov. to fall into separate clades based on the differing prefemoral processes. However, the form of the prefemoral process varies between species in several clades recovered in the molecular phylogeny (Fig. 7), suggesting that the prefemoral process may not be as taxonomically informative as we had previously believed.

Distribution. – the mcelroyorum clade extends from southwestern West Virginia into northeastern Kentucky (Fig. 114).

Nannaria caverna sp. nov.

http://zoobank.org/41468A49-456A-41C0-B973-D66E8B757919

Figs 24, 25 Vernacular name: “The Carter Caves Twisted-Claw Millipede”

Material examined

Holotype: United States – Kentucky • ♂; Carter Co., Carter Caves State Park; 38.3738°N, -83.1142°W; 24 Sep. 2017; hand collected; J. Means, D. Hennen leg.; VTEC MPE03139.

Paratypes: United States – Kentucky • 1 ♂; same collection data as holotype; VTEC MPE03157 • 1 ♀; same collection data as holotype; VTEC MPE03158 • 1 ♂; same collection data as holotype; VMNH MPE03160 • 1 ♀; same collection data as holotye; VMNH MPE03159. For detailed collection data see Suppl. material 7.

Diagnosis

Adult males of Nannaria caverna sp. nov. are distinct from other Nannaria and nearby N. shenandoa, based on the following combination of characters: Gonopods. Gonopodal acropodite linear, bent medially forming 130° angle with telopodite basal zone bent medially at tip, not continually curving throughout as N. serpens sp. nov. and N. shenandoa. Tip simple, rectangular, without laminate flanges as in N. shenandoa. Telopodite basal zone tall, > 1/2 length of acropodite, not < 1/2 as in N. serpens sp. nov. and N. shenandoa. Prefemur with sinuous, ventrally directed prefemoral process, bent at a 90° angle, arising from large, pronounced prefemoral spine (Fig. 24B, red arrow), not directed cephalically as in N. serpens sp. nov., or curving laterally as in N. shenandoa. Color. Tergites with light orange paranotal spots (Fig. 25). Light grey background, likely due to teneral condition of specimens. Collum smooth with orange margin.

Figure 24.

Nannaria caverna sp. nov. holotype ♂ (VTEC, MPE03139) left gonopod A anterior view B medial view; red arrow indicates ventrally directed prefemoral process C posterior view. Scale bar: 0.5 mm.

Figure 25.

Nannaria caverna sp. nov. holotype ♂ (VTEC, MPE03139) coloration. Scale bar: 4.0 mm.

Measurements

♂ holotype (VTEC, MPE03139): BL = 24.7, CW = 3.8, IW = 2.5, ISW = 0.8, B11W = 4.2, B11H = 2.9. ♀ paratype (VMNH, MPE03159): BL = 25.7, CW = 3.2, IW = 2.2, ISW = 1.2, B11W = 4.0, B11H = 2.9.

Variation

No known variation.

Distribution

Known only from the type locality (Kentucky: Carter Caves State Park; Suppl. material 7; Fig. 127). Distribution area: N/A; status: MRE.

Ecology

Individuals of N. caverna sp. nov. were collected as they emerged from hardpacked soil on the edge of a creek running through Carter Caves State Park. The surrounding forest was made up of hardwood trees and rhododendron thickets.

Etymology

This species is named for its type locality. The specific name is a noun in apposition from the Latin caverna for cave.

Type locality

United States, Kentucky, Carter Co., Carter Caves State Park, 38.3738°N, -83.1142°W.

Nannaria mcelroyorum sp. nov.

http://zoobank.org/37EFC360-6A08-41DD-848F-2C5C97C60E70

Figs 26, 27 Vernacular name: “The McElroy Twisted-Claw Millipede”

Material examined

Holotype: United States – West Virginia • ♂; Wayne County, Barboursville, Beech Fork State Park, Lost Trail near Moxley Branch Campground; 38.3047°N, -82.3512°W; elev. 194 m; 23 Sep. 2017; hand collected; J. Means and D. Hennen leg.; VTEC MPE03113.

Paratypes: United States – West Virginia • 3 ♂♂; same collection data as holotype; VTEC MPE03116, 3118, 3120 • 4 ♂♂; same collection data as holotype; VMNH MPE03135, 3151, 3154, 3156 • 3 ♀♀; same collection data as holotype; VTEC MPE03117, 3119, 3121 • 4 ♀♀; same collection data as holotype; VMNH MPE03122, 3136, 3137, 3155.

Other material

United States – Kentucky • 1 ♂; Boyd County, Fannin Park, off Bramble Drive, hillside across road from park; 38.3475°N, -82.6866°W; elev. 219 m; 24 Sep. 2017; J. Means, D. Hennen leg.; VTEC MPE03125; SCAU – West Virginia • 3 ♂♂; Boone County, Julian, along Big Pinnacle Road; 38.1802°N, -81.8384°W; elev. 223 m; 23 Sep. 2017; hand collected; J. Means, D. Hennen leg.; VTEC MPE03262, 63, 3289 • 2 ♀♀; same collection data as preceding; VTEC MPE03290, 91 • 7 ♂♂; same collection data as preceding; VTEC MPE02240, 2257, 2258, 2260–62, 3727 • same collection data as preceding; 5 ♀♀; same collection data as preceding;18 Nov. 2016; VTEC MPE02248–50, 3728, 3729. For detailed collection data see Suppl. material 7.

Diagnosis

Adult males of Nannaria mcelroyorum sp. nov. are distinct from other Nannaria and the nearby N. shenandoa, based on the following combination of characters: Gonopods. Gonopodal acropodite gently curving medially throughout, not straight as in N. shenandoa. Acropodite gradually tapering towards tip, with small lobed medial flange (Fig. 26A, red triangle), lacking swollen medial area as in N. serpens sp. nov. Tip directed medially, rounded and simple, not with folds, grooves, flanges as in N. shenandoa. Distal zone greatly reduced, not enlarged and curving caudolaterally as in N. shenandoa. Prefemur with thin, sinuous, acicular prefemoral process, not curved, saber-like as in N. shenandoa. Prefemoral spine small, medially directed, with secondary hump proximal to acropodite base (Fig. 26C, red arrow), not enlarged, cephalically directed as in N. serpens sp. nov., or lacking as in N. shenandoa. Telopodite basal zone height ca. 1/2 length of acropodite, not > 1/2 length of acropodite as in N. serpens sp. nov. or ca. 1/5 length as in N. shenandoa. Color. Tergites with light orange paranotal spots and faint orange stripes (Fig. 27). Light brown background. Dorsum of collum smooth with orange margin.

Figure 26.

Nannaria mcelroyorum sp. nov. holotype ♂ (VTEC, MPE03113) left gonopod A anterior view; red triangle indicates medial flange B medial view C posterior view; red arrow indicates prefemoral spine secondary hump. Scale bar: 0.5 mm.

Figure 27.

Nannaria mcelroyorum sp. nov. non-type ♂ (VTEC, MPE03125) coloration. Scale bar: 4.0 mm.

Measurements

♂ paratype (VTEC, MPE03116; holotype too damaged for measurement): BL = 33.9, CW = 4.7, IW = 2.3, ISW = 0.9, B11W = 5.4, B11H = 3.6; ♀ paratype (VTEC, MPE03117): BL = 33.2, CW = 4.4, IW = 2.5, ISW = 1.3, B11W = 5.9, B11H = 4.3.

Variation

Individuals of N. mcelroyorum sp. nov. from the Boone County, West Virginia site have prefemoral processes ca. 2/3 the length of those of the holotype.

Distribution

Known from a linear area in western West Virginia and Eastern Kentucky (Kentucky: Boyd County; West Virginia: Boone and Wayne counties, Suppl. material 7; Fig. 127). Distribution area: 105 km²; status: MRE.

Ecology

Individuals of N. mcelroyorum sp. nov. were collected from mesic deciduous forests dominated by oak, maple, laurel, and some pine, often under 1–2 cm soil.

Etymology

This species is named for Clint, Justin, Travis, and Griffin McElroy from Huntington, West Virginia, who provided endless hours of emotional support to JCM and DAH during field collections through their podcasts, ‘The Adventure Zone’ and ‘My Brother, My Brother, and Me’. The specific name is a plural genitive derived as a patronym.

Type locality

United States, West Virginia, Wayne County, Barboursville, Beech Fork State Park, Lost Trail near Moxley Branch Campground, 38.3047°N, -82.3512°W.

hardeni clade

Components. – Nannaria hardeni sp. nov. Based on the molecular phylogeny (Fig. 7) and the combination of morphological characters detailed in the below diagnosis, N. hardeni sp. nov. has no sister taxon and is in a monotypic clade (Fig. 113). Due to shared morphological characters between N. hardeni sp. nov. and N. monsdomia sp. nov., including a large, rounded prefemoral spine, the proximity of the prefemoral process to the acropodite and the wide separation between the prefemoral spine and process, we had expected the two species to be closely related. Instead, N. hardeni sp. nov. is sister to the serpens and castanea clades, and appears to be yet another example of morphological convergence confusing attempts to infer evolutionary relatedness through gonopod morphology alone.

Distribution. N. hardeni sp. nov. is known only from southern Virginia (Fig. 128).

Nannaria hardeni sp. nov.

http://zoobank.org/A276BC4F-2CB8-4822-9A23-5A368B6CE830

Figs 28, 29 Vernacular name: “Curt Harden’s Twisted-Claw Millipede”

Material examined

Holotype: United States – Virginia • ♂; Pittsylvania County, Angler’s Park; 36.5577°N, -79.3515°W; elev. 134 m; 23 Dec. 2016; C. W. Harden leg.; VMNH MPE02278.

Paratypes: United States – Virginia • 1 ♂; same collection data as holotype; VTEC MPE02282 • 1 ♂; same collection data as holotype; VMNH MPE02283 • 1 ♀; same collection data as holotype; VTEC MPE02279 • 1 ♀; same collection data as holotype; VMNH MPE02281.

Other material

United States – Virginia • 1 ♂; Henry County, vicinity of Figsboro; 36.7861°N, -79.8586°W; 9 Nov. 1980; R. Hoffman leg.; VMNH NAN0247 • 1 ♂; same collection data as preceding; 15 Nov. 1981; VMNH NAN0249 • 1 ♂; Henry County, Martinsville, VMNH; 36.6761°N, -79.8771°W; 2 Nov. 1992; J. Anderson, VMNH surveys leg.; VMNH NAN0270 • 1 ♂; same collection data as preceding; Dec. 1997; VMNH NAN0271 • 1 ♂; same collection data as preceding; 19 Dec. 1996; VMNH survey leg.; VMNH NAN0273 • 1 ♂; Martinsville, VMNH on sidewalk to Annex; 36.6761°N, -79.8771°W; 12 Nov. 1992; J. Anderson leg.; VMNH NAN0272 • 1 ♂; Martinsville, Dundee Lane; 36.6892°N, -79.8727°W; 5 Jan. 1990; P. Carter leg.; VMNH NAN0274 • 1 ♂; near Martinsville, Dupont prop.; 36.6680°N, -79.8922°W; 16 Oct. 1995; J. Anderson leg.; VMNH NAN0275 • 2 ♂♂; same collection data as preceding; 23 Oct. 1995; VMNH NAN0276 • 8 ♂♂; Pittsylvania County, Danville, Angler’s Park; 36.5500°N, -79.3500°W; elev. 126 m; 29 Oct. 2017; hand collected; C. Harden leg.; VTEC MPE03734–40, 4239 • 3 ♀♀; same collection data as preceding; VTEC MPE03741–3 • 18 ♂♂; Pittsylvania County, Lacy Farm DF site, 3 mi. ENE of Axton; 36.6760°N, -79.6621°W; 29 Mar. 1992; VMNH survey leg.; VMNH NAN0245 • 8 ♂♂; same collection data as preceding; 21 Dec. 1992; VMNH NAN0246 • 10 ♂♂; same collection data as preceding; 23 Apr. 1992; VMNH NAN0248 • 2 ♂♂; same collection data as preceding; 15 May 1992; VMNH NAN0250 • 8 ♂♂; Lacy Farm, ca 5 mi. ENE of Axton, pitfall trap; 36.6870°N, -79.6286°W; 13 May 1992; VMNH survey leg.; VMNH NAN0251; 28 ♂♂; Lacy Farm DF site, ca 4 mi. ENE of Axton; 36.6815°N, -79.6454°W; 13 Nov. 1992; VMNH survey leg.; VMNH NAN0252. For detailed collection data see Suppl. material 7.

Diagnosis

Adult males of Nannaria hardeni sp. nov. are distinct from other Nannaria and the nearby N. wilsoni, based on the following combination of characters: Gonopods. Gonopodal acropodite (Fig. 28) gently curving medially before apex, distal zone reduced, not laminate, curving caudally with flanges as in N. wilsoni or with lobed lateral flange as in N. monsdomia sp. nov. Telopodite basal zone height > 1/2 length of acropodite, not < 1/4 length of acropodite as in N. wilsoni. Tip simple, directed medially, without lateral flange as in N. monsdomia sp. nov., and not directed caudally with lateral flange as in N. wilsoni. Prefemur with straight, acicular prefemoral process, not laminate, curving medially as in N. monsdomia sp. nov. or N. wilsoni. Prefemoral process not crossing acropodite as in N. wilsoni or N. monsdomia sp. nov. Prefemoral process length subequal to height of telopodite basal zone, not greater than height of telopodite basal zone as in N. monsdomia sp. nov. and N. wilsoni. Prefemoral spine widely separated from prefemoral process (Fig. 28A, red arrow) and sharp, tooth-like, not rounded as in N. monsdomia sp. nov. or lacking as in N. wilsoni. Color. Tergites with orange paranotal spots (Fig. 29). Dark brown background. Dorsum of collum smooth with orange margin.

Figure 28.

Nannaria hardeni sp. nov. holotype ♂ (VMNH, MPE02278) left gonopod A anterior view; red arrow indicates wide separation between prefemoral process and prefemoral spine B medial view C posterior view. Scale bar: 0.5 mm.

Figure 29.

Nannaria hardeni sp. nov. holotype ♂ (VMNH, MPE02278) coloration. Scale bar: 4.0 mm.

Measurements

♂ holotype (VMNH, MPE02278): BL = 17.3, CW = 2.5, IW = 1.6, ISW = 0.6, B11W = 2.9, B11H = 1.9; ♀ paratype (VMNH, MPE02281): BL = 17.6, CW = 2.1, IW = 1.8, ISW = 0.6, B11W = 2.9, B11H = 1.9.

Variation

No known variation.

Distribution

Known from southern Virginia, near the North Carolina border (Virginia: Henry and Pittsylvania counties; Suppl. material 7; Fig. 128). Distribution area: 374 km²; status: MRE.

Ecology

Individuals of N. hardeni sp. nov. have been found in young, disturbed mesic hardwood forests under rocks, often in fern thickets. Interestingly, N. hardeni sp. nov. has also been collected from IPM sticky traps from inside the Virginia Museum of Natural History in Martinsville, Virginia.

Etymology

This species is named after its collector, Curt W. Harden. The specific name is a genitive noun derived as a patronym.

Type locality

United States, Virginia, Pittsylvania County, Angler’s Park, 36.5577°N, -79.3515°W.

serpens clade

Components. – Nannaria serpens sp. nov. Based on the molecular phylogeny (Fig. 7) and the morphological characters detailed in the below diagnosis, including the presence of a serpentine prefemoral process and enlarged prefemoral spine, coupled with a slight medial swelling before the acropodite apex, N. serpens sp. nov. is in a monotypic clade (Fig. 113).

Distribution. N. serpens sp. nov. is known solely from southwestern Virginia and northwestern North Carolina (Fig. 126).

Nannaria serpens sp. nov.

http://zoobank.org/33C19D59-CF22-4713-A660-D1A2FC2DE146

Figs 30, 31 Vernacular name: “The Serpentine Twisted-Claw Millipede”

Material examined

Holotype: United States – Virginia • ♂; Carroll County, Dugspur, side of hill southeast of Blacksnake Meadery; 36.7747°N, -80.5420°W; elev. 822 m; 17 Sep. 2014; hand collected; J. Means leg.; VTEC MPE00202.

Paratypes: United States – Virginia • 1 ♂; same collection data as holotype; VMNH MPE00210.

Other material

United States – North Carolina • 1 ♂; Ashe County, Three-Top Mtn., 2 miles SE of Creston; 36.4144°N, -81.6003°W; elev. 1219 m; 23 July 1963; R. Hoffman, Carico leg.; VMNH NAN0206 • 2 ♂♂; Avery County, Grandfather Mtn.; 36.1096°N, -81.8114°W; C. M. leg.; VMNH NAN0205; SCAU – Virginia • 1 ♂; Carroll County, Blacksnake Meadery, bottomland near creek at entrance; 36.7760°N, -80.5446°W; elev. 767 m; 14 Sep. 2014; hand collected; J. Means leg.; VTEC MPE00274 • 1 ♀; same collection data as preceding; VTEC MPE00204 • 1 ♂; Floyd County, on hill edge by side of road across from Rocky Knob Picnic Area; 36.8132°N, -80.3495°W; elev. 970 m; 18 Sep. 2015; hand collected; J. Means, P. Marek, K. Lawler, P. Shorter, V. Wong leg.; VTEC MPE00817 • 1 ♀; same collection data as preceding; VTEC MPE03717 • 6 ♂♂; Floyd County, 6 mi SE Willis, Felker’s property, Rt. 726, Sorex study; 36.8009°N, -80.3988°W; 7 June 1995; J. Anderson leg.; VMNH NAN0083 • 3 ♂♂; same collection data as preceding; 20 June 1995; VMNH survey leg.; VMNH NAN0087 • 2 ♂♂; same collection data as preceding; 7 June 1995; J. Anderson leg.; VMNH, NAN0089 • 12 ♂♂; same collection data as preceding; 4 June 1993; VMNH NAN0207 • 4 ♂♂; Floyd County, Buffalo Mountain N.A.P. south slope site, berleseate; 36.7958°N, -80.4772°W; elev. 1067 m; 9 Aug. – 6 Sep. 2000; berlese; Joint Survey leg.; VMNH NAN0084 • 1 ♂; same collection data as preceding; 23 Oct. 1996; VMNH survey leg.; VMNH NAN0088 • 1 ♂; same collection data as preceding; 18 Aug.1991; VMNH NAN0090 • 6 ♂♂; same collection data as preceding; 6 Sep.– 3 Oct. 2000; VMNH NAN0091 • 1 ♂♂; same collection data as preceding; 12 Jan. 1986; R. Hoffman leg.; VMNH NAN0092 • 6 ♂♂; same collection data as preceding; 10 Oct. – 8 Nov.; VMNH survey leg.; VMNH NAN0093 • 30 ♂♂; same collection data as preceding; 23 June – 15 July 2001; VMNH NAN0094, 95 • 9 ♂♂; same collection data as preceding; 23 Nov. 2000; Joint Survey; VMNH NAN0096 • 24 ♂♂; same collection data as preceding; 23 Aug. – 29 Sep. 2001; VMNH survey leg.; VMNH NAN0097 • 15 ♂♂; same collection data as preceding; 31 Oct. 2001; VMNH NAN0098 • 5 ♂♂; same collection data as preceding; 19 Aug. 1992; VMNH NAN0208 • 3 ♂♂; same collection data as preceding; 1 Apr. 1993; VMNH NAN0209 • 1 ♂; same collection data as preceding; 3 Sep. 1967; R. Hoffman leg.; VMNH NAN0296 • 1 ♂; Lee County, Powell Mountain at Va. Hy. 70, near crest; 36.6211°N, -83.1062°W; 5 June 1986; R. Hoffman leg.; VMNH NAN0153 • 2 ♂♂; Patrick County, Rock Castle Creek, Rock Castle Gorge Trail, Rocky Knob Recreation Center; 36.7858°N, -80.3717°W; elev. 839 m; 21 Sep. 2015; hand collected; J. Means, P. Marek leg.; VTEC MPE00830, 833 • 1 ♂; Patrick County, side of hill by Fred Clifton Park; 36.7181°N, -80.3246°W; elev; 869 m; hand collected; J. Means, P. Shorter; leg.; VTEC MPE02610 • 2 ♀; same collection data as preceding; VTEC MPE02629, 2630 • 3 ♂♂; Rockbridge County, vicinity of Natural Bridge, in dry limestone woods; 37.63°N, -79.5433°W; 14 May 1989; R. Hoffman leg.; VMNH NAN0193. For detailed collection data see Suppl. material 7.

Diagnosis

Adult males of Nannaria serpens sp. nov. are distinct from other Nannaria and the sympatric N. wilsoni, based on the following combination of characters: Gonopods. Gonopodal acropodite gently curving medially, not strongly curving medially as in N. wilsoni. Telopodite basal zone height < 1/2 length of acropodite, not heavily reduced as in N. wilsoni. Acropodite with slightly swollen medial area (Fig. 30C, red triangle) and small lobed medial flange (Fig. 30A, red arrow) basal to apex. Tip directed medially, blunt and simple, not with folds, grooves, flanges as in N. wilsoni. Distal zone greatly reduced, not enlarged and curving caudomedially as in N. wilsoni. Prefemur with sinuous, cephalically-directed prefemoral process, not curving medially as in N. wilsoni. Prefemoral spine enlarged, cephalically directed, lacking secondary hump as in N. mcelroyorum sp. nov. Color. Tergites with bright orange paranotal spots (Fig. 31). Dark brown background. Dorsum of collum smooth with orange margin.

Figure 30.

Nannaria serpens sp. nov. holotype ♂ (VTEC, MPE00202) left gonopod A anterior view; red arrow indicates medial flange B medial view C posterior view; red triangle indicates slight medial acropodite swelling. Scale bar: 0.5 mm.

Figure 31.

Nannaria serpens sp. nov. holotype ♂ (VTEC, MPE00202) coloration. Scale bar: 4.0 mm.

Measurements

♂ holotype (VTEC, MPE00202): BL = 30.9, IW = 2.4, ISW = 0.8, B11W = 4.7, B11H = 2.9; ♀ paratype (VTEC, MPE00204): BL = 29.4, CW = 3.4, IW = 1.8, ISW = 0.8, B11W = 4.4, B11H = 3.0.

Variation

No known variation.

Distribution

Known from southwestern Virginia and northwestern North Carolina, with disjunct populations ca. 112 km northeast at Natural Bridge State Park, and ca. 230 km southwest at Powell Mountain (North Carolina: Ashe and Avery counties; Virginia: Carroll, Floyd, Lee, Patrick and Rockbridge counties, Suppl. material 7; Fig. 126). Distribution area: 2,280 km²; status: SRE.

Ecology

Individuals of N. serpens sp. nov. have been collected from mesic hardwood forests composed of oak, maple, tuliptree, sassafras, and rhododendron, often found under leaves on wooded hillsides.

Etymology

This species is named for both the serpentine nature of its prefemoral process, and for its type locality, Blacksnake Meadery. The specific name is derived from the Latin serpens meaning snake, and is a noun in apposition.

Type locality

United States, Virginia, Carroll County, Dugspur, side of hill southeast of Blacksnake Meadery, 36.7747°N, -80.5420°W.

castanea clade

Components. Nannaria castanea (McNeill, 1887), N. davidcauseyi Causey, 1950, N. hokie, N. missouriensis Chamberlin, 1928, N. stellapolis sp. nov., N. stellaradix sp. nov. and a female from Poor Mountain, Virginia. Members of the castanea clade have a distinctly long, curving acropodite with a slight medial swelling before the acropodite apex and a medial flange at the tip which carries the prostatic groove, though this flange is much reduced in N. stellapolis sp. nov. and N. stellaradix sp. nov. No specimens of N. davidcauseyi were collected for this study, however due to close geographic proximity and the combination of shared gonopodal characters, including those typical for the clade mentioned above, we suggest that N. davidcauseyi belongs in the castanea clade.

Distribution. The castanea clade falls out into two distinct groups, one at the western extents of the Nannariini range and the other in southwestern Virginia. The castanea clade therefore has one of the largest distributions of all Nannaria clades, extending from southwestern Virginia, to central and northern Mississippi, and north into Arkansas, Indiana, and Missouri (Fig. 114). We expect that additional sampling of areas between these two geographic groups, such as Claytor Lake State Park, Natchez Trace State Park and throughout Tennessee, as well as male specimens from Poor Mountain, Virginia, will reveal additional forms of the castanea clade, and help explain its extensive geographic distribution.

Nannaria castanea (McNeill, 1887)

Figs 32, 33 Vernacular name: “The Bloomington Twisted-Claw Millipede”

Polydesmus castaneus McNeill, 1887: 329, fig. 8

Fontaria castanea: Bollman 1893: 123.

Mimuloria castanea: Chamberlin 1928: 155. Causey 1952: 8, fig. 6c; 1955: 30. Chamberlin and Hoffman 1958: 37. Hennen and Shelley 2015: 6, figs 3, 9–14.

Nannaria castanea: Chamberlin 1949: 4. Hoffman 1999: 365–366. Marek et al. 2014: 36. Means et al. 2021:17, S68.

Castanaria castanea: Causey 1950a: 1.

Castanaria depalmai: Causey 1950a: 1–3, fig. 1.

Mimuloria depalmai: Causey 1952: 9. Chamberlin and Hoffman 1958: 37–38.

Nannaria depalmai: Hoffman 1999: 366. Marek et al. 2014: 36.

Material examined

Lectotype (here designated): United States – Indiana • ♂; Monroe County, Bloomington; NMNH Type #38.

Paralectotype: United States – Indiana • 1 ♀; same collection data as holotype; NMNH Type #38.

Other material

United States – Arkansas • 11 ♀♀; Carroll County, Eureka Springs, Lake Leatherwood, Hyde Trail south of lake; 36.4307°N, -93.7576°W; elev. 317 m; 17 May 2017; hand collected; J. Means, D. Hennen, V. Wong leg.; VTEC MPE02765–70, MPE02797–99, MPE03766, MPE03767 • 1 ♂; Fulton County, Hwy. 62; 24 Apr. 1952; N. Causey leg.; VMNH NAN0196; SCAU – Mississippi • 1 ♂; Choctaw County, 4.5 mi. S. of Ackerman, upland woods; 33.2809°N, -89.1690°W; 2 Dec. 1961; L. Hubricht leg.; VMNH NAN0238 • 3 ♀♀; Tishomingo County, Burnsville, Divide Wildelife Management Area, forest beside pulloff on road; 34.8051°N, -88.3063°W; elev. 159 m; 14 May 2017; hand collected; J. Means, D. Hennen, V. Wong leg.; VTEC MPE02789–91 • 1 ♂; Tishomingo County, wooded hillside 1.6 miles W of Burnsville; 34.8388°N, -88.3298°W; 27 Feb. 1961; L. Hubricht leg.; VMNH NAN0241; SCAU – Missouri • 1 ♂; Crawford County, 5 miles W of Berryman; 37.9188°N, -91.1871°W; 2 Apr. 1955; R. Crabill leg.; VMNH NAN0242. For detailed collection data see Suppl. material 7.

Diagnosis

Adult males of Nannaria castanea are distinct from other Nannaria based on the following combination of characters: Gonopods. Gonopodal acropodite gently curving medially, not curving ventromedially before apex as in N. hokie. Distal zone with medial flange at 135° to solenomere, and lateral flange at 90° angle to solenomere (Fig. 32B, red triangle), not encircling tip as in N. hokie. Acropodite tip curving caudally. Acropodite robust, simple; not thin, with small medial flange near apex as in N. hokie. Telopodite basal zone height ca. 1/2 length of acropodite, not ca. 1/3 length as in N. hokie. Prefemoral process small, thin, curving medially, arising from top of projected, stout prefemoral spine (Fig. 32A, red arrow), not arising from thin, projected prefemoral spine as in N. hokie. Color. Tergites with orange paranotal spots (Fig. 33) and occasionally faint orange stripes (Fig. 33). Dark to light brown background. Dorsum of collum smooth with orange margin.

Figure 32.

Nannaria castanea lectotype ♂ (NMNH, Type #38) left gonopod A anterior view; red arrow indicates projected, stout prefemoral spine B medial view; red triangle indicates distal zone medial and lateral flanges C posterior view. Scale bar: 0.5 mm.

Figure 33.

Nannaria castanea (McNeill, 1887) ♀ (VTEC, MPE02789) coloration. Scale bar: 4.0 mm.

Measurements

Lectotype ♂ (NMNH, Type #38): CW = 3.5, IW = 2.2, ISW = 0.8. BL, B11W and B11H were unmeasurable due to the poor quality of the specimen.

Variation

Castanaria depalmai was synonymized with N. castanea by Hennen and Shelley (2015) without justification and the relationship between C. depalmai and N. castanea has not been tested using molecular evidence. Morphological differences between the two are extremely slight, however, so it is likely that the synonymy of C. depalmai and N. castanea will be supported by molecular phylogenetics. Interestingly, there are six specimens of N. castanea collected from two localities in Mississippi by Leslie Hubricht in 1961 that Hennen and Shelley (2015) were apparently unaware of and did not mention in their treatment of Mimuloria. The northern Mississippi specimen is remarkably similar in form to N. missouriensis, however specimens collected by the authors are genetically more similar to N. castanea from Arkansas than N. missouriensis, and we therefore identify the two Mississippi populations as N. castanea, awaiting more detailed investigations.

Distribution

Known from central Indiana west to Missouri and south into Arkansas and Mississippi (Indiana: Monroe County; Missouri: Jefferson, Crawford, Dent, Wayne, Wright counties; Arkansas: Carroll, Searcy, Stone, Fulton counties; Mississippi: Tishomingo, Choctaw counties; Suppl. material 7; Fig. 129). Distribution area: 113,102 km²; status: WRE.

Ecology

Individuals of N. castanea were collected from both mesic broadleaved and xeric semi-evergreen forests, often beneath 1–2 cm of soil.

Etymology

McNeill (1887) gave no etymology for his choice of castaneus but he mentioned the dark chestnut coloration of the species, suggesting that his choice of name may have been derived from the Latin castaneus for ‘of the color of chestnuts.’

Type locality

United States, Indiana, Monroe County, Bloomington.

Notes

In the original description, McNeill (1887: 329) stated that he examined three specimens but did not designate a holotype, and no lectotype has been designated by subsequent authors. Therefore, we consider two type specimens we examined as syntypes and here we designate a lectotype male. There is no information on who collected the type material and when. The label of “NMNH Type #38” was presumeably added by an unknown individual after the specimens were deposited at the NMNH.

Nannaria davidcauseyi Causey, 1950

Fig. 34 Vernacular name: “David Causey’s Twisted-Claw Millipede”

Nannaria davidcauseyi Causey, 1950b: 194, figs 3, 4. Hoffman 1999: 366. Marek et al. 2014: 36. Means et al. 2021: S69.

Mimuloria davidcauseyi: Chamberlin and Hoffman 1958: 37. Hennen and Shelley 2015: 10, figs 4–8, 16, 17.

Material examined

United States – Arkansas • 1 ♀; Newton County, Dismal Hollow; 35.8506°N, -93.2701°W; 2 Nov. 2013; pitfall trap; M. Skvarla leg.; NCSM NAN0539 • 2 ♂♂; Newton County, Steel Creek, Buffalo Nt River; 36.0380°N, -93.3402°W; 7 Nov. 2013; pitfall trap; M. Skvarla leg.; NCSM NAN0537. For detailed collection data see Suppl. material 7.

Diagnosis

Adult males of N. davidcauseyi are distinct from other Nannaria and the nearby N. castanea based on the following combination of characters: Gonopods. Gonopodal acropodite gently curving medially, and distal zone with medial and lateral flanges at 90° angle to solenomere, not with medial flange at 135° angle to solenomere as in N. castanea (Fig. 34A, red arrow). Acropodite with small medial flange near apex, not simple lacking flange as in N. castanea (Fig. 34A, red triangle). Telopodite basal zone height 1/2 length of acropodite. Prefemoral process pronounced, acicular, arising distantly from small blunt prefemoral spine, not from top of projected prefemoral spine as in N. castanea. Color. Hennen and Shelley (2015) described specimens of N. davidcauseyi as having clear paranota with irregular orange spots internally, and orange/pink collum margins.

Figure 34.

Nannaria davidcauseyi Causey, 1950 ♂ (VMNH, NAN0537) left gonopod A anterior view; red triangle indicates small acropodite medial flange, red arrow indicates medial flange at apex B medial view C posterior view. Scale bar: 0.5 mm.

Measurements

♂ (VMNH, NAN0537): BL = 23.3, CW = 3.0, IW = 1.5, ISW = 0.6, B11W = 3.8, B11H = 2.2.

Variation

Causey (1950b: 194) mentions that the male paratype of N. davidcauseyi has variation in the shape of both the prefemoral spine and the “keel at the end of the main blade” referring to the caudally-directed acropodite tip.

Distribution

Known only from the northwest corner of Arkansas (Arkansas: Johnson and Newton counties, Fig. 129). Distribution area: 131 km²; status: MRE.

Ecology

Causey (1950b) describes the habitat in which she and her husband found N. davidcauseyi as an eastern-facing hillside in an oak-hickory woodland. Two individuals of N. davidcauseyi were collected from Steel Creek Campground in Arkansas by M. Skvarla (NAN0537) who listed oak, hickory, American beech, and eastern red cedar as the primary tree species in the area (Skvarla et al. 2015).

Etymology

Causey (1950b) gave no explanation of her choice in name for this species, however it is presumed that this species is named after the author’s husband and its collector, Dr. David Causey.

Type locality

United States, Arkansas, Newton County, about three miles northwest of Jasper.

Notes

In the original description Causey (1950b: 194) designates a male holotype and one male and two immature paratypes collected by Dr. David Causey on 25 August 1950. Hennen and Shelley (2015: 10) refer to a female paratype and one immature paratype, which would suggest that Causey’s original identification of the female specimen as an immature was incorrect.

Nannaria hokie Means, Hennen & Marek, 2021

Figs 35, 36, 37 Vernacular name: “The Virginia Tech Twisted-Claw Millipede”

Nannaria hokie Means, Hennen & Marek, in Means et al. 2021: 4, 5, 7, 10, 17–19, 24, S70.

Material examined

Holotype: United States – Virginia • ♂; Montgomery County, Blacksburg, Virginia Tech campus, Stadium Woods; 37.2216°N, -80.4159°W; elev. 637 m; 17 Apr. 2019; hand collected; P. Marek leg.; VTEC MPE04803.

Paratypes: United States – Virginia • 5 ♂♂; Montgomery County, Blacksburg, Virginia Tech campus, south side of Duck Pond; 37.2250°N, -80.4276°W; elev. 625 m; 13 Oct. 2015; hand collected; D. Hennen leg.; VTEC MPE00880, 882, 883, VMNH, MPE00884, 886 • 2 ♀♀; same collection data as preceding; VTEC MPE00881, 885.

Other material

United States – Virginia • 4 ♂♂; Floyd County, along Goose Creek, ca. 3 mi. N of Simpsons, 1850 ft.; 37.0815°N, -80.2055°W; 9 Oct. 1971; R. Hoffman, Knight leg.; VMNH NAN0005, 0244 • 10 ♂♂; Montgomery County, Golden Hills Disc Golf course, in gully; 37.1729°N, -80.4078°W; elev. 628 m; 8 Nov. 2014; hand collected; J. Means, D. Hennen, P. Marek leg.; VTEC MPE00266, 275, 280, 283–89 • 5 ♂♂; Montgomery County, Blacksburg, Virginia Tech, patch of woods between Grove Ln. and Duck Pond; 37.2247°N, -80.4260°W, elev. 628 m; 9 Nov. 2017; hand collected; J. Means, D. Hennen, G. Schiermeyer leg.; VTEC MPE03464–68 • 1 ♂; Montgomery County, Blacksburg, end of Valley View Drive near quarry; 37.2218°N, -80.3894°W; elev. 671 m; 13 Sep. 2019; hand collected; J. Means, D. Hennen, P. Marek, F. Vasquez, I. Huerta leg.; VTEC MPE04998 • 2 ♂♂; Montgomery County, Gateway Trail, 0.5 mi in from main trail by stream; 37.2506°N, -80.4610°W; elev. 642 m; 26 Sep. 2014; hand collected; J. Means, P. Marek, E. Francis, K. Lawler, N. Zegler leg.; VTEC MPE00227, 229 • 4 ♂♂; Montgomery County, Blacksburg, 803 Airport Rd.; 37.2125°N, -80.4053°W; elev. 581 m; 3 Mar. 2018; hand collected; J. Means, P. Marek, G. Schiermeyer, C. Hall leg.; VTEC MPE03829, 3830, 3833, 3835 • 4 ♀♀; same collection data as preceding; VTEC MPE03831, 3832, 3834, 3836 • 1 ♂; Montgomery County, Blacksburg, 505 Fairview Ave., ex puddle of water at base of driveway in neighborhood; 37.212°N, -80.40663°W; elev. 640 m; 13 Mar. 2020; hand collected; P. Marek leg.; VTEC MPE05027 • 1 ♂; Montgomery County, Blacksburg, jct. Airport and Fairview Rds.; 37.2122°N, -80.4060°W; elev. 634 m; 19 Feb. 2018; hand collected; P. Marek leg.; VTEC MPE03808 • 1 ♀; Montgomery County, Blacksburg, Virginia Tech campus, southern end of Duck Pond; 37.2251°N, -80.4272°W; elev. 623 m; 8 June 2017; G. Scheirmeyer leg.; VTEC MPE03705 • 1 ♂; Montgomery County, Blacksburg; 37.1849°N, -80.4087°W; 28 Oct. 1956; R. Hoffman leg.; VMNH NAN0234 • 20 ♂♂; Montgomery County, south Blacksburg, nr jct. Rt. 775 and US 460, under stones in grassy field; 37.1849°N, -80.4087°W; 3 Nov. 1956; R. Hoffman leg.; VMNH NAN0320, 321 • 1 ♂; Montgomery County; Oct. 1950; R. Hoffman leg.; VMNH NAN0235 • 5 ♂♂; Montgomery County, Blacksburg, Trillium Vale; 37.2294°N, -80.4141°W; 10 Oct. 1950; R. Hoffman leg.; VMNH NAN0243 • 6 ♂♂; Pulaski County, Highland Farm, bottom of hill in dry stream bed; 37.1775°N, -80.6456°W; elev. 574 m; 27 Dec. 2014; hand collected; J. Means, K. Lawler leg.; VTEC MPE00253–55, 259–61 • 3 ♀♀; same collection data as preceding; MPE00262–64 • 1 ♂; Smyth County, 7 mi SE of Chilhowie, NW slope of Iron Mtn.; 36.7100°N, -81.6188°W; 4 May 1964; R. Hoffman leg.; VMNH NAN0003. For detailed collection data see Suppl. material 7.

Diagnosis

Adult males of Nannaria hokie are distinct from other Nannaria and the sympatric N. ericacea, based on the following combination of characters: Gonopods. Gonopodal acropodite long and curving ventromedially before apex, not medially as in N. castanea or straight as in N. ericacea. Distal zone bent ventroposteriorly, with laminate flange encircling tip forming a hood-like structure around dorso-posteriorly projected. Acropodite solenomere laminate, partially obscuring solenomere when viewed laterally (Fig. 35A, red arrow). Distal zone lacking flat laminate flange, at 90° angle to solenomere as in N. castanea or laminate, sinuous, directed ventro-cephalically as in N. ericacea. Acropodite with small medial flange near apex (Fig. 35C, red triangle), not with lateral projection as in N. ericacea. Telopodite basal zone height ca. 1/3 length of acropodite, not ca. 1/2 length as in N. castanea, or < 1/5 as in N. ericacea. Prefemoral process small, thin, curving medially, arising from top of projected acuminate prefemoral spine, not large, laminate as in N. ericacea or straight as in N. castanea. Color. Tergites with orange paranotal spots (Fig. 36). Dark brown background. Dorsum of collum smooth with orange margin.

Figure 35.

Nannaria hokie Means, Hennen & Marek in Means et al. 2021, paratype ♂ (VTEC, MPE00880) left gonopod. A anterior view; red arrow indicates hood-like flange partially obscuring view of projected solenomere B medial view C posterior view; red triangle indicates medial flange. Scale bar: 0.5 mm.

Figure 36.

Nannaria hokie Means, Hennen & Marek in Means et al. 2021, holotype ♂ (VTEC, MPE04803) coloration. Scale bar: 4.0 mm.

Measurements

♂ paratype (VTEC, MPE00880): BL = 18.0, CW = 2.9, IW = 2.3, ISW = 0.8, B11W = 3.5, B11H = 2.2; ♀ paratype (VTEC, MPE00881): BL = 22.9, CW = 3.1, IW = 1.6, ISW = 0.8, B11W = 3.9, B11H = 2.4.

Variation

Little morphological variation exists between individuals collected from Montgomery County, Virginia; however, populations in Floyd and Pulaski counties, Virginia display a level of variation that suggests a closer comparison of these populations may be warranted. Both the Floyd and Pulaski County specimens have a much reduced acropodite “hood” (Fig. 37A, red arrow), and the Pulaski County specimens have highly variable prefemoral processes, including forms with and without a prefemoral spine (Fig. 37B, red triangle).

Figure 37.

Nannaria hokie Means, Hennen & Marek, 2021 variation A Floyd County, Virginia non-type ♂ (VMNH, NAN0003; red arrow indicates reduced lateral and medial flanges) B Pulaski County, Virginia non-type ♂ (VTEC, MPE00253); red triangle indicates prefemoral process lacking prefemoral spine. Scale bar: 0.5 mm.

Distribution

Known only from southwestern Virginia, primarily in the vicinity of Blacksburg, though a single specimen collected from Smyth County, Virginia extends the distribution of the species considerably (Virginia: Floyd, Montgomery, Pulaski, and Smyth counties; Suppl. material 7, Fig. 126). Distribution area: 1,668 km²; status: SRE.

Ecology

Individuals of N. hokie have been collected from mesic hardwood forests, composed of maple, oak, tuliptree, beech, spicebush, and witch hazel, under deciduous leaf litter, logs, and fallen bark. The Pulaski County specimens were all collected from a 1 m² section of a dry creek, taken from under 1–2 cm hard packed soil on the bank of the creek.

Etymology

Means et al. (2021) named N. hokie for its type locality, the campus of Virginia Tech. A hokie is a popular term for a member of the Virginia Tech community.

Type locality

United States, Virginia, Montgomery County, Blacksburg, Virginia Tech campus, Stadium Woods, 37.2216°N, -80.4159°W.

Notes

In the original publication Means et al. (2021: 17) designated a male holotype (VTEC MPE04803), five male paratypes (VTEC MPE00880, 882, 883; VMNH MPE00884, 886) and two female paratypes (VTEC MPE00881, 885) from the multiple locations on the campus of Virginia Tech collected by the authors from various dates in 2015 and 2019.

Nannaria missouriensis Chamberlin, 1928

Figs 38, 39 Vernacular name: “The Missouri Twisted-Claw Millipede”

Mimuloria missouriensis Chamberlin, 1928: 155; Chamberlin and Hoffman 1958: 37; Hennen and Shelley 2015: 9, fig. 15.

Nannaria missouriensis: Hoffman, 1964: 33; Hoffman 1999: 367. Marek et al. 2014: 37. Means et al. 2021: S70–S71.

Material examined

Holotype: United States – Missouri • ♂ St. Charles; 1926; M. J. Brown leg.; NMNH P-6.

Paratypes: United States – Missouri • 3 ♂♂; same collection data as holotype; NMNH P-9 • 3 ♀♀; same collection data as holotype; NMNH P-15 • 1 ♀; [Allotype] same collection data as holotype; NMNH IC.

Other material

United States – Missouri • 1 ♂, 1 ♀; Cole County, Jefferson Cirty; 38.5767°N, -92.1735°W; 1 Oct. 1944; W. Dowdy leg.; NCSM NAN0535 • 1 ♂, 1 ♀; Morgan County, Versailles; 38.4314°N, -92.8410°W; 1 Apr. 1959; J. Brooks leg.; NCSM NAN0536 • 1 ♀; St. Charles County, Weldon Spring Conservation Area, Lost Valley Trail; 38.6706°N, -90.7515°W; elev. 164 m; 18 May 2017; hand collected; J. Means, D. Hennen, V. Wong; VTEC MPE02800. For detailed collection data see Suppl. material 7.

Diagnosis

Adult males of Nannaria missouriensis are distinct from other Nannaria, and the nearby N. castanea, based on the following combination of characters: Gonopods. Gonopodal acropodite gently curving medially. Distal zone with medial and lateral flanges at 90° angle to solenomere (Fig. 38B, red arrow), not with medial flange at 135° angle to tip as in N. castanea. Acropodite with small medial flange near apex (Fig. 38A, red triangle), not lacking as in N. castanea. Telopodite basal zone height ca. 1/3 length of acropodite, not ca. 1/2 length as in N. castanea. Prefemoral process reduced to small bump arising from top of projected, stout prefemoral spine (Fig. 38A, red circle), not projected, acicular as in N. castanea. Color. Tergites with orange paranotal spots (Fig. 39) and occasionally faint orange stripes (Fig. 39). Dark to light brown background. Dorsum of collum smooth with orange margin.

Figure 38.

Nannaria missouriensis Chamberlin, 1928, holotype ♂ (NMNH, P-6) left gonopod A anterior view; red triangle indicates small medial flange on acropodite; red circle indicates reduced prefemoral process B medial view; red arrow indicates medial and lateral flanges on tip at 90° to acropodite C posterior view. Scale bar: 0.5 mm.

Figure 39.

Nannaria missouriensis Chamberlin, 1928 ♀ (VTEC, MPE02800) coloration. Scale bar: 4.0 mm.

Measurements

♂ holotype (NMNH): BL = N/A, CW = 2.9, IW = 1.7, ISW = 0.7, B11W = N/A, B11H = N/A; ♀ paratype (NMNH, IC): BL = 25.4, CW = 1.6, IW = 1.7, ISW = 0.7, B11W = 3.4, B11H = 2.3.

Variation

As Hennen and Shelley (2015) noted, there is some slight variation in the prefemoral process and spine between populations of N. missouriensis.

Distribution

Known from central and eastern Missouri (Missouri: Cole, Morgan, Phelps, and St. Charles counties, Suppl. material 7; Fig. 129). Distribution area: 10, 977 km²; status: WRE.

Ecology

Individuals of N. missouriensis have been collected from mesic deciduous forests dominated by pawpaw, maple, ironwood, and oak, often found under 1–2 cm dark, crumbly soil on hillsides.

Etymology

Chamberlin (1928) gave no etymology for Nannaria missouriensis, but it is reasonable to assume that the specific name is in reference the state of Missouri.

Type locality

United States, Missouri, St. Charles.

Note

In the original publication, Chamberlin (1928: 155) mentions eight type specimens, one of which he designated as the holotype (NMNH P-6). Chamberlin (1928) did not mention the sex of the paratypes, but we examined the type material and found three male and four female paratypes, one of which (NMNH IC) was labeled as an allotype. Type specimens were collected in 1926 by M. J. Brown.

Nannaria stellapolis sp. nov.

http://zoobank.org/1C878FE9-4057-4633-81E4-2231D93D7380

Figs 40, 41 Vernacular name: “The Star City Twisted-Claw Millipede”

Material examined

Holotype: United States – Virginia • ♂; Roanoke County, Roanoke Mountain picnic area off Blue Ridge Parkway; 37.2309°N, -79.9502°W; elev. 449 m; 25 Oct.2014; J. Means leg.; VTEC MPE00252.

Paratype: United States – Virginia • 1 ♂; same collection data as holotype; VMNH MPE00258. For detailed collection data see Suppl. material 7.

Diagnosis

Adult males of Nannaria stellapolis sp. nov. are distinct from other Nannaria and the nearby N. wilsoni, based on the following combination of characters: Gonopods. Gonopodal acropodite long and curving medially before apex, not ventromedially as in N. hokie. Acropodite tip with small, triangular lateral flange (Fig. 40A, red arrow), not encircling tip forming a hood-like structure as in N. hokie. Acropodite simple, lacking triangular medial flange of N. hokie. Acropodite without heavily sclerotized laminate corkscrew as in N. wilsoni. Telopodite basal zone height ca. 1/2 length of acropodite, not < 1/3 length as in N. hokie or ca. 1/6 length as in N. wilsoni. Telopodite with small basal zone lateral bulge. Prefemoral process small, thin, curving medially and arising from top of large, projected, shelf-like prefemoral spine—not arising from sharp, acuminate prefemoral spine as in N. hokie, and not subequal to length of acropodite as in N. wilsoni. Color. Tergites with orange/red paranotal spots (Fig. 41). Dark brown background. Dorsum of collum smooth with orange anterior margin.

Figure 40.

Nannaria stellapolis sp. nov. holotype ♂ (VTEC, MPE00252) left gonopod A anterior view; red arrow indicates lateral flange B medial view C posterior view. Scale bar: 0.5 mm.

Figure 41.

Nannaria stellapolis sp. nov. holotype ♂ (VTEC, MPE00252) coloration. Scale bar: 4.0 mm.

Measurements

♂ holotype (VTEC, MPE00252): BL = 24.2, CW = 3.3, IW = 1.7, ISW = 0.7, B11W = 3.7, B11H = 2.3.

Variation

No known variation.

Distribution

Known only from the type locality (Virginia: Roanoke County, Suppl. material 7; Fig. 126). Distribution area: N/A; status: MRE.

Ecology

Individuals of N. stellapolis sp. nov. were found in mesic broadleaved forests composed of oak, walnut, and maple, under 1–2 cm of hard packed soil on the bank of a dry creek bed.

Etymology

This species is named for Star City, the common nickname of Roanoke city. The specific name is a noun in apposition derived from the Latin stella-, star, and Greek polis, city.

Type locality

United States, Virginia, Roanoke County, Roanoke Mountain picnic area off Blue Ridge Parkway, 37.2309°N, -79.9502°W.

Nannaria stellaradix sp. nov.

http://zoobank.org/8AA670CB-8404-494A-B46D-0EC1415B3867

Figs 42, 43, 44 Vernacular name: “Starroot’s Twisted-Claw Millipede”

Material examined

Holotype: United States – Virginia • ♂; Montgomery County, Riner, 222 Milky way NW, in wetland below lake; 36.9662°N, -80.4179°W; elev. 773 m; 17 Oct. 2014; hand collected; J. Means leg.; VTEC MPE002331.

Paratype: United States – Virginia • 1 ♂; same collection data as holotype; VMNH MPE00241 • 3 ♀♀; same collection data as holotype; VTEC MPE00238–240 • 2 ♀♀; same collection data as holotype; VMNH MPE00242, 244.

Other material

United States – Virginia • 1 ♂; Floyd County, 2 mi. SW of Copper Valley; 36.9702°N, -80.5425°W; 15 Oct. 1974; R. Hoffman leg.; VMNH NAN0146 • 1 ♂; Floyd County, Big Indian Creek, ca. 2 mi. S of Copper Valley; 36.9615°N, -80.5171°W; 16 Apr. 1970; B. Coombs leg.; VMNH NAN0147 • 1 ♂; Pulaski County, Powhatan Scout Reservation, ca. 1 mi. E. of Macks Creek P.O.; 36.9645°N, -80.6626°W; 6 Mar. 1976; R. Hoffman leg.; VMNH NAN0165. For detailed collection data see Suppl. material 7.

Diagnosis

Adult males of Nannaria stellaradix sp. nov. are distinct from other Nannaria and the sympatric N. wilsoni, based on the following combination of characters: Gonopods. Gonopodal acropodite continually curving medially before apex. Distal zone short, directed medially with small triangular lateral flange (Fig. 42A, red arrow), not directed caudally with enveloping lateral and medial flanges as in N. hokie, or enlarged, laminate, as in N. wilsoni. Acropodite with small laminate medial flange (Fig. 42A, red triangle). Basal zone > 1/3 length of acropodite, not < 1/5 as in N. wilsoni. Prefemur with enlarged prefemoral spine. Prefemur with small, ventrally curving prefemoral process arising dorsomedially from spine, not from top of spine as in N. hokie, or long, medially curving, paralleling acropodite as in N. wilsoni. Color. Tergites with light orange paranotal spots (Fig. 43). Dark brown background. Dorsum of collum smooth with orange and white margin.

Figure 42.

Nannaria stellaradix sp. nov. holotype ♂ (VTEC, MPE00233) left gonopod A anterior view; red arrow indicates lateral flange; red triangle indicates small medial flange B medial view C posterior view. Scale bar: 0.5 mm.

Figure 43.

Nannaria stellaradix sp. nov. holotype ♂ (VTEC, MPE00233) coloration. Scale bar: 4.0 mm.

Measurements

♂ holotype (VTEC, MPE00233): BL = 23.8, CW = 2.9, IW = 1.8, ISW = 0.8, B11W = 3.7, B11H = 2.6; ♀ paratype (VTEC, MPE00238): BL = 25.4, CW = 3.4, IW = 2.2, ISW = 0.9, B11W = 3.9, B11H = 2.8.

Variation

Three populations from Pulaski and Floyd counties in Virginia (VMNH, NAN0146, 147, 165) show some morphological variation (Fig. 44) that may indicate a separate species; however, without molecular evidence, specific recognition is not warranted. Notable variations in these populations include a prefemoral process arising from the top of the prefemoral spine (Fig. 44, red arrow) and the lack of an acropodite medial flange.

Figure 44.

Nannaria stellaradix sp. nov. non-type ♂ (VMNH, NAN0165) from Pulaski County, Virginia. Red arrow indicates prefemoral process arising from the top of the prefemoral spine. Scale bar: 0.5 mm.

Distribution

Known from a linear area in southern Virginia (Virginia: Floyd and Pulaski counties, Suppl. material 7; Fig. 126). Distribution area: 11 km²; status: MRE.

Ecology

Individuals of N. stellaradix sp. nov. were collected from mesic mixed hardwood and pine habitats predominately composed of maple and white pine; individuals were collected from under leaf litter.

Etymology

This species is named after the owner of the property from which the holotype was found, an artist named Starroot. The specific name is a noun in apposition derived from the Latin stella-, star, and -radix, root.

Type locality

United States, Virginia, Montgomery County, Riner, 222 Milky way NW, in wetland below lake, 36.9662°N, -80.4179°W.

laminata clade

Components. Nannaria cingulata sp. nov. and N. laminata. Members of the laminata clade share gonopodal characters, including a blunt acropodite tip and pronounced prefemoral spine. The molecular phylogeny (Fig. 7) would suggest that this clade is either one highly variable species, where eastern and northern forms have developed a cingulum, a basal bulge and a straight acropodite (N. cingulata sp. nov.) or, except for N. cingulata sp. nov., a fairly morphologically uniform but genetically diverse group of species. Sequencing of additional specimens throughout the range of the laminata clade will help illuminate which of the above scenarios are accurate, however, due to the number of morphological apomorphies found in N. cingulata sp. nov. specimens, we feel confident in naming N. cingulata sp. nov. as a separate species from N. laminata.

Distribution. The laminata clade extends from much of Virginia and the D.C. metro area into eastern West Virginia (Fig. 114).

Nannaria cingulata sp. nov.

http://zoobank.org/E1CC5FC5-266A-4B04-886B-22587B78B4C4

Figs 45, 46, 47 Vernacular name: “The Swamp-Dwelling Twisted-Claw Millipede”

Material examined

Holotype: United States – Virginia • ♂; Shenandoah County, Massanutten Trail nr. Signal Knob parking off Route 678; 38.9332°N, -78.3207°W; elev. 250 m; 24 Feb. 2017; hand collected; C. Harden leg.; VTEC MPE02324.

Paratypes: United States – Virginia • 11 ♂♂; same collection data as holotype; VTEC MPE02325-2335 • 11 ♂♂; same collection data as holotype; VMNH MPE02336-45, 3668 •1 ♀; Clarke County, nr. Mount Weather; 39.0717°N, -77.9120°W; elev. 326 m; 22 June 2017; hand collected; J. Means, D. Hennen leg.; VTEC MPE01881.

Other material

United States – Virginia • 1 ♂; Chesterfield County, Pocahontas State Park; 37.3679°N, -77.5755°W; 11 May 2002; A. Evans leg.; VMNH NAN0221 • 1 ♂; Clarke County, 1.7 miles by road N of Ashby Gap, U.S. Rt. 50 crossing of Blue Ridge; 39.0133°N, -77.9622°W; 10 May 1958; W. Highton leg.; VMNH NAN0227 • 2 ♂; Fairfax County, Hemlock Overlook Regional Park challenge course; 38.7666°N, -77.4104°W; elev. 101 m; 8 Apr. 2017; hand collected; P. Marek, C. Hall leg.; VTEC MPE02438, 2447 • 2 ♂; Fauquier County, Sky Meadows State park, near junction of North Ridge Trail and Appalachian Trail; 38.9850°N, -77.9900°W; elev. 549 m; 21 Apr. 2017; hand collected; C. Harden leg.; VTEC MPE02473, 77 • 2 ♂♂; Frederick County, Albin, ca. 3 mi. W of Winchester on VA. Hy 679; 39.2216°N, -78.1988°W; 10 Apr. 1990; LFCC class leg.; VMNH NAN0225 • 1 ♂; Prince Edward County, Hampden-Sydney College; 37.2381°N, -78.4606°W; 20 April 1983; J. White leg.; VMNH NAN0041 • 3 ♂♂; NAN0230 Prince George County, Lanham; 38.9669°N, -76.8622°W; 1 June; H. Loomis leg.; VMNH NAN0230 • 1 ♂; Shenandoah County, Massanuten Trail ca. 2.5 mi from Signal Knob parking lot by Rte. 698; 38.9324°N, -78.3249°W; elev. 260 m; 24 Feb. 2017; hand collected; C. Harden, G. Chapman leg.; VTEC MPE02348 • 2 ♂♂; Shenandoah County, George Washington National Forest near Eliz. Furn. Group Cmpgrd; 38.9310°N, -78.3220°W; elev. 233 m; 26 Feb. 2017; hand collected; C. Harden leg.; VTEC MPE03437, 03438 • 3 ♂♂; Shenandoah County, George Washington National Forest, Botts Trail, south of Elizabeth Furnace; 38.9240°N, -78.3280°W; elev. 250 m; 21 Apr. 2017; hand collected; C. Harden leg.; VTEC MPE02475, 76, 3673 • 2 ♂♂; Shenandoah County, dry creak near swamp just past Elizabeth Furnace Campground; 38.9122°N, -78.3326°W; elev. 248 m; 22 Apr. 2018; hand collected; J. Means, N. Larson leg.; VTEC MPE03899, 03900 • 1 ♀; same collection data as preceding; VTEC, MPE03901 • 1 ♂; Suffolk County, Magnolia, US Hys. 58-460; 36.7458°N, -76.5408°W; 23 Feb. 1954; R. Ragot leg.; VMNH NAN0229 • 1 ♂; Warren County, Smithsonian Conservation Biology Institute, High Knob Mountain; 38.8930°N, -78.1370°W; elev. 396 m; 13 Mar. 2017; hand collected; C. Harden leg.; VTEC MPE02463 • 6 ♂♂; Warren County, Racetrack Hill, gravel road/field; 38.8915°N, -78.1680°W; elev. 305 m; 4 Apr. 2017; hand collected; C. Harden leg.; VTEC MPE02453–56, 2786, 2787 • 1 ♂; Warren County, SCBI, High Knob Mountain; 38.8930°N, -78.1370°W; elev. 390 m; 6 Mar. 2017; hand collected; C. Harden leg.; VTEC MPE03744 • 1 ♂; Warren County, upper reaches of Stokes Branch 0.17 mi N of dirt forest road terminus, off Rte 613 nr jct w/608; 38.9156°N, -78.2994°W; elev. 265 m; 12 Feb. 2018; hand collected; C. Harden leg.; VTEC MPE03885 • 26 ♂♂; Warren County, 2 mi SE Front Royal, DF site at NZP-CRC, oak woods on hillside; 38.8975°N, -78.1683°W; 1 May 1996; VMNH Survey leg.; VMNH NAN0232 • 4 ♂; York County, Cheatham Annex NSD Jones Mill Pond; 37.2850°N, -76.6416°W; 17 Nov. 1989; Buhlmann, Pague leg.; VMNH NAN0222 • 1 ♂; same collection data as preceding; 2 Feb. 1997; S. Roble leg.; VMNH NAN0223 • 1 ♂; same collection data as preceding; 2 Nov. 1989; K. Buhlmann leg.; VMNH, NAN0224 • 2 ♂♂; same collection data as preceding; 20 Feb. 1990; VDNH survey leg.; VMNH NAN0228 • 1 ♂; same collection data as preceding; 17 Nov. 1989; K. Buhlmann leg.; VMNH NAN0359; SCAU – West Virginia • 2 ♂♂, 2 ♀♀; Berkeley County, Sleepy Creek Hunt Area; 39.5057°N, -78.1695°W; 6 May 1968; P. Martinat leg.; NCSM NAN0449, 450. For detailed collection data see Suppl. material 7.

Diagnosis

Adult males of Nannaria cingulata sp. nov. are distinct from other Nannaria and the sympatric N. shenandoa, based on the following combination of characters: Gonopods. Gonopodal acropodite gently curving medially before bending dorsomedially, not ventromedially as in N. paupertas sp. nov. and N. shenandoa. Distal zone short, bent dorsomedially at 45°, not bent dorsomedially at 90° as in N. paupertas sp. nov., or curving posterolaterally as in N. shenandoa. Acropodite tip blunt with small, lobed lateral flange (Fig. 45C, red arrow), not with large, pointed flange as in N. shenandoa. Telopodite basal zone height < 1/2 length of acropodite with lateral bulge (Fig. 45A, red triangle), not greatly reduced as in N. shenandoa, or lacking lateral bulge as in N. paupertas sp. nov. Acropodite shaft swollen before apex, with cingulum (Fig. 45A, red rectangle), not simple as in N. shenandoa, or with medial flange as in N. paupertas sp. nov. Prefemur with prefemoral process straight for first 3/4, and bent ventrally at 90° in last 1/4, not bent ventromedially as in N. paupertas sp. nov., or curving laterally as in N. shenandoa. Prefemoral process ca. 1/2 length of acropodite, not 2/3 length as in N. paupertas sp. nov. or N. shenandoa. Prefemoral spine greatly reduced to small, tooth-like projection (Fig. 45C, red oval), not prominent, acicular as in N. paupertas sp. nov., or absent as in N. shenandoa. Color. Tergites with orange paranotal spots (Fig. 46A) and occasionally faint orange stripes (Fig. 46B). Dark to light brown background. Dorsum of collum smooth with orange margin.

Figure 45.

Nannaria cingulata sp. nov. holotype ♂ (VTEC, MPE02324) left gonopod A anterior view; red triangle indicates basal lateral bulge; red rectangle indicates cingulum B medial view C posterior view; red arrow indicates small lateral flange on acropodite tip; red oval indicates reduced prefemoral spine. Scale bar: 0.5 mm.

Figure 46.

Nannaria cingulata sp. nov. coloration A holotype ♂ (VTEC, MPE02324) orange paranota B non-type ♀ (VTEC, MPE01881) faint orange stripes. Scale bars: 4.0 mm.

Measurements

♂ holotype (VTEC, MPE02324): BL = 24.7, CW = 3.9, IW = 2.1, ISW = 0.8, B11W = 4.4, B11H = 2.9. ♀ paratype (VTEC, MPE01881): BL = 28.6, CW = 3.9, IW = 2.7, ISW = 0.9, B11W = 4.8, B11H = 3.3.

Variation

The only Nannaria cingulata sp. nov. individual known from Prince Edward Co., Virginia has a cephalically directed prefemoral process and a pronounced prefemoral spine (Fig. 47, red arrow).

Figure 47.

Nannaria cingulata sp. nov. (VMNH, NAN0041) from Prince Edward Co., Virginia. Red arrow indicates projected prefemoral spine. Scale bar: 0.5 mm.

Distribution

Nannaria cingulata sp. nov. has a more extensive range than most species in the minor species group and can be found from eastern West Virginia south to central and southeastern Virginia, and Maryland in the Washington D.C. metropolitan area (West Virginia: Berkley County; Virginia: Frederick, Clarke, Shenandoah, Warren, Fairfax, Chesterfield, Prince Edward, Gloucester counties, and the City of Suffolk; Suppl. material 7; Fig. 128). Distribution area: 28,825 km²; status: WRE.

Ecology

The majority of Nannaria cingulata sp. nov. specimens collected by C. Harden have been taken from mesic hardwood forests at night and were found walking on top of leaf litter and on man-made paths, or were collected using pitfall traps. Several museum specimens of N. cingulata sp. nov. lack ecological notes.

Etymology

This species is named for the presence of a cingulum, a linear groove and possible point of flexion, on its gonopod, a characteristic unique within Nannaria. The specific name is a feminized adjective derived from the Latin cingulatus for belted.

Type locality

United States, Virginia, Shenandoah County, Massanutten Trail nr. Signal Knob parking off Route 678, 38.9332°N, -78.3207°W.

Nannaria laminata Hoffman, 1949

Figs 48, 49 Vernacular name: “The Laminate Twisted-Claw Millipede”

Nannaria laminata Hoffman, 1949: 383, figs 11, 12. Chamberlin and Hoffman 1958: 41. Hoffman 1999: 367. Marek et al. 2014: 37. Means et al. 2021: S70.

Material examined

Holotype: United States – West Virginia • ♂; Mercer County, ravine beside U.S. Route 460, ca. 2 miles south of Glen Lyn, Virginia; [37.3550°N, -80.8982°W]; 12 July 1947; H. H. Hobbs, C. M. Wilson leg.; NMNH Type #1806.

Other material

United States – Virginia • 1 ♂; Amherst County, Tarjacket Ridge FS 1167, 3500’; 37.7665°N, -79.1863°W; 13 Nov. 1999; VMNH Survey leg.; VMNH NAN0332 • 1 ♂; Augusta County, across creek from Barnwood Cabin, where old road flattens out by small stream; 37.89643°N, -79.0007°W; elev. 828 m; 18 Mar. 2017; J. Means leg.; VTEC MPE02401 • 1 ♀; same collection data as preceding; VTEC MPE03679 • 1 ♂; Augusta County, far corner of upper Sherando Lake, bank side by marsh; 37.9139°N, -79.0207°W; elev. 607 m; 19 Mar. 2017; J. Means leg.; VTEC MPE02392 • 1 ♂; Bedford County, Flattop Mtn., Peaks of Otter nr MP 82.8; 37.4498°N, -79.5834°W; 27 Oct. 1986; J. Mitchell leg.; VMNH NAN0330 • 3 ♂♂; Bland County, Hamilton’s Cave, ca. 6 km ENE of Mechanicsville, rich wooded hillside; 37.7032°N, -78.1627°W; 16 May 1980; R. Hoffman leg.; VMNH NAN0150 • 2 ♂♂; Botetourt County, Harkening Hill, Blue Ridge Parkway; 37.4577°N, -79.6175°W; 21 Oct. 1989; J. Mitchell leg.; VMNH NAN0333 • 1 ♂; Botetourt County, North Creek, 1–3 mi. east of Arcadia; 37.5411°N, -79.5866°W; 13 Oct. 1973; R. Hoffman leg.; VMNH NAN0334 • 1 ♂; Campbell County, 4 miles NW of Rustburg; 37.3176°N, -79.1526°W; 14 Oct. 1950; L. Hubricht leg.; VMNH NAN0331 • 1 ♂; Cumberland County, clearcut north DF site 2 ca 2 km SSW of Columbia; 37.7361°N, -78.1714°W; 17 May 1990; J. Mitchell leg.; VMNH NAN0253 • 3 ♂♂; same collection data as preceding; Nov. 1989; VMNH NAN0259 • 1 ♂; same collection data as preceding; 2 Apr. 1990; VMNH NAN0261 • 1 ♂; same collection data as preceding; 3 Dec. 1989; VMNH NAN0263 • 1 ♂; same collection data as preceding; 15 Feb. 1990; VMNH NAN0264 • 19 ♂♂; same collection data as preceding; 19 Oct. 1989 VMNH NAN0266 • 7 ♂♂; same collection data as preceding; 16 Nov. 1989; VMNH NAN0267 • 11 ♂♀; same collection data as preceding; Nov. 1989; VMNH NAN0361 • 6 ♂♂; same collection data as preceding; 1 May 1990; VMNH NAN0255 • 1 ♂; Cumberland County, hardwood site 1 (north), 2 km SSW of Columbia; 37.7361°N, -78.1715°W; 1 May 1990; J. Mitchell leg.; VMNH NAN0254 • 24 ♂♂; same collection data as preceding; 17 Mar. 1990; VMNH NAN0268 • 1 ♂; Cumberland County, hardwood site 4 (south), 7 km SSW of Columbia; 37.6953°N, -78,1959°W; 5 Oct. 1989; J. Mitchell; VMNH NAN0257 • 3 ♂♂; Cumberland County, hardwood site 2 – N 2 km SSW Columbia; 37.7361°N, -78.1714°W; VMNH NAN0258 • 2 ♂♂; Cumberland County, pinewoods DF site 5, ca. 5.5 km SSW of Columbia; 37.7048°N, -78.1889°W; 17 May 1990; J. Mitchell leg.; VMNH NAN0532 • 1 ♂; Giles County, Jefferson National Forest, Cascades Day Use Area, within a few hundred meters of the start of the Cascades Trail, near parking lot; 37.3500°N, -80.5835°W; elev. 544 m; 21 Oct. 2015; hand collected; D. Hennen leg.; VTEC MPE00891 • 1 ♂; Giles County, side of Upper Trail on way to Cascades; 37.3613°N, -80.5875°W; elev. 799 m; 18 Feb. 2018; hand collected; J. Means leg.; VTEC MPE03807 • 1 ♂; Giles County, Jefferson National Forest past Mountain Lake Biological Station, past Mini Ball Hill on Rt 613; 37.4209°N, -80.5093°W; elev. 1062 m; 3 Oct. 2016; hand collected; J. Means, D. Hennen, V. Wong leg.; VTEC MPE02124 • 2 ♀♀; same collection data as preceding; VTEC MPE02136, 2142 • 1 ♂; Prince Edward County, Farmville, Price Drive; 37.3019°N, -78.3922°W; 28 June 1988; W. Shear leg.; VMNH NAN0256 • 1 ♂; Prince Edward County, 13 mi S Farmville, dug up 12 cm under surface of clay; 37.1134°N, -78.3922°W; 22 Feb. 1975; W. Shear leg.; VMNH NAN0260 • 1 ♂; Prince Edward County, pitfall trap, old field at Rice; 37.2750°N, -78.2916°W; 17 June 1981; R. Bellinger leg.; VMNH NAN0265; SCAU – West Virginia • 1 ♀; Mercer County, just over the WV state line on 460, ca. 2 miles SW of Glen Lyn on 219/8; 37.3452°N, -80.9105°W; elev. 507 m; 12 Nov. 2017; hand collected; J. Means leg.; VTEC MPE03474 • 1 ♂; Monroe County, 5.8 air miles east of Linside, up logging road; 37.4749°N, -80.5673°W; elev. 934 m; 27 Apr. 2017; hand collected; J. Means, D. Hennen, P. Marek, P. Shorter, V. Wong leg.; VTEC MPE02520 • 3 ♀♀; same collection data as preceding; VTEC, MPE02578, 2579, 3675 • 2 ♂♂; Monroe County, along logging road off VA rt 613, down in bowl and on each side of dirt and gravel road; 37.4717°N, -80.5620°W; elev. 1107 m; 27 Apr. 2016; hand collected; J. Means, D. Hennen, P. Marek, P. Shorter, V. Wong leg.; VTEC MPE02528, 2561. For detailed collection data see Suppl. material 7.

Diagnosis

Adult males of N. laminata are distinct from other Nannaria and the nearby N. solenas sp. nov. and N. wilsoni, based on the following combination of characters: Gonopods. Gonopodal acropodite semi-circular, curving dorsomedially with abrupt 90° curve after apex, not straight before abrupt 90° curve as in N. solenas sp. nov. or with laminate corkscrew before apex as in N. wilsoni. Acropodite tip simple, directed posteriorly, not directed anteriorly as in N. solenas sp. nov. and not with triangular lateral flange as in N. wilsoni. Tip terminating in sharp claw-like point, not blunt point as in N. solenas sp. nov. and N. wilsoni. Height of telopodite basal zone > 1/3 length of acropodite, not ca. 1/2 length as in N. solenas sp. nov., or ca. 1/5 as in N. wilsoni. Prefemoral process large, laminate and serpentine, with prefemoral spine reduced to small, acuminate projection (Fig. 48C, red arrow), not lacking as in N. wilsoni. Color. Tergites with red paranotal spots (Fig. 49). Black background. Dorsum of collum smooth with red margin.

Figure 48.

Nannaria laminata holotype ♂ (NMNH, Type #1806) left gonopod A anterior view B medial view C posterior view; red arrow indicates reduced, acuminated prefemoral spine. Scale bar: 0.5 mm.

Figure 49.

Nannaria laminata non-type ♂ (VTEC, MPE02392) coloration. Scale bar: 4.0 mm.

Measurements

♂ holotype (NMNH, Type #1806): BL = 30.5, CW = 4.0, IW = 2.2, ISW = 0.9, B11W = 4.9, B11H = 3.4.

Variation

Nannaria laminata occupies a relatively large geographic area, > 10,000 km², and displays a fair amount of morphological variation; however, the general shape of the acropodite and laminate prefemoral process remain constant throughout. Variation is seen in the sharpness and curve of the acropodite tip; some individuals display a more medially directed, blunt tip than the holotype. Additionally, the width of the prefemoral process (when viewed anteriorly or posteriorly) varies widely between individuals.

Distribution

Known from a horizontal strip from central Virginia to just north of the border into West Virginia (Virginia: Amherst, Augusta, Bedford, Botetourt, Campbell, Cumberland, Giles, and Prince Edward counties; West Virginia: Mercer and Monroe counties, Suppl. material 7; Fig. 126). Distribution area: 9,297 km²; status: SRE.

Ecology

Individuals of N. laminata have been collected from mesic deciduous forests dominated by maple, oak, rhododendron, and some pine, often under 1–2 cm soil and/or leaf litter.

Etymology

Hoffman (1949) gives no explanation for the name laminata in his description of the species, however it is reasonable to assume that the name refers to the laminate prefemoral process, the major distinguishing morphological character.

Type locality

United States, West Virginia, Mercer County, ravine beside U.S. Route 460, ca. 2 miles south of Glen Lyn, Virginia; [37.3550°N, -80.8982°W].

Notes

In the original publication, Hoffman (1949: 384) designates a male holotype (NMNH Type #1806) collected by H. Hobbs and C. Wilson on July 12, 1947.

tasskelsoae clade

Components. Nannaria tasskelsoae sp. nov. Based on the molecular phylogeny (Fig. 7) N. tasskelsoae sp. nov. is in a monotypic clade (Fig. 113). The tasskelsoae clade can be diagnosed by the combination of morphological characters detailed in the below diagnosis, including an enlarged prefemoral spine from which arises a medially curving prefemoral process which parallels the medially curving, simple acropodite.

Distribution. the tasskelsoae clade is known only from eastern West Virginia (Fig. 127).

Nannaria tasskelsoae sp. nov.

http://zoobank.org/F3B40E44-28D4-4353-BF02-78B6398996D6

Figs 50, 51 Vernacular name: “Tass Kelso’s Twisted-Claw Millipede”

Material examined

Holotype: United States – West Virginia • ♂; Greenbrier Co., Monongahela National Forest, Summit Lake Campground; 38.2490°N, -80.4437°W; elev. 1062 m; 20 July 2005; hand collected; P. Marek, C. Spruill leg.; VTEC SPC000710.

Paratypes: United States – West Virginia • 1 ♂; same collection data as holotype; FMNH SPC000711 • 1 ♀; same collection data as holotype; VTEC SPC000712 • 1 ♀; same collection data as holotype; VMNH SPC000713 • 1 ♂; Pocahontas Co., Droop Mountain Battlefield State Park, near Little Levels between cemetery, lookout, and park area; 38.1101°N, -80.2733°W; elev. 942 m; 1 June 2015; hand collected; C. Hall leg.; VTEC MPE00578 • 1 ♀; same collection data as preceding; VTEC MPE00580.

Other material

United States – West Virginia • 4 ♂♂; Greenbrier County, Summit Lake Campground, small swift stream hollow downhill from campsite #20; 38.2498°N, -80.4455°W; elev. 1056 m; 19 May 2018; hand collected; C. Harden, G. Chapman leg.; VTEC MPE04022, 23, 31, 32 • 5 ♂♂; Greenbrier County, Kate’s Mountain summit, Greenbrier State Forest; 37.7374°N, -80.3324°W; 20 Apr. 1971; W. Shear leg.; VMNH NAN0042 • 1 ♂; Greenbrier County, Second Ck, 1 mi E, 1.4 mi S Ft. Spring; 37.7162°N, -80.5388°W; 12 Oct. 2000; W. Arnold leg.; VMNH NAN0079 • 1 ♂; Greenbrier County, White Sulfur Springs, Kate’s Mountain summit; 37.7622°N, -80.3013°W; 20 Apr. 1968; W. Shear leg.; VMNH NAN0082 • 1 ♂; Monroe County, Laurel Creek Cave, Greenville; 37.5440°N, -80.6818°W; 16 Apr. 1972; W. Shear leg.; VMNH NAN0040 • 1 ♂; Nicholas County, Monongahela NF, Woodbine Picnic Area; 38.2897°N, -80.5353°W; elev. 646 m; 10 May 2011; Ashley Bailey leg.; VTEC SPC001100 • 1 ♀; Nicholas County, Monongahela NF, FR-76 (E side), 5.15 rd km N WV-39/55; 38.2728°N, -80.5250°W; elev. 806 m; 20 July 2005; hand collected; P. Marek, C. Spruill leg.; VTEC SPC000715 • 1 ♂; Pocahontas County, Kennison Mountain Trail, ca.0.5 mi from trailhead on Hwy 39; 38.1906°N, -80.2858°W; elev. 1261 m; 19 May 2018; hand collected; C. Harden, G. Chapman leg.; VTEC MPE04021 • 1 ♀; Pocahontas County, Watoga State Park, Dragon Draft Trail; 38.1190°N, -80.1540°W; elev. 739 m; 18 July 2005; hand collected; P. Marek, C. Spruill leg.; VTEC SPC000689 • 2 ♀♀; Pocahontas County, Watoga State Park, Riverside Campground; 38.1119°N, -80.1783°W; elev. 636 m; 19 July 2005; hand collected; P. Marek, C. Spruill leg.; VTEC SPC000694, 695 • 1 ♂; Pocahontas County, Cranberry Glades Natural Area, Monongahela Nat. Forest; 38.2000°N, -80.2716°W; 20 May 1967; W. Shear leg.; VMNH NAN0016 • 1 ♂; Pocahontas County, Hills Creek Falls, Monongahela Nat. Forest; oak-hickory-beech-maple transition; 38.1761°N, -80.3365°W; 20 May 1967; W. Shear leg.; VMNH NAN0021 • 2 ♂♂; same collection data as preceding; 8 July 1967; W. Shear leg.; VMNH NAN0022 • 1 ♂; same collection data as preceding; 18 May 1968; W. Shear leg.; VMNH NAN0023 • 1 ♂; same collection data as preceding; 24 Sep. 1972; W. Shear leg.; VMNH NAN0039 • 2 ♂♂; same collection data as preceding; 8 May 1971; W. Shear leg.; VMNH NAN0044 • 10 ♂♂; same collection data as preceding; 5 May 1970; VMNH NAN0078 • 1 ♂; Pocahontas County, Droop Mtn State Park; 38.1150°N, -80.2694°W; 30 Apr. 1967; W. Shear leg.; VMNH NAN0024 • 1 ♀; Randolph County, Kumbrabow State Forest, off Kumbrabow Rd. near Mill Creek; 38.6550°N, -80.0713°W; elev. 922 m; 1 June 2015; hand collected; C. Hall leg.; VTEC MPE00585 • 1 ♀; Randolph County, Gatewood CpGrnd; 37.7622°N, -80.3013°W; elev. 1322 m; 20 Apr. 1968; W. Shear leg.; VMNH NAN0080 • 1 ♂; Randolph County, between 1–2 miles W of Bear Heaven Picnic Area on road to Bickle’s Knob nr. Alpena; 38.9367°N, -79.7029°W; 12 June 1986; Highton, Barry leg.; VMNH NAN0152 • 3 ♂♂; Randolph County, along US 33, 2.1 W Alpena; 38.9101°N, -79.6900°W; 23 Aug. 1978; R. Shelley, C. Withrow leg.; NCSM NAN0486 • 3 ♂♂; Summers County, 9 miles SE of Bellepoint on W. Va. Hy. 12; 37.5385°N, -80.7897°W; 22 Sep. 1962; R. Hoffman leg.; VMNH NAN0149 • 1 ♂; Tucker County, Backbone Mtn.; 39.1470°N, -79.5700°W; elev. 1067 m; 30 June 1968; W. Muchmore leg.; VMNH NAN0148 • For detailed collection data see Suppl. material 7.

Diagnosis

Adult males of Nannaria tasskelsoae sp. nov. are distinct from other Nannaria and the sympatric N. shenandoa, based on the following combination of characters: Gonopods. Gonopodal acropodite gently curving medially before apex, not straight as in N. shenandoa. Distal zone short, bent medially forming 130° angle with acropodite (Fig. 50). Telopodite basal zone ca. 1/3 length of acropodite, not < 1/3 as in N. fowleri and N. shenandoa. Acropodite tip and distal zone simple, rectangular, blunt, < 1/8 length of acropodite, not large, curving with flange as in N. shenandoa. Prefemoral process arising from top of prefemoral spine (Fig. 50A, red arrow), paralleling medial curve of acropodite, not arising from bottom of prefemoral spine as in N. fowleri, or curving laterally as in N. shenandoa. Prefemoral spine large, curving cephalically, not stout, tooth-like as in N. fowleri. Color. Tergites with orange paranotal spots (Fig. 51). Black background. Dorsum of collum smooth with orange margin.

Figure 50.

Nannaria tasskelsoae sp. nov. holotype ♂ (VTEC, SPC000710) left gonopod A anterior view; red arrow indicates pronounced, curving prefemoral spine B medial view C posterior view. Scale bar: 0.5 mm.

Figure 51.

Nannaria tasskelsoae sp. nov. non-type ♂ (VTEC, MPE04021) coloration. Scale bar: 4.0 mm.

Measurements

♂ holotype (VTEC, SPC000710): BL = 30.5, CW = 3.9, IW = 2.4, ISW = 0.9, B11W = 5.0, B11H = 3.3; ♀ paratype (VTEC, SPC000712): BL = 34.4, CW = 4.3, IW = 2.2, ISW = 1.1, B11W = 5.3, B11H, 4.1.

Variation

The most notable variation amongst individuals of N. tasskelsoae sp. nov. is found in the southern range of the species’ distribution. One specimen from Greenville, West Virginia (NAN0040) has a sudden lateral curve in the tip of its prefemoral process, forming a cat-claw like bend, while another specimen from near Forest Hill, West Virginia (NAN0149) has a laminate medial flange that is distal to the acropodite apex. Additional variation between populations is minimal, and is primarily found in the degree to which the prefemoral process parallels the acropodite and the size of the prefemoral spine.

Distribution

Nannaria tasskelsoae sp. nov. has a linear distribution running the length of the eastern edge of West Virginia (West Virginia: Randolph, Nicholas, Greenbrier, Monroe, Pocahontas, Summers, and Tucker counties; Suppl. material 7; Fig. 127). Distribution area: 3,623 km²; status: SRE.

Ecology

Specimens of N. tasskelsoae sp. nov. were collected from mesic hardwood forests composed of beech, black cherry, oak, sugar maple, tuliptree, and yellow birch. Several specimens were found on hillsides, including those from Summit Lake Campground which were active on top of leaf litter at night.

Etymology

This species is named after Dr. Tass Kelso, systematic botanist. Dr. Kelso was mentor to the discoverer of this species, Charity Hall. The specific name is a genitive noun derived as a matronym.

Type locality

United States, West Virginia, Greenbrier Co., Monongahela National Forest, Summit Lake Campground; 38.2490°N, -80.4437°W.

tennesseensis clade

Components. Nannaria breweri sp. nov., N. equalis Chamberlin, 1949, N. fritzae sp. nov., N. monsdomia sp. nov., N. tennesseensis, and a female specimen from Dungannon, Virginia. The tennesseensis clade represents one of the more heterogeneous Nannaria clades, with a variety of gonopod morphologies and a widespread geographic distribution, including the most southern species of the minor group, N. fritzae sp. nov. There are a few generally shared gonopodal characters, however, including a blunt acropodite tip and a medially directed prefemoral process, with the exception of N. monsdomia sp. nov. and N. equalis (Fig. 113). No specimens of N. equalis were collected for this study, however due to close geographic proximity and shared gonopodal characters, such as a blunt acropodite tip and a prominent prefemoral spine (as seen in N. monsdomia sp. nov.), we place N. equalis within the tennesseensis clade.

Distribution. The tennesseensis clade extends from northwestern Georgia, into eastern Tennessee, and southwestern Virginia (Fig. 114).

Nannaria breweri sp. nov.

http://zoobank.org/6B041BC2-5674-4133-993F-FFCDCAC54252

Figs 52, 53 Vernacular name: “Michael Brewer’s Twisted-Claw Millipede”

Material examined

Holotype: United States – Tennessee • ♂; Hamblen County, Morristown, Panther Creek State Park, in gully below parking lot at Spoone Recreation Area, near Ore Mine Trail; 36.2167°N, -83.4055°W; elev. 373 m; 9 Oct. 2016; hand collected; J. Means and D. Hennen leg.; VTEC MPE02191.

Paratypes: United States – Tennessee • 1 ♂; same collection data as for holotype; VTEC MPE02198 • 2 ♂♂; same collection data as for holotype; VMNH MPE02205, 3730 • 5 ♀♀; same collection data as for holotype; VTEC MPE02197, 2202–4, 3719 • 4 ♀♀; same collection data as for holotype; VMNH MPE03720, 3731–3.

Other material

United States – Tennessee • 1 ♂; Grainger County, 6.8 miles S of Rutledge; 36.1822°N, -83.5150°W; 18 May 1956; Lund, Keeton, R. Hoffman leg.; VMNH NAN0324 • 2 ♀♀; Hamblen County, Panther Creek State Park, Seven Sinkholes Trail; 36.2160°N, -83.4057°W; elev. 343 m; 14 Oct. 2007; hand collected; P. Marek, L. Swafford, M. Brewer, C. Hall, K. Bader leg; VTEC SPC001167, 1179 • 5 ♂♂; Hamblen County, Cherokee Lake Bluff, 3.5 miles N of Morristown; 36.2646°N, -83.2950°W; 19 May 1961; L. Hubricht leg.; VMNH NAN0309; SCAU – Virginia • 1 ♂; Tazewell County, Burkes Garden, on Garden Mountain; 37.0809°N, -81.4145°W; elev. 1158 m; 27 Oct. 1970; R. Hoffman leg.; VMNH NAN0001 • 1 ♂; same collection data as preceding; 16 Oct. 1966; A. Q. Field Trip leg; VMNH NAN0004 • 1 ♂; same collection data as preceding; 14 Apr. 1965; Herp Class leg.; VMNH NAN0008 • Tazewell County, Burkes Garden, Station Spring Creek; 37.0809°N, -81.4145°W; elev. 1219 m; 23 Aug. 1981; R. Hoffman leg.; VMNH NAN0006 • 1 ♂; Tazewell County, Burkes Garden, NW slope Beartown Mountain; 36.9361°N, -81.8861°W; elev. 1341 m; 12 July 1971; R. Hoffman leg.; VMNH NAN0007 • Tazewell County, Burkes Garden; 37.0980°N, -81.3411°W; 20 March 1954; R. Hoffman leg.; VMNH, NAN0009 • 2 ♂♂; same collection data as preceding; 30 March 1954; VMNH NAN0070 • 2 ♂♂; Tazewell County, Burkes Garden, Cassell Farm; 37.0980°N, -81.3411°W; 26 April 1981; R. Hoffman leg.; VMNH NAN0142 • SCAU – West Virginia • 1 ♂; Mercer County, Concord College campus; 37.4245°N, -81.0060°W; 14 Nov. 1966; W. Shear leg.; VMNH NAN0036. For detailed collection data see Suppl. material 7.

Diagnosis

Adult males of Nannaria breweri sp. nov. are distinct from other Nannaria and the nearby N. scutellaria Causey, 1942, based on the following combination of characters: Gonopods. Acropodite curving medially throughout, not sinuous, only curving slightly medially as in N. tennesseensis and not straight, with 90° medial bend at apex as in N. scutellaria. Distal zone with large, lobed lateral flange (Fig. 52A, red arrow), not with small, triangular lateral flange as in N. tennesseensis, or medial flange as in N. scutellaria. Tip directed medially, not cephalically as in N. tennesseensis. Telopodite basal zone height < 1/2 length of acropodite, not > 1/2 as in N. tennesseensis or ca. 1/4 length as in N. scutellaria. Prefemur with medially curving prefemoral process, not straight as in N. tennesseensis. Prefemoral tip bent dorsally (Fig. 52B). Prefemoral spine fused to prefemoral process throughout length, forming ridge along base of prefemoral process (Fig. 52C, red triangle). Color. Tergites with orange paranotal spots (Fig. 53). Brown background. Collum smooth with orange margin.

Figure 52.

Nannaria breweri sp. nov. holotype ♂ (VTEC, MPE02191) left gonopod A anterior view; red arrow indicates lateral flange B medial view C posterior view; red triangle indicates prefemoral spine fused to the prefemoral process. Scale bar: 0.5 mm.

Figure 53.

Nannaria breweri sp. nov. holotype ♂ (VTEC, MPE02191) coloration. Scale bar: 4.0 mm.

Measurements

♂ holotype (VTEC, MPE02191): BL = 29.9, CW = 4.8, IW = 2.3, ISW = 0.9, B11W = 5.4, B11H = 3.6; ♀ paratype (VTEC, MPE02203): BL = 35.9, CW = 4.5, IW = 2.5, ISW = 1.0, B11W = 5.8, B11H = 4.4.

Variation

No known variation.

Distribution

Known from a linear area from southern West Virginia to northeastern Tennessee (Tennessee: Grainger and Hamblen counties; West Virginia: Mercer County; Virginia: Russell and Tazewell counties, Suppl. material 7; Fig. 126). Distribution area: 635 km²; status: MRE.

Ecology

Individuals of N. breweri sp. nov. have been collected from mesic, broadleaved forests composed of pawpaw, maple, spicebush, oak, and elm, often under deciduous leaf litter and/or beneath 1–2 cm dark soil.

Etymology

This species is named for its co-collector and West Virginian, Dr. Michael Brewer (Suppl. material 7). The specific name is a genitive noun derived as a patronym.

Type locality

United States, Tennessee, Hamblen County, Morristown, Panther Creek State Park, in gully below parking lot at Spoone Recreation Area, near Ore Mine Trail, 36.2167°N, -83.4055°W.

Nannaria equalis Chamberlin, 1949

Fig. 54 Vernacular name: “The Knoxville Twisted-Claw Millipede”

Nannaria equalis Chamberlin, 1949: 4, fig. 4. Chamberlin and Hoffman 1958: 40. Hoffman 1999: 366. Marek et al. 2014: 37. Means et al. 2021: S69–S70.

Material examined

United States – Tennessee • 1 ♂; Knox County, site 2 mesic cove hardwoods; from label: 17.233902E 3993447N; 10 May 2005; J. Sevier leg.; VMNH NAN0335. For detailed collection data see Suppl. material 7.

Diagnosis

Adult males of N. equalis are distinct from other Nannaria and the nearby N. monsdomia sp. nov. and N. scutellaria based on the following combination of characters: Gonopods. Gonopodal acropodite straight before apex, not slightly curving before apex as in N. monsdomia sp. nov. Distal zone much reduced, rounded, simple, not with small, lobed lateral flange as in N. monsdomia sp. nov., or bent at 90° with pronounced medial flange as in N. scutellaria. Acropodite with medial swelling, lacking in N. scutellaria. Telopodite basal zone height ca. 1/2 length of acropodite, not > 2/3 length as in N. monsdomia sp. nov. or < 1/3 length as in N. scutellaria. Prefemur with large, straight prefemoral process (Fig. 54A, red arrow), not curving medially, crossing acropodite, and expanding before tip, as in N. monsdomia sp. nov., or curving laterally as in N. scutellaria. Prefemoral spine sharp, curving laterally (Fig. 54B, red triangle), not widely separated from prefemoral process and rounded, as in N. monsdomia sp. nov., or lacking as in N. scutellaria. Acropodite appearing as straight line in medial view, therefore the medial view was not illustrated. Color. Chamberlin reports that the paranota of Nannaria equalis are yellow in color, though this would be unique in the Nannaria and has not been confirmed by the authors. It is likely Chamberlin was referring to a preserved specimen whose color had faded in alcohol.

Figure 54.

Nannaria equalis ♂ (VMNH, NAN0335) left gonopod A anterior view; red arrow indicates large, straight prefemoral process B posterior view; red triangle indicates sharp prefemoral spine. Scale bar: 0.5 mm.

Figure 55.

Nannaria fritzae sp. nov. ♂ holotype (VTEC, MPE02359) left gonopod A anterior view; red arrow indicates tooth-like medial flange B medial view; red triangle indicates small, thorn-like prefemoral spine C posterior view. Scale bar: 0.5 mm.

Measurements

♂ (VMNH, NAN0335): BL = 27.2, CW = 4.0, IW = 1.7, ISW = 0.9, B11W = 4.7, B11H = 2.8.

Variation

No known variation.

Distribution

Known only from the type locality (Tennessee: Knox County; Suppl. material 7; Fig. 126). Distribution area: N/A; status: MRE.

Ecology

Chamberlin (1949) gives no information on the ecology of N. equalis.

Etymology

Chamberlin (1949) provided no etymology for the name equalis, though it is reasonable to assume that it is in reference to the prefemoral process nearly equaling in length the acropodite, as Chamberlin mentions in his description.

Type locality

United States, Tennessee, Knox County, Knoxville.

Notes

In the original publication Chamberlin (1949: 4) designated a male holotype and female paratype (originally as an “allotype”).

Nannaria fritzae sp. nov.

http://zoobank.org/735CBEDE-9F71-4971-9601-BCE0ADF3F53B

Figs 55, 56 Vernacular name: “Kathlyn Fritz’s Twisted-Claw Millipede”

Material examined

Holotype: United States – Georgia • ♂ Chattooga County, hillside by Lake Marvin Rd.; 34.5628°N, -85.0681°W; elev. 385 m; 5 Mar. 2017; hand collected; J. Means, K. Means leg.; VTEC MPE02359.

Paratypes: United States – Georgia • 1 ♂; same collection data as holotype; VTEC MPE02377 • 1 ♂; same collection data as holotype; VMNH MPE02378.

Diagnosis

Adult males of Nannaria fritzae sp. nov. are distinct from other Nannaria and the nearby Nannaria sp. nov. ‘Amicolola’ (wilsoni species group) based on the following combination of characters. Gonopods. Gonopodal acropodite gently curving medially throughout, not straight with abrupt medial bend at apex as in Nannaria sp. nov. ‘Amicolola.’ Distal zone short, rounded, not serpentine as in Nannaria sp. nov. ‘Amicolola.’ Acropodite with small, tooth-like medial flange near apex (Fig. 55A, red arrow). Prefemur with long acicular prefemoral process bent at base and directed medially—not curving laterally, crossing acropodite, as in N. minor or not large, laminate as in Nannaria sp. nov. ‘Amicolola.’ Prefemoral spine reduced to thorn-like structure, arising from prefemoral process, directed ventrally (Fig. 55B, red triangle)—not curving cephalically as in N. minor. Telopodite basal zone ca. 1/2 length of acropodite, not < 1/2 as in N. minor or > 1/4 as in Nannaria sp. nov. ‘Amicolola.’ Color. Tergites with orange paranotal spots (Fig. 56), reduced pigmentation of paranotal spots of terminal segments. Light brown background. Dorsum of collum smooth with orange and white margin.

Figure 56.

Nannaria fritzae sp. nov. ♂ holotype (VTEC, MPE02359) coloration. Scale bar: 4.0 mm.

Measurements

♂ holotype (VTEC, MPE02359): BL = 28.2, CW = 3.8, IW = 2.3, ISW = 0.8, B11W = 4.6, B11H = 3.0.

Variation

No known variation.

Distribution

Known only from the type locality (Georgia: Chattooga County; Suppl. material 7; Fig. 126). Distribution area: N/A; status: MRE.

Ecology

Individuals of N. fritzae sp. nov. were collected on a hillside in a mesic hardwood forest dominated by oak and maple. Specimens were found under leaf litter in a dry creek bed near the base of an oak tree.

Etymology

This species is named for its co-collector, Kathlyn Fritz Means. The name is a genitive noun derived as a matronym.

Type locality

United States, Georgia, Chattooga County, hillside by Lake Marvin Rd.; 34.5628°N, -85.0681°W.

Nannaria monsdomia sp. nov.

http://zoobank.org/B78C1537-424E-42E7-9100-9385966C69D9

Figs 57, 58 Vernacular name: “The House Mountain Twisted-Claw Millipede”

Material examined

Holotype: United States – Tennessee • ♂; Knox County, Mascot, House Mountain State Natural Area, along dried up stream beside Hogskin Rd.; 36.1032°N, -83.7642°W; elev. 357 m; 8 Oct. 2016; hand collected; J. Means, D. Hennen leg. VTEC MPE02188.

Paratypes: United States – Tennessee • 2 ♂♂; same collection data as holotype; VTEC MPE02186, 2199 • 2 ♂; same collection data as holotype; VMNH MPE02201, 3712 • 2 ♀♀; same collection data as holotype; VTEC MPE02187, 2189 • 2 ♀♀; same collection data as holotype; VMNH MPE02196, 2200 • 1 ♂; Knox County, Mascot, House Mountain State Natural Area, under large strips of oak bark on top of soil at top of hill; 36.1043°N, -83.7633°W; elev. 360 m; 8 Oct. 2016; hand collected; J. Means, D. Hennen leg.; VMNH MPE03677 • 2 ♀♀; same collection data as preceding; VMNH MPE02180, 2192 For detailed collection data see Suppl. material 7.

Diagnosis

Adult males of Nannaria monsdomia sp. nov. are distinct from other Nannaria and the nearby N. scutellaria, based on the following combination of characters. Gonopods. Gonopodal acropodite slightly curving medially basal to apex, not straight basal to apex as in N. scutellaria. Distal zone much reduced, rounded, with small, lobed lateral flange (Fig. 57A, red triangle)—not simple as in N. hardeni sp. nov., or bent at 90° with pronounced medial flange as in N. scutellaria. Telopodite basal zone height > 2/3 length of acropodite, not < 1/3 length as in N. scutellaria. Prefemur with large, laminate prefemoral process, curving medially, crossing acropodite and expanding before tip, not straight, acicular as in N. hardeni sp. nov., or curving laterally as in N. scutellaria. Prefemoral process length greater than height of telopodite basal zone, not subequal as in N. hardeni sp. nov. Prefemoral spine widely separated from prefemoral process (Fig. 57A, red arrow) and rounded, not sharp or tooth-like as in N. hardeni sp. nov., or lacking as in N. scutellaria. Color. Tergites with bright orange paranotal spots and faint orange stripes (Fig. 58). Dark brown background. Dorsum of collum smooth with orange margin.

Figure 57.

Nannaria monsdomia sp. nov. holotype ♂ (VTEC, MPE02188) left gonopod A anterior view; red triangle indicates small lateral flange on tip; red arrow indicates wide separation between prefemoral process and rounded prefemoral spine B medial view C posterior view. Scale bar: 0.5 mm.

Figure 58.

Nannaria monsdomia sp. nov. holotype ♂ (VTEC, MPE02188) coloration. Scale bar: 4.0 mm.

Measurements

♂ holotype (VTEC, MPE02188): BL = 30.7, CW = 4.2, IW = 2.2, ISW = 0.9, B11W = 4.2, B11H = 3.1; ♀ paratype (VTEC, MPE02187): BL = 33.5, CW = 4.1, IW = 2.5, ISW = 1.1, B11W = 5.3, B11H = 3.9.

Variation

No known variation.

Distribution

Known only from the type locality, Suppl. material 7; Fig. 126. Distribution area: N/A; status: MRE.

Ecology

Individuals of N. monsdomia sp. nov. were collected from a mesic hardwood forest dominated by beech, maple, magnolia, pawpaw, and poison ivy. All specimens were found under 1–4 cm of hardpacked soil at the edge of boulders near a dried up, rocky stream.

Etymology

The specific epithet is an arbitrary combination of letters from the Latin mons, meaning mountain, and domus, meaning house, which refers to the type locality House Mountain State Natural Area. It is to be treated as a noun in apposition.

Type locality

United States, Tennessee, Knox County, Mascot, House Mountain State Natural Area, along dried up stream beside Hogskin Rd., 36.1032°N, -83.7642°W.

Nannaria tennesseensis (Bollman, 1888)

Figs 59, 60 Vernacular name: “The Tennessee Twisted-Claw Millipede”

Fontaria tennesseensis Bollman, 1888: 340. Bollman 1893: 91. Attems 1898: 263. Attems 1938: 199.

Nannaria tennesseensis: Chamberlin and Hoffman 1958: 42. Hoffman 1999: 368. Marek et al. 2014: 38. Means et al. 2021: S73.

Material examined

Syntype: United States – Tennessee • ♀; Jefferson County, Mossy Creek; C. Branner leg.; NMNH No. 203.

Other material

United States – Tennessee • 1 ♂; Blount County, on dirt rd. off 2422 on Blount-Sevier co. line, 14 E Maryville; 35.7450°N, -83.7188°W; 11 Oct. 1978; R. Shelley, W. Jones leg.; NCSM NAN0484 • 6 ♂♂; Blount County, GSMNP, Hwy 129/Lake Chilhowee Site 2; 35.5472°N, -84.01277°W; 14 Oct. 2006; hand collected; B. Snyder leg.; Bruce Snyder Pers. Coll., B+3, B+9, B+15 • 2 ♂♂ and 1 ♀; same collection data as preceding; 27 October 2006; Bruce Snyder Pers. Coll. • 1 ♂; Jefferson County, Cherokee Dam Campground, roped off road (gravel) next to parking lot by tent camping; 36.1515°N, -83.5170°W; elev. 345 m; 24 May 2016; hand collected; J. Means, D. Hennen leg.; VTEC MPE01237 • 2 ♀♀; same collection data as preceding; VTEC MPE01315, 131 • 1 ♂; Jefferson County, Mossy Creek Wildlife Viewing Area, south end of Cherokee Reservoir, followed road past parking area, turned into a dirt road; 36.1301°N, -83.5056°W; elev. 336 m; 25 May 2016; hand collected; J. Means, D. Hennen leg.; VTEC MPE01245 • 3 ♀♀; same collection data as preceding; VTEC MPE01290–92 • 1 ♂; Jefferson County, on unnumb. rd. off US 411, 16 air mi SE Jefferson City; 35.9543°N, -83.2928°W; 10 Oct. 1978; R. Shelley, W. Jones leg.; NCSM NAN0471 • 1 ♀; Sevier County, Great Smoky Mountains National Park, Ramsey Cascades Trail; 35.7032°N, -83.3536°W; elev. 691 m; 24 Apr. 2019; hand collected; D. Hennen leg.; VTEC MPE04825 • 2 ♂♂; Sevier County, Great Smoky Mountains National Park, forest beside Greenbrier Rd.; 35.7257°N, -83.4020°W; elev. 471 m; 26 Apr. 2019; hand collected; D. Hennen leg.; VTEC MPE04812, 4828 • 3 ♀♀; same collection data as preceding; VTEC MPE04813–15. For detailed collection data see Suppl. material 7.

Diagnosis

Adult males of Nannaria tennesseensis are distinct from other Nannaria and the nearby species N. scutellaria, based on the following combination of characters: Gonopods. Gonopodal acropodite sinuous, curving slightly medially, not curving medially as in N. breweri sp. nov. and not straight with 90° medial bend at apex as in N. scutellaria. Distal zone heavily reduced, with small triangular lateral flange (Fig. 59B, red arrow)—without large lobed lateral flange as in N. breweri sp. nov., or medial flange as in N. scutellaria. Acropodite tip directed cephalically, not medially as in N. breweri sp. nov. or N. scutellaria. Telopodite basal zone long, > 1/2 length of acropodite, not < 1/2 as in N. breweri sp. nov. or ca. 1/4 length as in N. scutellaria. Prefemur with straight prefemoral process, not medially curving as in N. breweri sp. nov. Prefemoral spine fused to prefemoral process throughout length (Fig. 59A, red triangle). Color. Tergites with sherbet orange paranotal spots (Fig. 60). Brown background. Dorsum of collum smooth with orange and white caudal margin.

Figure 59.

Nannaria tennesseensis ♂ (VTEC, MPE01245) left gonopod A anterior view; red triangle indicates prefemoral spine fused to the prefemoral process B medial view; red arrow indicates lateral flange C posterior view Scale bar: 0.5 mm.

Figure 60.

Nannaria tennesseensis ♂ (VTEC, MPE01245) coloration. Scale bar: 4.0 mm.

Measurements

♂ (VTEC, MPE01245): BL = 29.2, CW = 3.2, IW = 1.8, ISW = 0.7, B11W = 4.2, B11H = 2.5; ♀ (VTEC, MPE01291): BL = 34.8, CW = 3.9. IW = 2.5, ISW = 1.2, B11W = 4.7. B11H = 3.5.

Variation

No known variation.

Distribution

Known from eastern Tennessee in the vicinities of Jefferson City and the Great Smoky Mountains National Park (Tennessee: Blount, Jefferson, and Sevier counties; Suppl. material 7; Fig. 126). Distribution area: 1,440 km²; status: SRE.

Ecology

Individuals of N. tennesseensis have been collected from a variety of habitats, including mesic mixed hemlock and hardwood forests primarily composed of oak, maple, tuliptree, hemlock, and rhododendron, and semi-xeric hardwood forests composed of oak and maple. In the former habitats, specimens were often found under leaf litter and/or under 1–2 cm of dark, loose soil; in the latter habitats, specimens were found walking on top of hard packed soil, under thin oak leaf litter.

Etymology

Bollman (1888) gave no etymology for the name tennesseensis, but it is reasonable to assume that the specific name refers to the state in which it is found.

Type locality

United States, Tennessee, Jefferson County, Mossy Creek.

Notes

In the original publication Bollman (1888: 340) inspected two lots of specimens, Nos. 203 & 388. No. 203 contained just one specimen, a female syntype, while No. 388 consisted of five specimens, two males and three females. Bollman (1888: 340) notes that he deposited two of the lot No. 388 specimens at the USNM (now NMNH) though he does not mention the sex of these specimens. Bollman (1888: 340) mentioned two “types”, one from each of the lots. The type from lot No. 388 was deposited in Bollman’s private collection, and its current whereabouts are unknown. Both lots were collected by C. Branner on unspecified dates.

ignis clade

Components. Nannaria ignis sp. nov., N. tenuis sp. nov., and a female from Mercer Co., West Virginia (Fig. 113). Members of the ignis clade share gonopodal characters, including a wide division between the prefemoral process and acropodite, and a small lateral flange near the acropodite apex. Beyond these characters, however, there is a high degree of variability in the shape and curve of the prefemoral process and the width of the trunk. Additionally, the members of the ignis clade occupy a very small geographic range, ~ 500 square km. This suggests that this clade either represents a closely related group of species which have not dispersed widely or one isolated, morphologically variable species which may be in the process of speciating. Based on a combination of morphological and molecular evidence, we feel confident in naming N. ignis sp. nov. and N. tenuis sp. nov. as separate species, however additional sampling, including in and around Mercer Co., West Virginia, will help elucidate relationships within the ignis clade.

Distribution. the ignis clade extends from southeastern West Virginia into southwestern Virginia (Fig. 114).

Nannaria ignis sp. nov.

http://zoobank.org/12052629-9913-403E-BE70-4EB35E719A9E

Figs 61, 62 Vernacular name: “The Dragon-Headed Twisted-Claw Millipede”

Material examined

Holotype: United States – Virginia • ♂; Bland County, base of Big Walker Mountain off powerline access road; 37.0383°N, -81.1090°W; elev. 828 m; 28 Mar. 2016; hand collected; J. Means, P. Marek, A. Prewitt leg.; VTEC MPE01063 •

Paratype: United States – Virginia • 1 ♀; same collection data as holotype; VMNH MPE01064.

Other material

United States – Virginia • 1 ♀; Bland County, 0.5 mi. down road on south side of Little Walker Mtn. near powerline; 37.0273°N, -81.0933°W; elev. 871 m; 6 Jan. 2016; hand collected; J. Means, D. Hennen, R. Jean, P. Marek leg.; VTEC MPE00917 • 2 ♂♂; Bland County, uphill of road, under mixed pine and hardwood ca. 0.3 m from power line on north side of LWM; 37.0302°N, -81.0977°W; elev. 879 m; 8 Jan. 2016; hand collected; J. Means, D. Hennen, P. Marek, R. Jean leg.; VTEC MPE00921, 922 • 1 ♂; Bland County; 37.0324°N, -81.0947°W; elev. 827 m; 21 Nov. 2016; hand collected; J. Means, P. Marek, A. Prewitt leg.; VTEC MPE01037 • 2 ♂♂; Bland County, next to rock face, outcrop going down from access road; 37.0478°N, -81.1155°W; elev. 1184 m; 17 May 2016; hand collected; D. Hennen, P. Shorter, D. Krall, A. Prewitt leg.; VTEC MPE01198, 1202 • 1 ♂; Bland County, Big Walker Mtn., 2 mi E of Sharon Spgs.; 37.0585°N, -81.0988°W; 28 July 1962; R. Hoffman leg.; VMNH NAN0010 • 2 ♂♂; Wythe County; 37.0257°N, -81.0901°W; elev. 838 m; Jan. 2016; hand collected; J. Means, D. Hennen, R. Jean, P. Marek leg.; VTEC MPE00912, 918 • 1 ♂; Wythe County; 37.0315°N, -81.0950°W; elev. 841 m; 21 Mar. 2016; hand collected; J. Means, P. Marek, A. Prewitt leg.; VTEC MPE01036. For detailed collection data see Suppl. material 7.

Diagnosis

Adult males of Nannaria ignis sp. nov. are distinct from other Nannaria and the sympatric N. aenigma based on the following combination of characters: Gonopods. Gonopodal acropodite gently curving medially before apex, not strongly curved as in N. ohionis Loomis & Hoffman, 1948. Distal zone curving dorsomedially (Fig. 61B, red arrow), not medially as in N. ohionis or N. aenigma. Tip with small, rounded lateral flange (Fig. 61A, red triangle),not simple as in N. ohionis, or serpentine as in N. aenigma. Telopodite basal zone reduced, ca. 1/4 length of acropodite, not > 1/3 length as in N. ohionis, or ca. 1/6 length as in N. aenigma. Prefemur with serpentine prefemoral process, bending ventrally before curving cephalolaterally (Fig. 61B), not straight, acicular as in N. ohionis or N. aenigma. Prefemoral process ca. 3/4 length of acropodite, not ca. 1/2 length as in N. aenigma. Prefemoral process arising dorsomedially from large, sharp prefemoral spine, not arising from top of prefemoral spine as in N. ohionis. Color. Tergites with white (Fig. 62A) or orange (Fig. 62B) paranotal spots. Dark brown background. Dorsum of collum smooth with either white or orange margin, depending on color morph.

Figure 61.

Nannaria ignis sp. nov. holotype ♂ (VTEC, MPE01063) left gonopod A anterior view; red triangle indicates small lateral flange B medial view; red arrow indicates dorsomedially curving acropodite tip C posterior view. Scale bar: 0.5 mm.

Figure 62.

Nannaria ignis sp. nov. coloration A non-type ♂ (VTEC, MPE01198) white paranota B non-type ♂ (VTEC, MPE01202) orange paranota. Scale bars: 4.0 mm.

Measurements

♂ holotype (VTEC, MPE01063): BL = 25.3, CW = 3.2, IW = 2.3, ISW = 0.8, B11W = 3.9, B11H = 2.5; ♀ paratype (VMNH, MPE01064): BL = 27.6, CW = 3.6, IW = 1.9, ISW = 0.9, B11W = 4.6, B11H = 3.2.

Variation

Individuals of N. ignis sp. nov. from the ridge of Big Walker Mountain have slightly wider, more laminate prefemoral processes, and a more pronounced lateral flange at the tip of the acropodite.

Distribution

Known only from a small area on the border of Bland and Wythe counties in southwestern Virginia, and on both Big and Little Walker Mountains (Virginia: Bland and Wythe counties; Suppl. material 7; Fig. 126). Distribution area: 3.5 km²; status: MRE.

Ecology

Individuals of N. ignis sp. nov. were collected from mesic broadleaved forests composed of oak, maple, laurel, and some pine.

Etymology

This species is named for the shape of its acropodite in the medial view, which resembles a dragon’s head and neck. The specific name is a noun in apposition derived from the Latin ignis meaning fire.

Type locality

United States, Virginia, Bland County, base of Big Walker Mountain off powerline access road, 37.0383°N, -81.1090°W.

Nannaria tenuis sp. nov.

http://zoobank.org/8280982C-467A-4445-B146-53014CFAB831

Figs 63, 64 Vernacular name: “The Svelte Twisted-Claw Millipede”

Material examined

Holotype: United States – Virginia • ♂; Bland Co., 1.0 km southeast of Bastian; 37.1453°N, -81.1417°W; elev. 728 m; 2 Feb. 2016; hand collected; J. Means, P. Marek, T. Price leg.; VTEC MPE00990.

Paratype: United States – Virginia • ♀; Bland County, 1.6 km southeast of Bastian; 37.1407°N, -81.1394°W; elev. 797 m; 15 Jan. 2016; hand collected; J. Means, P. Marek, V. Wong, T. Price leg.; VMNH MPE00925.

Other material

United States – Virginia • 1 ♂; Bland county, in gully, side of hill, by creek near power line; 37.1537°N, -81.1450°W; elev. 712 m; 8 Feb. 2016; hand collected; J. Means, P. Marek, T. Price, Kyle leg.; VTEC MPE00991 • 3 ♀♀; Bland County; 37.1193°N, -81.1357°W; elev. 874; 1 Apr. 2016; J. Means, P. Marek, A. Prewitt leg.; VTEC MPE01087–89 • 1 ♀; Bland County; 37.1184°N, -81.1360°W; elev. 861 m; 1 Apr. 2016; J. Means, P. Marek, A. Prewitt leg.; VTEC MPE01095 • 1 ♂; Bland County; 37.1159°N, -81.1363°W; elev. 890 m; 1 Apr. 2016; hand collected; J. Means, P. Marek, A. Prewitt leg.; VTEC MPE01086 • 1 ♂; Bland County; 37.1149°N, -81.1347°W; elev. 864 m; 1 Apr. 2016; hand collected; J. Means, P. Marek, A. Prewitt leg.; VTEC MPE01106 • 1 ♀; Bland County; 37.1134°N, -81.1339°W; elev. 842 m; 1 Apr. 2016; hand collected; J. Means, P. Marek, A. Prewitt leg.; VTEC MPE01109 • 1 ♀; Bland County; 37.1200°N, -81.1360°W; elev. 894 m; 7 Apr. 2016; hand collected; J. Means, P. Marek, A. Prewitt, Tyler, Renea leg.; VTEC MPE01115 • 1 ♀; Bland County; 37.1218°N, -81.1356°W; elev. 889 m; 7 Apr. 2016; hand collected; J. Means, P. Marek, A. Prewitt, Tyler, Renea leg.; VTEC MPE01114 • 1 ♂; Bland County; 37.1234°N, -81.1355°W; elev. 889 m; 7 Apr. 2016; hand collected; J. Means, P. Marek, A. Prewitt, Tyler, Renea leg.; VTEC MPE01111 • 2 ♀♀; same collection data as preceding; VTEC MPE01112, 1113 • 1 ♂; Bland County, off AT down from the road; 37.1369°N, -81.1372°W; elev. 939 m; 20 May 2016; hand collected; J. Means, P. Shorter, D. Krall leg.; VTEC MPE01319 • 1 ♀; same collection data as preceding; VTEC MPE01320 • 1 ♂; Bland County, small creek with forest on either side, gully; 37.1449°N, -81.1412°W; elev. 748 m; 20 May 2016; hand collected; J. Means, P. Shorter, D. Krall leg.; VTEC MPE01317 • 1 ♀; same collection data as preceding; VTEC, MPE01318. For detailed collection data see Suppl. material 7.

Diagnosis

Adult males of Nannaria tenuis sp. nov. are distinct from other Nannaria and the sympatric N. aenigma, based on the following combination of characters: Gonopods. Gonopodal acropodite gently curving medially before apex, distal zone curving dorsomedially, with caudally directed tip, not medially directed as in N. ignis sp. nov. Tip rounded with small lateral flange (Fig. 63A, red arrow), not thin and serpentine, lacking flanges as in N. aenigma. Acropodite with medial swelling before apex, not simple as in N. ignis sp. nov. and N. aenigma. Telopodite basal zone thin and elongate, > 1/3 length of acropodite, not ca. 1/4 length of acropodite as in N. ignis sp. nov., or ca. 1/6 length as in N. aenigma. Prefemur with dorsomedially curving prefemoral process, not serpentine, curving ventrally and bending cephalo-laterally at tip as in N. ignis sp. nov., or straight, acicular as in N. aenigma. Prefemoral spine reduced to slight swelling at base of prefemoral process (Fig. 63C, red triangle), not large, projected as in N. ignis sp. nov. Color. Tergites with orange paranotal spots (Fig. 64). Black background. Dorsum of collum smooth with caudal orange margin.

Figure 63.

Nannaria tenuis sp. nov. ♂ holotype (VTEC, MPE00990) left gonopod A anterior view; red arrow indicates small lateral flange B medial view C posterior view; red triangle indicates reduced prefemoral spine. Scale bar: 0.5 mm.

Figure 64.

Nannaria tenuis sp. nov. ♂ non-type (VTEC, MPE00991) coloration. Scale bar: 4.0 mm.

Measurements

♂ holotype (VTEC, MPE00990): BL = 30.6, CW = 4.1, IW = 2.4, ISW = 0.9, B11W = 5.1, B11H = 3.3; ♀ paratype (VMNH, MPE00925): BL = 35.4, CW = 3.9, IW = 2.6, ISW = 1.0, B11W = 5.1, B11H = 3.6.

Variation

No known variation.

Distribution

Known only from southwestern Virginia (Virginia: Bland County; Suppl. material 7; Fig. 126). The distribution of N. tenuis sp. nov. is restricted to the area just southeast of Bastian, Virginia. Distribution area: < 1 km²; status: MRE.

Ecology

Individuals of N. tenuis sp. nov. were collected in winter and spring from mesic broadleaved forests, were found under hardwood leaf litter, and were occasionally beneath 1–2 cm of soil. Two individuals, including the female holotype, were found under ca. 7 cm of snow when temperatures were below freezing, suggesting that there ostensibly exists some cold tolerance in Nannaria.

Etymology

This species is named for its strikingly thin telopodite basal zone. The specific name is an adjective derived from the Latin tenuis, meaning thin.

Type locality

United States, Virginia, Bland Co., 1.0 km southeast of Bastian; 37.1453°N, -81.1417°W.

ohionis clade

Components. Nannaria ohionis, N. sheari sp. nov., N. suprema sp. nov., and female specimens from Stone Mtn. State Park in North Carolina and The Blue Hole, Tennessee (Fig. 114; Supplemental Material 2). Members of the ohionis clade share gonopodal characters, including an acropodite tip with a small lateral flange. The ohionis clade is closely related to the ignis clade, and may represent one, larger group, as evidenced by the low PP (< 0.70) separating the two clades. We expect that further sampling and genetic evidence may prove this group to be polyphyletic, perhaps uniting N. sheari sp. nov. with the closely related ignis clade, or discovering transitional forms between these species.

Distribution. the ohionis clade extends from southeastern Ohio south into West Virginia, southwestern Virginia, northeastern Tennessee, and northwestern North Carolina (Fig. 114).

Nannaria ohionis Loomis & Hoffman, 1948

Figs 65, 66. Vernacular name: “The Southern Ohio Twisted-Claw Millipede”

Fontaria castanea: Williams & Hefner, 1928: 106, fig. 9b

Nannaria ohionis Loomis & Hoffman, 1948: 53. Hoffman 1999: 367. Marek et al. 2014: 37. Means et al. 2021: S71–S72.

Mimuloria ohionis: Causey 1952: 8. Chamberlin and Hoffman 1958: 38.

Material examined

Neotype (here designated): United States – Ohio • ♂; Athens County, Coolville, Hennen Ln.; 39.2107°N, -81.8421°W; elev. 242 m; 25 Nov. 2015; hand collected; D. Hennen leg.; VTEC MPE00906.

Other material

United States – Ohio • 1 ♂; Hocking County, Hocking Hills State Park, Cantwell Cliffs; 39.5420°N, -82.5755°W; elev. 270 m; 22 Oct. 2016; hand collected; D. Hennen leg.; VTEC MPE02270 • 2 ♂♂; same collection data as preceding; 29 Sep. 1963; FAC leg.; VMNH NAN0026 • 2 ♂♂; same collection data as preceding; 15 May 1961; J. Crites leg.; VMNH NAN0326, 327 • 5 ♀; Hocking County, Crane Hollow Nature Preserve, in hollow behind Ellis House; 39.4913°N, -82.5797°W; elev. 290 m; 15 June 2016; hand collected; J. Means, D. Hennen leg.; VTEC MPE01707–11 • 1 ♂; Lawrence County, Pedro, Lake Vesuvius Rec Area, hillside across street from Roadside Group Picnic Shelter; 38.6064°N, -82.6289°W; elev. 195 m; 6 July 2017; hand collected; D. Hennen leg.; VTEC MPE03678 • 1 ♂; Meigs County, Morgan’s Cave, under rocks; 39.0874°N, -81.9971°W; 5 Mar. 1964; L. Carr leg.; VMNH NAN0169 • 1 ♂; Monroe County, Sardis, Narrows Run Rd.; 39.6150°N, -80.9369°W; elev. 219 m; 7 Oct. 2018; L. Hughes leg.; VTEC MPE04611 • 2 ♂♂; Washington County, Little Hocking Nature Trail, next to Little Hocking Elementary School, off Newbury Road; 39.2583°N, -81.7044°W; elev. 195 m; 27 Dec. 2016; hand collected; D. Hennen leg.; VTEC MPE02271, 2273 • 1 ♂; Washington County, Hune Bridge, hillside above parking lot; 39.5092°N, -81.2508°W; elev. 210 m; 19 Aug. 2017; hand collected; D. Hennen, K. Lustofin, M. Spring leg.; VTEC MPE03003 • 1 ♀; same collection data as preceding; VTEC MPE03004 • 2 ♂♂; Washington County, Marietta, Washington County Career Center; 39.4307°N, -81.5003°W; elev. 263 m; 28 Nov. 2015; hand collected; D. Hennen leg.; VTEC MPE00907; SCAU – West Virginia • 1 ♂; Wood County, Waverly, Mountwood Park, 1100 Volcano Rd., north-facing hillside near parking lot; 39.2420°N, -81.2991°W; elev. 292 m; 25 Nov. 2017; hand collected; D. Hennen leg.; VTEC MPE03643. For detailed collection data see Suppl. material 7.

Figure 65.

Nannaria ohionis Loomis & Hoffman, 1948 neotype ♂ (VTEC, MPE00906) left gonopod. A anterior view; red arrow indicates prefemoral process arising from top of prefemoral spine B medial view C posterior view. Scale bar: 0.5 mm.

Diagnosis

Adult males of Nannaria ohionis are distinct from other Nannaria, the sympatric N. terricola, and the nearby N. shenandoa, based on the following combination of characters: Gonopods. Gonopodal acropodite gently curving medially before apex (Fig. 65), not straight or slightly curved as in N. terricola, or strongly curved as in N. shenandoa. Distal zone and tip short, simple, bending medially, not bending dorsally as in N. terricola or large with flanges, curving posterolaterally as in N. shenandoa. Telopodite basal zone height ca. 1/3 length of acropodite, not ca. 1/2 as in N. terricola, or ca. 1/4 as in N. shenandoa. Prefemur with straight, acicular prefemoral process, not curving laterally as in N. shenandoa. Prefemoral process arising from top of prefemoral spine, not from the prefemur as in N. terricola and N. shenandoa (Fig. 65A, red arrow). Prefemoral spine large and projecting, acicular, –not reduced, shelf-like, fused with prefemoral process as in N. terricola. Color. Tergites with orange paranotal spots (Fig. 66). Background variable between individuals, varying from dark brown to black. Dorsum of collum smooth with orange margin.

Figure 66.

Nannaria ohionis Loomis & Hoffman, 1948 neotype ♂ (VTEC, MPE00906) coloration. Scale bar: 4.0 mm.

Measurements

♂ neotype (VTEC, MPE00906): BL = 32.0, CW = 4.0, IW = 2.0, ISW = 0.9, B11W = 4.7, B11H = 3.4; ♀ (VTEC, MPE03004): BL = 32.6, CW = 4.1, IW = 2.4, ISW = 1.0, B11W = 5.2, B11H = 3.9.

Variation

No known variation.

Distribution

Known from southeastern Ohio and northwestern West Virginia (Ohio: Athens, Hocking, Lawrence, Meigs, and Washington counties; West Virginia: Wood County, Suppl. material 7, Fig. 127). Distribution area: 5,823 km²; status: SRE.

Ecology

Individuals of N. ohionis have been collected from mesic broadleaved forests composed of red maple, birch, tuliptree, oak, hickory, pawpaw, American beech, sycamore, buckeye, spicebush, sourwood, and hemlock, often from hillsides and beside walking trails, under leaf litter and logs.

Etymology

Loomis and Hoffman (1948) give no justification for their naming, however it is reasonable to assume this species is named after its type locality, Ohio.

Type locality

United States, Ohio, Athens County, Coolville, Hennen Ln., 39.2107°N, -81.8421°W.

Notes

We designate a neotype for N. ohionis because a holotype was never designated by either Williams and Hefner (1928) or by Loomis and Hoffman (1948). Williams and Hefner (1928) illustrated a millipede found in Ohio and identified it as Fontaria castanea (McNeill, 1887). Loomis and Hoffman (1948) recognized that this illustration was not of F. castanea, but rather a yet-undescribed species of Nannaria, which they named Nannaria ohionis. Chamberlin and Hoffman (1958, p. 38) stated that the “type” was at Miami University; however, searches of the collection by the current curator at our behest revealed no millipede specimens. Causey (1952: 8) mentions and illustrates a “paratype” of N. ohionis loaned to her by Hefner, but does not mention where the “paratype” was held. Because no type material was ever designated for N. ohionis, and due to the lack of any known location of material collected by Williams or Hefner, we felt a neotype designation was warranted.

Nannaria sheari sp. nov.

http://zoobank.org/2A2E7202-5E4E-4915-9533-5FD5235834E8

Figs 67, 68 Vernacular name: “William Shear’s Twisted-Claw Millipede”

Material examined

Holotype: United States – West Virginia • ♂; Mercer County, Brush Creek Preserve, along trail to waterfall; 37.4647°N, -81.0623°W; elev. 628 m; 14 June 2016; hand collected; J. Means, D. Hennen leg.; VTEC MPE01684.

Paratypes: United States – West Virginia • 1 ♀; same collection data as holotype; VMNH MPE01685 • 1 ♂; Mercer County, Brush Creek Falls, Tullgren, sort of Rhododendron litter; 37.4671°N, -81.0597°W; 5 Oct.1967; W. Shear leg.; VMNH NAN0012.

Other material

United States – West Virginia • 1 ♀; Mercer County, Speedway Hemlock Grove, oak-pine litter; 37.4564°N, -81.0105°W; 4 Apr. 1967; W. Shear leg.; VMNH NAN0002 • 2 ♂♂; same collection data as preceding; VMNH NAN0048 • 13 ♂♀; Mercer County, Speedway Roadside Park, route 20, 3 mi. north of Athens, hibernating aggregation of dead animals under log; 37.4646°N, -81.0112°W; 9 Mar. 1968; W. Shear leg.; VMNH NAN0020 • 3 ♂♂; Mercer County, Brush Creek Preserve, along trail to waterfall; 37.4647°N, -81.0623°W; elev. 628 m; 14 June 2016; hand collected; J. Means, D. Hennen leg.; VTEC MPE01665–7 • 1 ♂ and 1 ♀; Mercer County, Athens, Jackson’s Park; 37.4222°N, -81.0163°W; 30 Mar. 1968; W. Shear leg.; VMNH NAN0013 • 1 ♂; Mercer County, Athens, 218 W. Broadway; 37.4254°N, -81.0211°W; 29 Mar. 1973; W. Shear leg.; VMNH NAN0046. For detailed collection data see Suppl. material 7.

Diagnosis

Adult males of Nannaria sheari sp. nov. are distinct from other Nannaria and the nearby N. aenigma, based on the following combination of characters: Gonopods. Gonopodal acropodite continually curving medially, not relatively straight before curving medially at half-way point as in N. castra sp. nov. Distal zone curving medially, not dorsally as in N. castra sp. nov. Acropodite tip directed caudally (Fig. 67B), not medially as in N. aenigma. Acropodite tip with lateral flange (Fig. 67, red arrow). Acropodite simple in N. sheari sp. nov., lacking slight twist and swelling at midpoint as found in N. castra sp. nov., and lateral flange as found in N. aenigma. Telopodite basal zone with slight lateral bulge and wider than space between prefemoral process and acropodite, not thinner as in N. castra sp. nov. Prefemur with dorsomedially curving prefemoral process (Fig. 67B), not straight as in N. aenigma. Prefemoral spine fused with prefemoral process, reduced to shelf-like ridge at base of prefemoral process (Fig. 67C, red triangle). Color. Tergites with orange paranotal spots (Fig. 68). Dark brown background. Dorsum of collum smooth with orange and white margin.

Figure 67.

Nannaria sheari sp. nov. paratype ♂ (VMNH, NAN0012) left gonopod A anterior view; red arrow indicates lateral flange B medial view C posterior view; red triangle indicates reduced, ridge-like prefemoral spine. Scale bar: 0.5 mm.

Figure 68.

Nannaria sheari sp. nov. holotype ♂ (VTEC, MPE01684) coloration. Scale bar: 4.0 mm.

Measurements

♂ holotype (VTEC, MPE01684): BL = 30.7, CW = 3.5, IW = 1.9, ISW = 0.7, B11W = 4.1, B11H = 2.4. ♀ paratype (VMNH, MPE01685): BL = 30.2, CW = 3.2, IW = 1.9, ISW = 0.8, B11W = 4.5, B11H = 3.3.

Variation

No known variation.

Distribution

Known from a small area in and around Athens, West Virginia (West Virginia: Mercer County, Suppl. material 7; Fig. 126). Distribution area: 12 km²; status: MRE.

Ecology

Individuals of N. sheari sp. nov. have been collected from mesic hardwood forests composed of oak, tuliptree, birch, maple, buckeye, rhododendron, and some pine and hemlock. Individuals were found under deciduous leaf litter by the side of hiking trails.

Etymology

This species is named after its original collector, and a longtime mentor to the authors, Dr. William Shear. The specific name is a genitive noun derived as a patronym.

Type locality

United States, West Virginia, Mercer County, Brush Creek Preserve, along trail to waterfall, 37.4647°N, -81.0623°W.

Nannaria suprema sp. nov.

http://zoobank.org/33E2F68E-ACCA-4370-B759-3ECA81A6DDA9

Figs 69, 70 Vernacular name: “The High Elevation Twisted-Claw Millipede”

Material examined

Holotype: United States – Virginia • ♂; Smyth County, Mount Rogers Natural Rec Area, FR 84; 36.70672°N, -81.60284°W; elev. 1318 m; 25 June 2014; hand collected; J. Means, P. Marek and E. Francis leg.; VTEC MPE00075.

Paratypes: United States – Virginia • 1 ♀; same collection data as holotype; VTEC MPE00066 • 1 ♀; same collection data as holotype; VMNH MPE00068.

Other material

United States – Tennessee • 2 ♂♂; Johnson County, 2 NE Shady Valley McQueen Gap Rd., 1 jct. Hwy 133 Harp Mtn.; 36.5580°N, -81.9097°W; May 2002; A. Gagan leg.; VMNH NAN0459; SCAU – Virginia • 1 ♂; Grayson County, Grayson Highlands State Park, Haw Orchard Mountain, Twin Pinnacles trail; 36.6250°N, -81.5020°W; elev. 1530 m; 10 June 2017; hand collected; C. Harden leg.; VTEC MPE03436 • 1 ♂; Grayson County, Grayson Highlands State Pk Haw Orchard Mtn., DF site 1, behind water tank; 36.6251°N, -81.5008°W; 17 Sep. 1990; VMNH survey leg.; VMNH NAN0121 • 6 ♂♂; Grayson County, Grayson Highlands State Park, DF site 1, nr Visitor Center; 36.6242°N, -81.4979°W; 17 Oct. 1990; VMNH survey leg.; VMNH NAN0132 • 1 ♂; same collection data as preceding; 20 May 1991; VMNH NAN0357 • 7 ♂♂; Grayson County, Grayson Highlands State Park DF site 2, below picnic area, Haw Orchard Mountain; 36.6242°N, -81.4979°W; 17 Sep. 1990; VMNH survey leg.; VMNH NAN0122 • 11 ♂♂; same collection data as preceding; 17 Oct. 1990; VMNH NAN0127 • 1 ♂; Grayson County, Grayson Highland State Park, site 1, above water tank, Haw Orchard Mtn.; 36.6250°N, -81.5008°W; 2 June 1991; VMNH survey leg.; VMNH NAN0123 • 1 ♂; same collection data as preceding; 30 Aug. 1990; VMNH NAN0415 • 2 ♂♂; Grayson County, drift fence site 1 Grayson Highlands State Park; 36.6211°N, -81.4845°W; 2 Oct. 1990; VMNH survey leg.; VMNH NAN0413 • 3 ♂♂; Grayson County, Whitetop Mtn., DF site off FS 89, beechwoods; 36.6387°N, -81.6055°W; elev. 1524 m; 7 Sep. 1990; VMNH survey leg.; VMNH NAN0125 • 2 ♂♂; same collection data as preceding; 18 Nov. 1993; VMNH NAN0129 • 6 ♂♂; same collection data as preceding; 11 July 1993; VMNH NAN0131 • 2 ♂♂; same collection data as preceding; 15 Mar. – 23 Apr. 1994; VMNH NAN0139 • 1 ♂; Grayson County, S. slope Mt. Rogers; 36.6597°N, -81.5447°W; elev. 1372 m; 19 Apr. 1970; Karren leg.; VMNH NAN0134 • 1 ♂; same collection data as preceding; 19 May 1957; Highton, R. Hoffman leg.; VMNH NAN0135 • 1 ♂; same collection data as preceding; 27 Sep. 1969; group trip leg.; VMNH NAN0136 • 2 ♂♂; from the border of Johnson and Sullivan counties, top of Holston Mtn., 2 mi. NW of Shady Valley, U.S. Hyw. 421; 36.5394°N, -81.9535°W; 17 May 1974; R. Hoffman leg.; VMNH NAN0171 • 1 ♂; Grayson County, Grayson Highlands State Park at Massie’s Gap; 36.6242°N, -81.4979°W; elev. 1372 m; 23 Aug. 1984; A. Garland leg.; VMNH NAN0179 • 1 ♂; Smyth County, Sugar Grove, Raccoon Branch Wilderness campground, near beginning of Raccoon Branch Trail by campsite 8; 36.7462°N, -81.4247°W; elev. 855 m; hand collected; D. Hennen leg.; VTEC MPE02634 • 1 ♂; Smyth County, Va. Hy. 600, halfway between Konnarock and Elk Garden; 36.7003°N, -81.6109°W; elev. 1219 m; 10 May 1982; R. Hoffman et alia leg; VMNH NAN0128. For detailed collection data see Suppl. material 7.

Diagnosis

Adult males of Nannaria suprema sp. nov. are distinct from other Nannaria and the sympatric N. aenigma, based on the following combination of characters: Gonopods. Acropodite simple and curving medially, without lateral flange as in N. aenigma. Distal zone short, directed dorsomedially (Fig. 69C), with large, triangular lateral flange (Fig. 69A, red arrow)—not serpentine as in N. aenigma. Telopodite basal zone thin with lateral bulge, not simple, straight as in N. ambulatrix sp. nov. and N. aenigma. Telopodite basal zone > 1/2 length of acropodite, not subequal to length as in N. ambulatrix sp. nov., or ca. 1/6 length as in and N. aenigma. Prefemur with serpentine, ventrally curving prefemoral process (Fig. 69B, red triangle), not medially curving as in N. ambulatrix sp. nov., or straight, acicular as in and N. aenigma. Prefemoral spine reduced, sharp. Color. Tergites with orange paranotal spots with lateral white trim (Fig. 70). Individuals variably with a dark brown to black background. Dorsum of collum smooth with orange margin with white trim.

Figure 69.

Nannaria suprema sp. nov. holotype ♂ (VTEC, MPE00075) left gonopod A anterior view; red arrow indicates small, triangular lateral flange B medial view; red triangle indicates ventrally curving prefemoral process C posterior view. Scale bar: 0.5 mm.

Figure 70.

Nannaria suprema sp. nov. holotype ♂ (VTEC, MPE00075) coloration. Scale bar: 4.0 mm.

Measurements

♂ holotype (VTEC, MPE00075): BL = 30.1, CW = 3.4, IW = 2.2, ISW = 0.7, B11W = 4.4, B11H = 2.9; ♀ paratype (VTEC, MPE00066): BL = 31.8, CW = 4.0, IW = 2.2, ISW = 0.9, B11W = 5.3, B11H = 3.6.

Variation

Some individuals from Grayson Highlands, Virginia, display a phenotypic variation wherein the acropodite tip is directed dorsally in the anterior view, rather than medially; though this is extremely rare.

Distribution

Known from a small area in southwestern Virginia and northeastern Tennessee (Tennessee: Johnson and Sullivan counties; Virginia: Grayson and Smyth counties, Suppl. material 7; Fig. 126). Distribution area: 342 km²; status: MRE.

Ecology

Individuals of N. suprema sp. nov. have been found in mesic hardwood forests dominated by oak, maple, rhododendron, hemlock, and red spruce; they are often found under logs and leaf litter.

Etymology

This species is named for its occurrence at high elevations, including the peak of Mount Rogers, the highest point in Virginia. The specific name is derived from the Latin supremus, highest, and is a feminized adjective.

Type locality

United States, Virginia, Smyth County, Mount Rogers Natural Rec Area, FR 84; 36.70672°N, -81.60284°W.

blackmountainensis clade

Components. Nannaria alpina sp. nov., N. blackmountainensis sp. nov. and females from Turkey Foot Campground in Kentucky and Bamboo, North Carolina (Fig. 114, Supplemental Material 2). Members of the blackmountainensis clade share gonopodal characters, including a rectangular gonopod basal zone and a lateral flange on the acropodite apex, which becomes hooked in N. blackmountainensis sp. nov. While the two N. blackmountainensis sp. nov. specimens used in our genetic analysis were both from southeastern Kentucky, the VMNH collection includes specimens from a disparate population near Bamboo, North Carolina. This suggests that the female collected from Bamboo may either represent a third, basal species with similar morphology to N. blackmountainensis sp. nov., or an as-yet unseen species which is sympatric with the North Carolina N. blackmountainensis sp. nov.

Distribution. the blackmountainensis clade extends from eastern Kentucky, into southwestern Virginia, northeastern Tennessee, and northwestern North Carolina (Fig. 114).

Nannaria alpina sp. nov.

http://zoobank.org/D44E0F4B-C192-4706-8445-04E373AC9F66

Figs 71, 72 Vernacular name: “The Alpine Twisted-Claw Millipede”

Material examined

Holotype: United States – Kentucky • ♂; Pulaski County, Boone National Forest, Alpine Recreation Area; 36.9156°N, -84.5182°W; elev. 360 m; 27 Sept. 2017; J. Means, D. Hennen leg.; hand collected; VTEC MPE03150.

Paratypes: United States – Kentucky • 1 ♂; same data as for holotype; VMNH MPE03199 • 3 ♀♀; same data as for holotype; VTEC MPE03200, 3201; VMNH, MPE03768. For detailed collection data see Suppl. material 7.

Diagnosis

Adult males of Nannaria alpina sp. nov. are distinct from other Nannaria based on the following combination of characters. Gonopods. Acropodite straight, curving at 45° angle at apex, not gently curving throughout or with medial swelling as in Nannaria blackmountainensis sp. nov. Acropodite tip with small, triangular lateral flange (Fig. 71A, red arrow), not with large, hooked lateral flange as in N. blackmountainensis sp. nov. Telopodite basal zone with slight lateral bulge (Fig. 71A, red triangle). Prefemur with laterally curving, acuminate prefemoral process; not straight, acicular prefemoral process as in N. blackmountainensis sp. nov. Gap between prefemoral process and acropodite greater than width of acropodite basal zone, not less than width of acropodite basal zone as in N. blackmountainensis sp. nov. Prefemur lacking prefemoral spine. Color. Tergites with pale orange paranotal spots (Fig. 72). White background (most likely due to teneral condition of the specimen). Dorsum of collum smooth with orange margin.

Figure 71.

Nannaria alpina sp. nov. holotype ♂ left gonopod (VTEC, MPE03150) A anterior view; red arrow indicates triangular lateral flange; red triangle indicates lateral bulge B medial view C posterior view. Scale bar: 0.5 mm.

Figure 72.

Nannaria alpina sp. nov. holotype ♂ (VTEC, MPE03150) coloration. Scale bar: 4.0 mm.

Measurements

♂ holotype (VTEC, MPE03150): BL = 29.5, CW = 4.2, IW = 2.0, ISW = 0.9, B11W = 4.9, B11H = 3.1; ♀ paratype (VTEC, MPE03200): BL = 30.8, CW = 3.7, IW = 2.3, ISW = 0.9, B11W = 5.0, B11H = 3.5.

Variation

No known variation.

Distribution

Known only from the type locality, Alpine Recreation Area (Kentucky: Pulaski County; Suppl. material 7; Fig. 126). Distribution area: N/A; status: MRE.

Ecology

Specimens of Nannaria alpina sp. nov. were collected from the side of a hiking path in a mesic broadleaf deciduous forest under ca. 3 cm of soil. Specimens were primarily found within molting chambers from a 1 m² area.

Etymology

This species is named for its type locality. The specific name is a feminine adjective derived from the Latin alpinus for alpine.

Type locality

United States, Kentucky, Pulaski County, Boone National Forest, Alpine Recreation Area, 36.9156°N, -84.5182°W.

Nannaria blackmountainensis sp. nov.

http://zoobank.org/2ACF8697-BA97-4E11-BFBD-233FE0452FAB

Figs 73, 74 Vernacular name: “The Black Mountain Twisted-Claw Millipede”

Material examined

Holotype: United States – Kentucky • ♂; Harlan Co., Black Mountain summit, radar station access road off Black Mountain Ridge Rd., about 2.4 rd. km E jct w/ KY-160; 36.9156°N, -82.8930°W; elev. 1250 m; 10 May 2011; hand collected; P. Marek, C. Hall and D. & M. Beamer leg.; VTEC SPC001090.

Paratypes: United States – Kentucky • 1 ♂; Harlan Co., Stone Mountain State Natural Area, wooded slope above entrance ex US-421; 36.7607°N, -83.1400°W; elev. 570 m; 31 May 2006; hand collected; P. Marek leg.; VMNH SPC001002 • 1 ♀ same collection data as for preceding; VTEC SPC001009 • 1 ♂ Kentucky, Harlan Co., Cumberland Mountain (N Slope), Wagonroad Tunnel Trail; 36.7328°N, -83.2216°W; elev. 691 m; 25 July 2005; hand collected; P. Marek and C. Spruill leg.; FMNH SPC000652.

Other material

United States – Kentucky • 1 ♂; Bell County, Pine Mountain State Park, trail to Honeymoon Falls; 36.7434°N, -83.7121°W; 26 Sept.1976; R. Hoffman leg.; VMNH NAN0184 • 1 ♂; Bell County, Pine Mtn. State Park; 36.6844°N, -83.8351°W; 10 May 1975; J. Ettman leg.; VMNH NAN0493 • 2 ♂♂; Bell County, Pine Mtn. State Park, Wildflower Garden Area; 36.7359°N, -83.7379°W; 1 Feb. 1976; J. Ettman leg.; NCSM NAN0545 • 1 ♀; Harlan County, Cumberland Mtn (N slope), Wagonroad Tunnel Trail (36.7389°N, -83.2197°W, elev. 528 m), 26 July 2005 (Coll: P. Marek, C. Spruill; VTEC SPC000794. SCAU – North Carolina • 1 ♂; Avery County, wooded hillside at Plumetree; 36.0269°N, -82.0080°W; 4 June 1964; L. Hubricht leg.; VMNH NAN0183 • 11 ♂♂♀♀; Avery County, 3.7 SSE Banner Elk, NC 105 & 184, Grandfather Mtn.; 36.1170°N, -81.8386°W; 8 Sept. 1973; R. Shelley; NCSM NAN0498 • 6 ♂♂; same collection data as for preceding; 1978; R. Shelley leg.; NCSM NAN0518 • 20 ♂♂♀♀; Avery County, Newland; 36.0873°N, -81.9273°W; 20 Aug. 1984; D. Massee leg.; NCSM NAN0522 • 1 ♂; Watauga County, Valle Crucis, NC 194, ca. 6S 1113; 36.2092°N, -81.7783°W; 11 Oct. 1975; J. Clamp leg.; NCSM NAN0515; SCAU – Tennessee • 1 ♂; Carter County, 2.5 miles S of Burbank; 36.1123°N, -82.1019°W; 2 May 1951; L. Hubricht leg.; VMNH, NAN0185 • 6 ♂; Carter County, Roan Mtn.; 36.1943°N, -82.0710°W; 9 Aug. 1941; Brooks leg.; VMNH, NAN0186 • 5 ♂♂; Carter County, N slope, Roane Mtn; 36.1079°N, -82.1280°W; 9 Aug. 1941; Dr. & Mrs. Brooks leg.; VMNH, NAN0294 • 1 ♂; Cumberland Mtns.; E. Cope leg.; VMNH, NAN0181; SCAU – Virginia • 1 ♀; Lee County, Poor Valley, Pennington Gap, VA. 621, ca.1.6 km E US. 421, on N facing slope; 36.7768°N, -83.0157°W, elev. 460 m; 10 June 2005; hand collected; P. Marek leg.; VTEC, SPC000540 • 1 ♂; Scott County, Powell Mtn. off FS 642, ca. 1 mi NW of Duffield, JNF; 36.7311°N, -82.8071°W; 13 Sept. 1994; C. Hobson, D. Stevenson leg.; VMNH NAN0182. For detailed collection data see Suppl. material 7.

Diagnosis

Adult males of Nannaria blackmountainensis sp. nov. are distinct from other Nannaria and the nearby N. domestica based on the following combination of characters: Gonopods. Gonopodal acropodite gently curving dorsomedially with pronounced medial swelling (Fig. 73A). Acropodite tip with large, hooked lateral flange (Fig. 73A, red arrow), not small, triangular lateral flange as in N. domestica. Tip terminating in small, dorsally directed rectangular point, not sharp, caudally directed as in N. domestica. Height of telopodite basal zone ca. 1/2 length of prefemoral process, not < 1/3 length as in N. domestica. Prefemoral process straight, acuminate, not laminate and serpentine as in N. domestica. Prefemoral spine reduced and fused to prefemoral process, forming small ridge (Fig. 15B, red triangle). Color. Tergites with hot orange/red paranotal spots (Fig. 74). Jet black background. Dorsum of collum smooth with orange margin.

Figure 73.

Nannaria blackmountainensis sp. nov. holotype ♂ left gonopod (VTEC, SPC001090) A anterior view; red arrow indicates hooked lateral flange B medial view; red triangle indicates reduced prefemoral spine C posterior view. Scale bar: 0.5 mm.

Figure 74.

Nannaria blackmountainensis sp. nov. paratype ♂ (VMNH, SPC001002) coloration. Scale bar: 4.0 mm.

Measurements

♂ holotype (VTEC, SPC001090): BL = 31.2, CW = 3.6, IW = 2.3, ISW = 0.7, B11W = 4.7, B11H = 3.2. ♀ paratype (VTEC, SPC001009): BL = 35.4, CW = 4.1, IW = 2.8, ISW = 1.0, B11W = 5.5, B11H = 3.9.

Variation

There exists a fair amount of variation amongst individuals of N. blackmountainensis sp. nov. from throughout the species’ range. Individuals from the northern border of Tennessee and North Carolina have medially curving prefemoral processes and sharp, pronounced prefemoral spines, while the only known specimen from near Crummies, Kentucky (SPC000652) has a small, triangular prefemoral spine. The southern populations of N. blackmountainensis may be discovered to be a separate species; however, such a discovery is contingent on the further collection of genetic material.

Distribution

Nannaria blackmountainensis sp. nov. has a disjunct distribution in the confluence of eastern Kentucky and Tennessee and western Virginia and North Carolina (Kentucky: Harlan and Bell counties; Virginia: Scott County; Tennessee: Cumberland and Carter counties; North Carolina: Avery and Watauga counties; Suppl. material 7; Fig. 126). Distribution area: 5,582 km²; status: SRE.

Ecology

Individuals of N. blackmountainensis sp. nov. were collected from mesic forests of hemlock, tuliptree, maple, and oak, with an understory of jewelweed and stinging nettle.

Etymology

This species is named for its type locality. The specific name is an adjective.

Type locality

United States, Kentucky, Harlan Co., Black Mountain summit, radar station access road off Black Mountain Ridge Rd., about 2.4 rd. km E jct w/ KY-160, 36.9156°N, -82.8930°W.

terricola clade

Components. Nannaria bobmareki sp. nov., N. dilatata (Hennen & Shelley, 2015), N. fracta sp. nov., N. solenas sp. nov., N. spruilli sp. nov., N. terricola, and females from Little Coal River Campground, Boone County, West Virginia, Crane Hollow, Hocking County, Ohio, and Raven Run, Fayette County, Kentucky (Fig. 114; Suppl. material 2). Members of the terricola clade share gonopodal characters, including straight gonopods which do not cross in situ (with the exception of N. dilatata), the presence of a lateral basal bulge, and a medial flange near the acropodite apex. Nannaria dilatata and the female specimens from Crane Hollow, Ohio, and Raven Run, Kentucky, form a clade which is sister to the rest of the terricola clade, and may represent a separately evolving lineage. The gonopods of N. dilatata are not as straight as in the other members of the terricola clade, and we look forward to the discovery of males from both the Crane Hollow and Raven Run populations. As with the ohionis and several other clades in the minor group, more sampling is needed to resolve the relationships within the terricola clade, but the terricola clade is one of the more morphologically homogenous clades in the minor species group.

Distribution. the terricola clade extends from southeastern Ohio, south into eastern Kentucky, West Virginia, southwestern Virginia, and central Tennessee (Fig. 114).

Nannaria bobmareki sp. nov.

http://zoobank.org/485F1B62-722C-4887-B5D7-E47E34BAFF43

Figs 75, 76, 77 Vernacular name: “Bob Marek’s Twisted-Claw Millipede”

Nannaria ‘Blanton’: Marek and Bond 2006: 721.

Material examined

Holotype : United States – Kentucky • ♂; Leslie County, Cawood Recreation Site, about 5.4 rd km N jct KY-221 & US-421; 36.9364°N, -83.3729°W; elev. 417; 26 July 2006; hand collected; P. Marek and B. Marek leg.; VTEC SPC001019.

Paratypes : United States – Kentucky • 1 ♂; same collection data as for holotype; VMNH SPC001025 • 1 ♂; same collection data as for holotype; FMNH SPC001026 • 1 ♂; same collection data as for holotype; VTEC SPC001027 • 1 ♀; same collection data as for holotype; VTEC SPC001028.

Other material

United States – Kentucky • 6 ♂♂ and 1 ♀; Harlan County Pine Mtn., Blanton Forest State Nature Preserve, High Fork Br., nr campground ranger station; 36.8594°N, -83.3823°W; elev. 411 m; 10 Aug. 2003; hand collected; P. Marek leg.; VTEC SPC000177–183 • 1 ♂; Harlan County, N slope Pine Mtn., James E Bickford Nature Preserve, Pine Mtn. Settlement School; 36.9473°N, -83.1807°W; elev. 587 m; 9 May 2011; hand colleted; P. Marek, C. Hall, D. Beamer, M. Beamer leg.; VTEC SPC001083. For detailed collection data see Suppl. material 7.

Diagnosis

Adult males of Nannaria bobmareki sp. nov. are distinct from other Nannaria and the nearby N. aenigma, based on the following combination of characters: Gonopods. Gonopodal acropodite acicular, bending abruptly medially at 90° at tip, not slightly curving medially before tip as in N. fracta sp. nov. or gently curving medially as in N. aenigma. Distal zone quadrate, not short, rounded, as in N. fracta sp. nov., or long and serpentine as in N. aenigma. Acropodite with small, shelf-like medial flange just before tip (Fig. 75C, red arrow). Prefemur with long, acicular prefemoral process with a pronounced, sharp prefemoral spine, partially fused to prefemoral process forming a ridge (Fig. 75C, red triangle), not free from prefemoral process as in N. fracta sp. nov. or lacking as in N. aenigma. Telopodite basal zone ca. 1/2 length of acropodite, not > 1/2 length of acropodite as in N. fracta sp. nov. or ca. 1/6 length of acropodite as in N. aenigma. Telopodite basal zone with slight medial swelling (Fig. 75A, red circle), not with pronounced medial swelling as in N. fracta sp. nov., or lacking as in N. aenigma. Color: Tergites with white paranotal spots (Fig. 76). Dark brown background. Dorsum of collum smooth with white margin.

Figure 75.

Nannaria bobmareki sp. nov. ♂ holotype left gonopod (VTEC, SPC001019) A anterior view; red circle indicates slight basal swelling B medial view C posterior view; red arrow indicates shelf-like medial flange; red triangle indicates prefemoral spine partially fused with prefemoral process. Scale bar: 0.5 mm.

Figure 76.

Nannaria bobmareki sp. nov. ♂ holotype (VTEC, SPC001019) coloration. Scale bar: 4.0 mm.

Measurements

♂ holotype (VTEC, SPC001019): BL = 26.6, CW = 3.7, IW = 2.1, ISW = 0.9, B11W = 4.3, B11H = 2.8; ♀ paratype (VTEC, SPC001028): BL = 30.4, CW = 4.1, IW = 2.4, ISW = 1.2, B11W = 5.1, B11H = 3.6.

Variation

No known variation.

Distribution

Known from a small triangular area in southeastern Kentucky (Kentucky: Harlan and Leslie counties; Suppl. material 7; Fig. 126). Distribution area: 74 km²; status: MRE.

Ecology

Nannaria bobmareki sp. nov. is the only species of Nannaria which is known to engage in swarming behavior. PEM observed a swarm of an estimated 400 N. bobmareki sp. nov. individuals covering 4 m² in the Blanton Forest State Nature Preserve in August of 2003 (Fig. 77). Nannaria are rarely found in high abundance, making swarming behavior especially notable. Additionally, N. bobmareki sp. nov. was encountered co-occurring with, but not participating in, a putative Müllerian mimicry ring composed of the xystodesmid species Apheloria polychroma Marek, Means & Hennen, 2018 and Brachoria flammipes Marek, 2010 (Marek 2010).

Figure 77.

Small subset of Nannaria bobmareki sp. nov. ♂ swarm observed in the Blanton Forest State Nature Preserve, Harlan Co., Kentucky.

Etymology

This species is named after its co-collector, Bob Marek. The specific name is a genitive noun derived as a patronym.

Type locality

United States, Kentucky, Leslie County, Cawood Recreation Site, about 5.4 rd km N jct KY-221 & US-421, 36.9364°N, -83.3729°W.

Nannaria dilatata (Hennen & Shelley, 2015)

Figs 78, 79 Vernacular name: “The Mossy Rock Twisted-Claw Millipede”

Mimuloria dilatata dilatata Hennen & Shelley, 2015: 1–16, figs 18, 19.

Nannaria dilatata: Means et al. 2021: S69.

Material examined

Holotype : United States – Tennessee • ♂; Marshall County, Henry Horton State Park, campground; [35.5875°N, -86.7035°W]; 9 May 1979; R. M. Shelley leg.; FSCA.

Paratype : United States – Tennessee • 1 ♀; same collection data as holotype; NCSM NCSM27945.

Other material

United States – Tennessee • 1 ♀; Marshall County, Henry Horton State Park, to the right of the main office, under moss on top of large boulder; 35.5914°N, -86.7029°W; 13 May 2017; hand collected; D. Hennen, J. Means, V. Wong leg.; VTEC MPE02788. For detailed collection data see Suppl. material 7.

Diagnosis

Adult males of N. dilatata are distinct from other Nannaria, including the nearby N. hippopotamus sp. nov. and Nannaria sp. nov. ‘Cratagae’ (wilsoni species group) based on the following combination of characters: Gonopods. Gonopodal acropodite gently curving anteromedially, not straight as in N. hippopotamus sp. nov. and Nannaria sp. nov. ‘Cratagae.’ Acropodite tip with prominent triangular lateral flange curving abruptly at a 90° angle towards tip (Fig. 78A, red triangle), not curving gently and rounded as in N. hippopotamus sp. nov. or lacking as in Nannaria sp. nov. ‘Cratagae.’ Acropodite with laminate medial flange just proximal to tip, not lacking as in N. hippopotamus sp. nov. and Nannaria sp. nov. ‘Cratagae.’ Acropodite simple, without medial swelling as in N. hippopotamus sp. nov. Telopodite basal zone ca. ¼ length of acropodite, not ca. ½ as in N. hippopotamus sp. nov. Prefemur with dorsomedially curving prefemoral process, not straight, acicular as in N. hippopotamus sp. nov. and Nannaria sp. nov. ‘Cratagae.’ Prefemoral spine reduced to small rounded lobe at base of prefemoral process (Fig. 78, red arrow), not sharp as in N. hippopotamus sp. nov. or lacking as in Nannaria sp. nov. ‘Cratagae.’ Color. Tergites with light orange paranotal spots and light pink stripes (Fig. 79). Light grey background. Dorsum of collum smooth with light pink margin.

Figure 78.

Nannaria dilatata (Hennen & Shelley, 2015) paratype ♂ (NCSM, NCSM27945) left gonopod A anterior view; red triangle indicates acropodite tip lateral flange B medial view C posterior view; red arrow indicates reduced prefemoral spine. Scale bar: 0.5 mm.

Figure 79.

Nannaria dilatata (Hennen & Shelley, 2015) non-type ♀ (VTEC, MPE02788) coloration. Scale bar: 4.0 mm.

Measurements

♂ holotype (FSCA): BL = 30.9, CW = 4.1, IW = 2.1, ISW = 0.8, B11W = 4.7, B11H = 3.5; ♀ paratype (NCSM, NCSM27945): BL = 28.1, CW = 3.5, IW = 2.0, ISW = 0.9, B11W = 4.5, B11H = 2.9.

Variation

Hennen and Shelley (2015) noted some slight variation between the type locality males and the male collected from Davidson Co., Tennessee, including the latter having a smaller prefemoral spine, a more gradual distal curve of the acropodite, and a reduced acropodite medial swelling.

Distribution

Known only from central Tennessee (Tennessee: Davidson and Marshall counties, Fig. 129). Distribution area: N/A; status: MRE.

Ecology

Hennen and Shelley (2015) provided no ecological notes in their description of N. dilatata, but the single individual collected by DAH for this revision was found under a damp mat of moss on a large boulder.

Etymology

Hennen and Shelley (2015: 14) state “The specific name references the apical dilation on the outer/anterior acropodite surface.”

Type locality

United States, Tennessee, Marshall County, Henry Horton State Park, campground.

Notes

In the original publication, Hennen and Shelley (2015, 14) designated a male holotype (FSCA) and one male and two female paratypes (FSCA, NCSM), all collected by R. M. Shelley on May 9, 1979. Which paratypes were sent to either the FSCA or NCSM was not mentioned in the original publication; however, the NCSM had only one paratype (female, NCSM27945), implying that the other male and female paratypes are deposited at the FSCA.

Nannaria fracta sp. nov.

http://zoobank.org/8D8D39A0-7084-4BA9-8F47-9D0C7C99F55B

Figs 80, 81 Vernacular name: “The Breaks Interstate Park Twisted-Claw Millipede”

Material examined

Holotype : United States – Virginia • ♂; Dickenson County, Haysi, Breaks Interstate Park, Laurel Branch Trail at intersection with Cold Spring Trail; 37.2897°N, -82.2999°W; elev. 565 m; 28 Sep. 2017; hand collected; J. Means, D. Hennen leg.; VTEC MPE031781.

Paratypes : United States – Virginia • ♂; same collection data as holotype; VTEC, MPE03179 • 1 ♂; same collection data as holotype; VMNH MPE03756 • 2 ♂♂; same collection data as holotype; VTEC MPE03183, 84; 1 ♂; Dickenson County, Breaks Interstate Park, in camping area; 37.2936°N, -82.3005°W; 16 Apr. 1983; D. Ogle leg.; VMNH NAN0156 • SCAU – Kentucky • 2 ♀♀; Pike County, Pikeville, Bob Amos Park, 424 Bob Amos Dr. WW Gearheart Hiking Trail; 37.4690°N, -82.5462°W; elev. 359 m; 28 Sep. 2017; hand collected; J. Means, D. Hennen leg.; VTEC MPE03185, 86.

Other material

United States – Virginia • 1 ♂; Dickenson County, Breaks Interstate Park; 37.2936°N, -82.3005°W; 7 Sep. 1967; Neff, R. Hoffman leg.; VMNH NAN0154 • 1 ♂; Russell County, 1 mile NW of Lynn Spring; 37.1153°N, -81.9411°W; 20 Apr. 1962; R. Hoffman leg.; VMNH NAN0155 • 1 ♂; Washington County, Clinch Mountain Wildlife Management Area, deciduous forest above Big Tumbling Creek; 36.9931°N, -81.7368°W; 21 Sep. 2011; S. Roble leg.; VMNH NAN0157. For detailed collection data see Suppl. material 7.

Diagnosis

Adult males of Nannaria fracta sp. nov. are distinct from other Nannaria and the nearby N. aenigma, based on the following combination of characters: Gonopods. Gonopodal acropodite acicular, slightly curving medially before tip, not bending abruptly medially at 90° at tip as in N. bobmareki sp. nov. or gently curving medially as in N. aenigma. Distal zone short, rounded—not quadrate as in N. bobmareki sp. nov., or long and serpentine as in N. aenigma. Acropodite with small, shelf-like medial flange just before tip (Fig. 80C, red arrow). Prefemur with long, acicular prefemoral process with a pronounced, sharp prefemoral spine (Fig. 80C, red triangle), not fused with prefemoral process as in N. bobmareki sp. nov. or lacking as in N. aenigma. Telopodite basal zone > ½ length of acropodite, not ca. 1/2 length of acropodite as in N. bobmareki sp. nov. or ca. 1/6 length of acropodite as in N. aenigma. Telopodite basal zone basal zone with pronounced medial swelling (Fig. 80A, red circle), not with slight medial swelling as in N. bobmareki sp. nov., or lacking as in N. aenigma. Color: Tergites with red paranotal spots (Fig. 81). Dark brown background. Dorsum of collum smooth with red margin.

Figure 80.

Nannaria fracta sp. nov. paratype ♂ (VMNH, NAN0156) left gonopod A anterior view; red circle indicates pronounced basal swelling B medial view C posterior view; red arrow indicates shelf-like medial flange; red triangle indicates sharp, pronounced prefemoral spine. Scale bar: 0.5 mm.

Figure 81.

Nannaria fracta sp. nov. paratype ♂ (VTEC, MPE03185) coloration. Scale bar: 4.0 mm.

Measurements

♂ holotype (VTEC, MPE03178): BL = 28.9, CW = 3.7, IW = 1.8, ISW = 0.85, B11W = 4.4, B11H = 2.5; ♀ paratype (VTEC, MPE03185): BL = 35.0, CW = 4.5, IW = 2.7, ISW = 1.2, B11W = 5.6, B11H = 4.2.

Variation

No known variation.

Distribution

Nannaria fracta sp. nov. has a linear distribution extending from eastern Kentucky into western Virginia (Kentucky: Pike County; Virginia: Dickenson, Russell, and Tazewell counties; Suppl. material 7, Fig. 126). Distribution area: 98 km²; status: MRE.

Ecology

Individuals of N. fracta sp. nov. have been collected from mesic hardwood forests composed of beech, maple, tuliptree, hemlock, and rhododendron. Specimens taken from Bob Amos Park in Kentucky were found under 1–2 cm of hardpacked, dark soil on the side of a hiking path, while specimens collected from Breaks Interstate Park in Virginia were found in a very moist rhododendron cove along the bank of Laurel Branch Creek, under 1–2 cm of sandy, dark soil.

Etymology

This species is named for Breaks Interstate Park, where it was originally collected by R. L. Hoffman in 1962. The specific name is an adjective derived from the Latin fractura, meaning break or fracture.

Type locality

United States, Virginia, Dickenson County, Haysi, Breaks Interstate Park, Laurel Branch Trail at intersection with Cold Spring Trail, 37.2897°N, -82.2999°W.

Nannaria solenas sp. nov.

http://zoobank.org/D0A9A027-576C-4BB4-9B51-0EE1178377A7

Figs 82, 83, 84 Vernacular name: “The Pipestem Twisted-Claw Millipede”

Material examined

Holotype : United States – West Virginia • ♂; Summers County, Pipestem Resort State Park, path to lake, ca. 12 km northeast of Athens; 37.5278°N, -80.9889°W; 835 m; 21 Aug. 2014; hand collected; J. Means, E. Francis leg.; VTEC MPE00128.

Paratypes : United States – West Virginia • 4 ♂♂; same collection data as holotype; VTEC MPE00130, 132–134 • 4 ♂♂; same collection data as holotype; VMNH, MPE00135, 137, 139, 142 • 3 ♀♀; same collection data as holotype; VMNH MPE00129, 131, 136 • 3 ♀♀; same collection data as holotype; VMNH MPE00138, 140, 141.

Other material

United States – Virginia • 2 ♂♂; Hamilton’s Cave, ca. 4 mi. E Mechanicsburg; 37.1726°N, -80.8780°W; 11 Apr. 1967; Herpetology Class leg.; VMNH NAN0151 • 1 ♂; same collection data as preceding; 29 Apr. 1956; R. Hoffman leg.; VMNH NAN0177 • 19 ♂♂; Bland County, Hamilton’s Cave, 5 mi. ENE of Mechanicsburg; 37.1834°N, -80.8624°W; 20 Sep. 1967; Knight, Rushin, Liscombe, R. Hoffman leg.; VMNH NAN0322 • 5 ♂♂; Bland County, hillside outside Hamilton’s Cave, ca. 6 km east of Mechanicsville (presumably a typo of Mechanicsburg); 37.1468°N, -80.8727°W; 16 May 1980; R. Hoffman leg.; VMNH NAN0323 • 10 ♂♂; Bland County, Big Walker Mountain, on 3 mi/SE of Mechanicsville (presumably a typo of Mechanicsburg); 37.1307°N, -80.8916°W; 21 Oct. 1978; R. Hoffman leg.; VMNH NAN0220 • 10 ♂; Russell County, Pinnacles Nature Preserve, nr preserve tr.; 36.9533°N, -82.0550°W; elev. 625 m; 20 Aug. 2006; hand collected; P. Marek, C. Spruill leg.; VTEC MMC0201, 203–207, 209, 212, 214, 215 • 5 ♀♀; same collection data as preceding; VTEC MMC0200, 208, 210, 211, 213 • 1 ♂; Tazewell County, SE slope East River Mtn., near Cove Creek; 37.2060°N, -81.3070°W; 29 Mar. 1971; C. Chapman leg.; VMNH NAN0049 • 3 ♂♂; Tazewell County, Burkes Garden, e. slope of Beartown Mtn.; 37.0114°N, -81.6990°W; elev. 1219 m; 19 Feb. 1971; W. Shear leg.; VMNH NAN0328 • 1 ♂; Wythe County, Crawfish Valley, Channel Rock Hollow trail 1 mile from Strawberry Rd. end; 36.9526°N, -81.3247°W; elev. 772 m; 24 Mar. 2017; hand collected; C. Harden leg.; VTEC MPE02416 • 1 ♂; Wythe County, Crawfish Valley, Channel Rock Hollow trail 1.5 mile from Strawberry Rd. end; 36.9585°N, -81.3189°W; elev. 770 m; 24 Mar. 2017; hand collected; C. Harden leg.; VTEC MPE02428 • SCAU – West Virginia • 1 ♂; Fayette County, Fayette Station; 38.0672°N, -81.0834°W; 4 Oct. 1989; W. Arnold leg.; NCSM NAN0455 • 1 ♂; Mercer County, Athens, Jackson’s Park, Unity Rd.; 37.4267°N, -81.0402°W; 2 Oct. 1966; W. Shear leg.; VMNH NAN0031. For detailed collection data see Suppl. material 7.

Diagnosis

Adult males of Nannaria solenas sp. nov. are distinct from other Nannaria and the sympatric N. asta sp. nov. and N. aenigma, based on the following combination of characters: Gonopods. Gonopodal acropodite straight, not gently curving throughout as in N. asta sp. nov. or N. aenigma. Distal zone short, rectangular, bent medially at 90° angle with acropodite with slight cephalically-directed upturn at terminal edge (Fig. 82A, red arrow), not rounded, directed caudally with lateral flange as in N. asta sp. nov., or sinuous as in N. aenigma. Acropodite simple with slight swelling on inner margin but lacking dimple on outer margin as in N. asta sp. nov., or hooked lateral flange as in N. aenigma. Prefemur with long, acicular prefemoral process, not stout as in N. asta sp. nov., or laterally curving as in N. aenigma. Prefemoral spine pronounced and tooth-like (Fig. 82C, red triangle), not cephalically-curving as in N. asta sp. nov. or lacking as in N. aenigma. Telopodite basal zone simple, ca. 1/2 length of acropodite, not with lateral bulge, < ½ length of acropodite as in N. asta sp. nov., or < 1/6 length of acropodite as in N. aenigma. Color. Tergites with orange paranotal spots (Fig. 83). Black background. Dorsum of collum smooth with orange and white margin.

Figure 82.

Nannaria solenas sp. nov. holotype ♂ (VTEC, MPE00128) left gonopod A anterior view; red arrow upturned acropodite tip B medial view C posterior view; red triangle indicates pronounced prefemoral spine. Scale bar: 0.5 mm.

Figure 83.

Nannaria solenas sp. nov. non-type ♂ (VTEC, MPE02420) coloration. Scale bar: 4.0 mm.

Measurements

♂ holotype (VTEC, MPE00128): BL = 30.8, CW = 4.1, IW = 2.1, ISW = 1.0, B11W = 5.0, B11H = 3.3; ♀ paratype (VMNH, MPE00140): BL = 34.7, CW = 4.3, IW = 2.5, ISW = 1.1, B11W = 5.5, B11H = 3.1.

Variation

Individuals from Crawfish Valley, Wythe County, Virginia, have reduced, shelf-like prefemoral spines (Fig. 84, red triangle).

Figure 84.

Nannaria solenas sp. nov. non-type ♂ (VTEC, MPE02420) from Crawfish Valley left gonopod; red triangle indicates reduced prefemoral spine.

Distribution

Known from southwestern Virginia and southeastern West Virginia, with an individual from central West Virginia (West Virginia: Fayette, Mercer, and Summers counties; Virginia: Bland, Giles, Russell, Tazewell, and Wythe counties, Suppl. material 7; Fig. 126). Distribution area: 3,009 km²; status: SRE.

Ecology

Individuals of Nannaria solenas sp. nov. have been collected from mesic hardwood forests, dominated by oak, maple, and pine. Specimens from Crawfish Valley were collected at night while walking on top of grass in an overgrown road.

Etymology

This species is named after its type locality, Pipestem Resort State Park in West Virginia. The specific name is a noun in apposition derived from the Greek solínas, meaning pipe.

Type locality

United States, West Virginia, Summers County, Pipestem Resort State Park, path to lake, ca. 12 km northeast of Athens, 37.5278°N, -80.9889°W.

Nannaria spruilli sp. nov.

http://zoobank.org/B60DEA74-0FFB-4C8E-AFC0-06769D39AA52

Fig. 85 Vernacular name: “Chad Spruill’s Twisted-Claw Millipede”

Material examined

Holotype : United States – Virginia • ♂; Wise County, Osborn Rock, FR238; 36.8949°N, -82.5902°W; elev. 1112 m; 17 Aug. 2006; hand collected; P. Marek & C. Spruill leg.; VTEC MMC0035.

Paratype : United States – Virginia • 1 ♀; same collection data as holotype; VTEC MMC0021 • 2 ♀♀; same collection data as holotype; VMNH MMC0027, 32. For detailed collection data see Suppl. material 7.

Diagnosis

Adult males of Nannaria spruilli sp. nov. are distinct from other Nannaria and the nearby N. aenigma, based on the following combination of characters: Gonopods. Gonopodal acropodite straight, curving medially at nearly 90° angle at apex, not gently curving or with medial swelling as in N. blackmountainensis sp. nov. Acropodite tip blunt with small, lobed lateral flange (Fig. 85C, red arrow), not with large, hooked lateral flange as in N. blackmountainensis sp. nov., or thin, sinuous tip as in N. aenigma. Height of telopodite basal zone > 1/2 length of acropodite, not < ½ length as in N. blackmountainensis sp. nov., or ca. 1/6 length as in N. aenigma. Prefemur with straight, acicular prefemoral process and reduced prefemoral spine (Fig. 85C, red triangle). Color. Color in life unknown.

Figure 85.

Nannaria spruilli sp. nov. holotype ♂ (VTEC, MMC0035) left gonopod A anterior view B medial view C posterior view; red arrow indicates small lateral flange; red triangle indicates reduced prefemoral spine. Scale bar: 0.5 mm.

Measurements

♂ holotype (VTEC, MMC0035): BL = 29.9, CW = 3.8, IW = 2.1, ISW = 0.8, B11W = 5.1, B11H = 3.1; ♀ paratype (VMNH, MMC0027; head taken for DNA): BL = N/A, CW = 4.1, IW = N/A, ISW = N/A, B11W = 5.4, B11H = 3.7.

Variation

No known variation.

Distribution

Known only from the type locality (Virginia: Wise County, Suppl. material 7; Fig. 126). Distribution area: N/A; status: MRE.

Ecology

Individuals of N. spruilli sp. nov. were collected from mesic broadleaved forests from underneath deciduous leaf litter. Nannaria spruilli sp. nov. was encountered co-occurring with a Mullerian mimicry ring composed of the xystodesmid species Apheloria polychroma, Brachoria cedra Keeton, 1959, and Brachoria insolita Keeton, 1959 (Marek and Bond 2009).

Etymology

This species was named after its co-collector, Chad Spruill. The specific name is a genitive noun derived as a patronym.

Type locality

United States, Virginia, Wise County, Osborn Rock, FR238; 36.8949°N, -82.5902°W.

Nannaria terricola (Williams & Hefner, 1928)

Figs 86, 87 Vernacular name: “The Northern Ohio Twisted-Claw Millipede”

Fontaria terricola Williams & Hefner, 1928: 106, fig. 9c.

Nannaria terricola: Loomis and Hoffman 1948: 53. Chamberlin and Hoffman 1958: 42. Hoffman 1999: 368. Marek et al. 2014: 38. Means et al. 2021: S73.

Material examined

Syntypes : United States – Ohio • 6 ♀♀; labeled as cotypes, Butler County, Oxford, Hueston’s Woods; [39.5800°N, -84.7600°W]; NMNH #2269.

Other material

United States – Ohio • 1 ♂; Adams County, West Union, Edge of Appalachia Preserve, Abner Hollow Trail, not far up the trail, after the uphill walk; 38.7213°N, -83.4335°W; elev. 231 m; 16 June 2016; hand collected; J. Means, D. Hennen leg.; VTEC MPE03714 • 5 ♀♀; same collection data as preceding; VTEC MPE01721–24, 1738 • 9 ♂♂; Butler County, Oxford (39.5069°N, -84.7452°W), 1928, Coll: R. Hefner leg.; VMNH NAN0329 • 3 ♂; Harrison County, Hopedale, in old field; 40.3252°N, -80.9013°W; 29 Apr. 1979; R. Urbanek leg.; VMNH NAN0325 • 2 ♀♀; Hocking County, Crane Hollow Nature Preserve, in hollow behind Ellis House; 39.4913°N, -82.5797°W; elev. 290 m; 15 June 2016; hand collected; J. Means, D. Hennen leg.; VTEC MPE01690, 1691 • 3 ♂♂; Logan County, Bellefontaine, Fred Corker Park; 40.3640°N, -83.7330°W; elev. 435 m; 3 Nov. 2016; hand collected; J. Brown leg.; VTEC MPE02234, 2235, 2238 • 2 ♀♀; same collection data as preceding; VTEC MPE02236, 2237 • 1 ♂; Stark County, Stark Wilderness Center; 40.6720°N, -81.6420°W; 1 May 1971; W. Shear leg.; VMNH NAN0047. For detailed collection data see Suppl. material 7.

Diagnosis

Adult males of Nannaria terricola are distinct from other Nannaria, the sympatric N. ohionis, and the nearby N. shenandoa, based on the following combination of characters: Gonopods. Gonopodal acropodite very slightly curving medially before apex, nearly straight, not strongly curving medially as in N. shenandoa, or obviously curving medially as in N. ohionis. Distal zone and tip short, simple, bent at 90° angle to acropodite and curving dorsally—not bending medially as in N. ohionis, or large, with flanges, curving posterolaterally as in N. shenandoa. Telopodite basal zone ca. 1/2 length of acropodite, not ca. 1/3 as in N. ohionis, or ca. 1/4 as in N. shenandoa. Telopodite basal zone with lateral bulge (Fig. 86A, red arrow), lacking in both N. ohionis and N. shenandoa. Prefemur with straight acicular prefemoral process, not curving laterally as in N. shenandoa. Prefemoral process arising from prefemur, not from top of prefemoral spine as in N. ohionis. Prefemoral spine reduced to small ridge, fused with prefemoral process (Fig. 86B, red triangle), not large, projecting, acicular as in N. ohionis. Color. Tergites with either white or pale orange paranotal spots (Fig. 87). Tan to dark brown background. Dorsum of collum smooth with either white or pale orange caudal margin, depending on color morph.

Figure 86.

Nannaria terricola ♂ (VTEC, MPE02234) left gonopod A anterior view; red arrow indicates basal zone lateral bulge B medial view; red triangle indicates reduced, ridge-like prefemoral spine C posterior view. Scale bar: 0.5 mm.

Figure 87.

Nannaria terricola coloration A ♀ (VTEC, MPE01690) white paranota B ♀ (VTEC, MPE01691) pale orange paranota. Scale bars: 4.0 mm.

Measurements

♀ syntype (NMNH, #2269): BL = 21.7 CW = 3.1, IW = 1.8, ISW = 0.9, B11W = 4.0, B11H = 3.1.

Variation

No known variation.

Distribution

Known from throughout central and southern Ohio (Ohio: Adams, Butler, Harrison, Hocking, Logan, Preble, and Stark counties; Suppl. material 7; Fig. 127). Distribution area: 33,812 km²; status: WRE.

Ecology

Individuals of N. terricola have been collected from mesic hardwood forests composed of sycamore, maple, beech, cherry, hemlock, oak, birch, pawpaw, spicebush, and alder. They were often found under leaf litter and logs on hillsides, occasionally under 1–2 cm of dark soil.

Etymology

Williams and Hefner (1928) gave no explanation for the name terricola in their description of the species, but it is assumed that it is derived from the Latin terricolus, terrestrial.

Type locality

United States, Ohio, Butler County, Oxford, Hueston’s Woods; [39.5800°N, -84.7600°W].

Notes

In the original publication, Williams and Hefner (1928: 106, 107) did not designate type specimens, but mentioned that adults were collected, implying the existence of a type series. Upon investigation of the type material deposited at the NMNH we found that six female syntypes existed (NMNH #2269). Where the male specimen which Williams and Hefner (1928) illustrated and described currently resides is unknown.

ambulatrix clade

Components. Nannaria ambulatrix sp. nov., N. asta sp. nov., N. botrydium sp. nov., N. tsuga, sp. nov., and female specimens from Warriors Path State Park, Tennessee and Brumley Gap, Virginia (Fig. 114, Suppl. material 2). Members of the ambulatrix clade share gonopodal characters, including a thin gonopod with a straight and tall basal zone, a prominent prefemoral spine, and a lobed or triangular lateral flange near the acropodite apex. For the minor species group, the ambulatrix clade is uniquely cohesive morphologically, genetically (average PP = 0.95) and geographically.

Distribution. the ambulatrix clade extends from southwestern Virginia into northeastern Tennessee (Fig. 114).

Nannaria ambulatrix sp. nov.

http://zoobank.org/F4DFC180-0793-4435-9E2A-31BD35809325

Figs 88, 89 Vernacular name: “The Big Walker Twisted-Claw Millipede”

Material examined

Holotype: United States – Virginia • ♂; Smyth County, west slope of Big Walker Mtn., VA-16; 36.9112°N, -81.5317°W; elev. 1054 m; 9 Sept. 2014, J. Means leg.; hand collected; VTEC MPE00178.

Paratypes: United States – Virginia • 1 ♀; same data as for holotype; VTEC, MPE00174 • 2 ♂; same data as for holotype; VTEC MPE00166, 167 • 2 ♂ and 1 ♀; same data as for holotype; VMNH, MPE00169, 179, 180.

Other material

United States – Virginia • 1 ♂ Washington County, Mendota, steep hillside beside intersection of CR 614 and CR 620, near Holston River; 36.7146°N, -82.2806°W; elev. 381 m; 12 Feb. 2018; hand collected J. Means, D. Hennen, P. Marek leg.; VTEC MPE03789 • 1 ♂; Washington County, Mendota, Anderson Rd.; 36.7219°N, -82.2771°W; elev. 464 m; 11 Feb. 2017; J. Means, D. Hennen, V. Wong leg.; hand collected; VTEC MPE02286 • 1 ♂ Washington County, Mendota, VA-620, ca.1 road km NE jct w/VA-614; 36.7192°N, -82.2775°W; elev. 287 m; 12 Feb. 2018; J. Means, D. Hennen, P. Marek leg.; hand collected; VTEC MPE04240 • 3 ♀; same collection data as for preceding; VTEC MPE03797, 3799, 4270 • 5 ♂♂; same collection data as for preceding; VTEC MPE03794, 3800-3803 • 2 ♀♀; Washington County, Mendota, Fugate Gap; 36.7285°N, -82.3026°W; elev. 609 m; 12 Feb. 2018; hand collected; J. Means, D. Hennen, P. Marek leg.; VTEC MPE03795, 3804 • 1 ♂ Washington County, DF site off VA. 620, 1 km E of Mendota; 36.7108°N, -82.2904°W; 11 June 1998; VMNH Survey leg.; VMNH NAN0137 • 1 ♂; Washington County, DF site off VA. 620, ca. 2 mi NE of Mendota; 36.7313°N, -82.2760°W; 8 Aug.–2 Dec. 1997;VMNH Survey leg.; VMNH NAN0307 • 1 ♂; Washington County, Abram’s Fall, ca. 2 mi SW of Benhams, end of VA. 614; 36.6580°N, -82.2439°W; 15 June 1960; R. Hoffman leg.; VMNH NAN0308 • 1 ♂; Smyth County, Hungry Mother State Park, inside Hemlock Haven conference center; 36.8914°N, -81.5249°W; 5 Aug. 2009; J. Beard leg.; VMNH NAN0124 • 1 ♂; Smyth County, Big Walker Mountain, ca. 3 mi west of Hungry Mother State Park on VA. 16, west slope; 36.9104°N, -81.5333°W; 9 Sept. 1956; R. Hoffman leg.; VMNH NAN0310. For detailed collection data see Suppl. material 7.

Diagnosis

Adult males of Nannaria ambulatrix sp. nov. are distinct from other Nannaria and the co-occurring N. aenigma, based on the following combination of characters: Gonopods. Gonopodal acropodite very slightly curving medially before apex, nearly straight (Fig. 88). Distal zone short, directed dorsomedially, with large, triangular lateral flange (Fig. 88A, red arrow), not simple as in N. terricola, or serpentine as in N. aenigma. Telopodite basal zone large, subequal to length of acropodite, not ca. 1/2 length as in N. terricola, or ca. 1/6 length as in N. aenigma. Telopodite basal zone straight, without lateral bulge as in N. terricola. Prefemur with medially curving prefemoral process, not acicular, straight as in N. terricola and N. aenigma. Prefemoral process arising dorsomedially from small, sharp, prefemoral spine (Fig. 88A, red triangle), not arising from prefemur or with spine reduced to a shelf-like ridge as in N. terricola. Color. Tergites with either white/light yellow or orange paranotal spots (Fig. 89). Gray to black background. Dorsum of collum smooth with either white/light yellow or orange margin, depending on color morph.

Figure 88.

Nannaria ambulatrix sp. nov. holotype ♂ left gonopod (MPE00178) A anterior view; red arrow indicates large, triangular lateral flange; red triangle indicates prefemoral process arising dorsomedially from prefemoral spine B medial view C posterior view. Scale bar: 0.5 mm.

Figure 89.

Nannaria ambulatrix sp. nov. coloration A non-type ♀ (VTEC, MPE03795) white/tan paranota B paratype ♂ (VTEC, MPE00167) orange paranota. Scale bars: 4.0 mm.

Measurements

♂ holotype (VTEC, MPE00178): BL = 28.0, CW = 4.3, IW = 2.2, ISW = 1.0, B11W = 5.0, B11H = 3.4; ♀ paratype (VTEC, MPE00174): BL = 32.3, CW = 4.3, IW = 2.5, ISW = 0.9, B11W = 5.6, B11H = 3.7.

Variation

Specimens of Nannaria ambulatrix sp. nov. collected from the top of Clinch Mountain (36.7285°N, -82.3026°W, Elev. 845 m) were noticeably smaller than those collected from other localities, suggesting a possible adaptation in the higher elevation population to a drier and/or less suitable habitat. This is the only instance of a stark difference in size between populations of the same species that we have observed in Nannaria.

Distribution

Known only from southwestern Virginia (Russell, Smyth and Washington counties; Suppl. material 7; Fig. 126). The distribution of N. ambulatrix sp. nov. follows the Brumley and Little Brushy Mountain ranges. Distribution area: 488 km²; status: MRE.

Ecology

Nannaria ambulatrix sp. nov. individuals were collected from mesic broadleaf deciduous forests, often in groups occupying 1–2 m² patches, suggesting that the species is distributed in a highly aggregated manner.

Etymology

This species is named for its type locality (Big Walker Mountain), and was informally proposed by R.L. Hoffman. The specific name is a noun in apposition derived from the Latin ambulo, to walk, and trix, ‘a female doer of an action.’

Type locality

United States, Virginia, Smyth County, west slope of Big Walker Mtn., VA-16, 36.9112°N, -81.5317°W.

Nannaria asta sp. nov.

http://zoobank.org/79F53717-C02F-4A97-825F-F2F7F5C8D474

Figs 90, 91 Vernacular name: “The Crawfish Valley Twisted-Claw Millipede”

Material examined

Holotype: United States – Virginia • ♂; Wythe County, Crawfish Valley, Channel Rock, ca. 1.5 mi down trail from Strawberry Rd. end; 36.9585°N, -81.3189°W; elev. 770 m; 24 Mar. 2017; hand collected; C. Harden leg.; VTEC MPE02419.

Paratypes: United States – Virginia • 3 ♂♂; Wythe Co. Crawfish Valley; 36.9811°N, -81.2934°W; elev. 715 m; 24 Mar. 2017; C. Harden leg.; VTEC MPE02421, VMNH, MPE02429, 2430. For detailed collection data see Suppl. material 7.

Diagnosis

Adult males of Nannaria asta sp. nov. are distinct from other Nannaria and the sympatric N. solenas sp. nov. and N. aenigma based on the following combination of characters: Gonopods. Gonopodal acropodite gently curving medially, not straight as in N. solenas sp. nov. Distal zone curving dorsomedially, tip rounded, directed caudally with small lateral flange (Fig. 90A), not rectangular, directed medially at 90° angle with acropodite and slight cephalically-directed upturn at terminal edge as in N. solenas sp. nov., and not sinuous, without flange as in N. aenigma. Acropodite with slight swelling on inner margin (Fig. 90B, red triangle) and dimple on outer margin (Fig. 90A, red arrow). Prefemur with stout, acicular prefemoral process, arising dorsomedially from projected, cephalically-curving prefemoral spine, not arising from prefemur as in N. solenas sp. nov. Telopodite basal zone with small lateral bulge, lacking in N. solenas sp. nov. Color. Tergites with orange paranotal spots (Fig. 91). Black background. Dorsum of collum smooth with orange margin.

Figure 90.

Nannaria asta sp. nov. holotype ♂ left gonopod (VTEC, MPE02419) A anterior view; red arrow indicates outer margin dimple B medial view; red triangle indicates slightly swollen inner margin C posterior view. Scale bar: 0.5 mm.

Figure 91.

Nannaria asta sp. nov. holotype ♂ (VTEC, MPE02419) coloration. Scale bar: 4.0 mm.

Measurements

♂ holotype (VTEC, MPE02419): BL = 26.2, CW = 4.1, IW = 2.1, ISW = 0.9, B11W = 4.9, B11H = 3.1.

Variation

No known variation.

Distribution

Known only from southwestern Virginia (Wythe and Bland counties; Suppl. material 7; Fig. 126). The distribution of N. asta sp. nov. is restricted to the Crawfish Valley area and south of Wytheville, Virginia. Distribution area: 171 km²; status: MRE.

Ecology

The majority of Nannaria asta sp. nov. individuals were found at night, walking along the side of a path on top of predominantly pine litter, a notably odd behavior for Nannaria, which typically remain beneath the leaf litter and are more cryptic in their behavior.

Etymology

The specific epithet is an arbitrary combination of letters derived from the Infraorder Astacidea (which includes crayfish). It is to be treated as a noun in apposition.

Type locality

United States, Virginia, Wythe County, Crawfish Valley, Channel Rock, ca. 1.5 mi down trail from Strawberry Rd. end, 36.9585°N, -81.3189°W.

Nannaria botrydium sp. nov.

http://zoobank.org/AA57E2AC-54E7-4B80-B9E0-44EE3F1038EC

Figs 92, 93 Vernacular name: “The Grapefield Road Twisted-Claw Millipede”

Material examined

Holotype: United States – Virginia • ♂; Bland County, 2.6 km west of Hicksville; 37.1892°N, -81.1633°W; elev. 738 m; 12 Feb. 2016; hand collected; D. Hennen, P. Marek leg.; VTEC MPE00993.

Paratypes: United States – Virginia • 1 ♂; Bland County, 2.7 km west of Hicksville; 37.1913°N, -81.165°W; elev. 754 m; 12 Feb. 2016; hand collected; D. Hennen, P. Marek leg.; VMNH MPE00992 • 1 ♀; Bland County, 4.2 km northwest of Hicksville; 37.19314°N, -81.18221°W; elev. 900 m; 2 Mar. 2016; hand collected; J. Means, P. Marek leg.; VTEC MPE01014.

Other material

United States – Virginia • 1 ♂; Bland County; 37.1940°N, -81.1733°W, elev. 823 m; 2 Mar. 2016; hand collected; J. Means, P. Marek, Tyler leg.; VTEC MPE01012 • 1 ♂; Bland County; 37.1937°N, -81.1739°W; elev. 815 m; 2 Mar. 2016; hand collected; J. Means, P. Marek, Tyler leg.; VTEC MPE01013 • 2 ♂♂; Bland County; 37.1930°N, -81.1827°W; elev. 905 m; 2 Mar. 2016; hand collected; J. Means, P. Marek, Tyler leg.; VTEC MPE01009, 1011 • 1 ♂; Bland County; 37.1830°N, -81.1607°W; elev. 675 m; 7 Mar. 2016; hand collected; P. Marek, V. Wong, Tyler leg.; VTEC MPE01016 • 1 ♂ and 1 ♀; Bland County; 37.1234°N, -81.1355°W; elev. 889 m; 7 Apr. 2016; hand collected; J. Means, P. Marek, A. Prewitt, Tyler, Renea leg.; VTEC MPE03721, 3722 • 4 ♂; Tazewell County, Summit of East River Mtn., above Bluefield, West Virginia; 37.2112°N, -81.3164°W; elev. 914–1067 m; 9 Sep.1972; W. Shear leg.; VMNH NAN0015 • 2 ♂♂; Tazewell County, SE slope of East River Mountain near Cove Creek; 37.2067°N, -81.3072°W; 29 Mar. 1973; C. Chapman leg.; VMNH NAN0043; SCAU – West Virginia • 1 ♂; Mercer County, just over the West Virginia state line on 460, ca. 2 mi. SW of Glen Lyn on 219/8; 37.3452°N, -80.9105°W; elev. 570 m; 12 Nov. 2017; hand collected; J. Means; VTEC MPE03925 • 1 ♂; Randolph County, Cheat Bridge, TNC Preserve, Showers Fk at US 250; 38.6166°N, -79.8707°W; elev. 1174 m; 8 Oct. 2000; W. Arnold leg.; VMNH NAN0081. For detailed collection data see Suppl. material 7.

Diagnosis

Adult males of Nannaria botrydium sp. nov. are distinct from other Nannaria and the sympatric N. aenigma, based on the following combination of characters: Gonopods. Gonopodal acropodite gently curving medially before apex, with lateral flange on tip, not thin and undulating as in N. aenigma. Acropodite tip directed caudally. Acropodite without medial swelling as in N. tenuis sp. nov. Telopodite basal zone > 1/3 length of acropodite, not ca. 1/6 as in N. aenigma. Prefemur with long, thin prefemoral process, bent 90° and directed medially at half-way point (Fig. 92A, red arrow)—not nearly linear as in N. tenuis sp. nov., or straight as in N. aenigma. Prefemoral spine reduced and fused for entire length with prefemoral process (Fig. 92C, red triangle), not lacking as in N. aenigma and N. tenuis sp. nov. Color. Tergites with bright red paranotal spots (Fig. 93). Black background. Dorsum of collum smooth with red margin.

Figure 92.

Nannaria botrydium sp. nov. holotype ♂ (VTEC, MPE00993) left gonopod A anterior view; red arrow indicates 90° bend in prefemoral process B medial view C posterior view; red triangle indicates reduced, prefemoral spine fused with prefemoral process. Scale bar: 0.5 mm.

Figure 93.

Nannaria botrydium sp. nov. holotype ♂ (VTEC, MPE00993) coloration. Scale bar: 4.0 mm.

Measurements

♂ holotype (VTEC, MPE00993): BL = 27.0, CW = 3.8, IW = 1.9, ISW = 0.8, B11W = 4.7, B11H = 3.0; ♀ paratype (VTEC, MPE01014): BL = 26.7, CW = 3.2, IW = 1.9, ISW = 0.9, B11W = 3.9, B11H = 2.8.

Variation

No known variation.

Distribution

Known only from a small area (< 10 km²) west of Hicksville, Virginia (Virginia: Bland and Tazewell counties; Suppl. material 5; Fig. 126. A single male is recorded from Randolph County, West Virginia (ca. 200 km away from type locality), but as this is such a disjunct record, the authors are treating this specimen as an N. botrydium sp. nov. with an incorrect locality label (marked with an asterisk in Fig. 127). Distribution area: 10 km²; status: MRE.

Ecology

Individuals of N. botrydium sp. nov. were collected from mesic hardwood forests composed of oak, maple, and rhododendron, found under leaf litter and occasionally 1–2 cm in dark soil.

Etymology

This species is named after Grapefield Road, which runs near the type locality. The specific name is a Latinized diminutive of the Greek botrys, meaning a cluster of grapes.

Type locality

United States, Virginia, Bland County, 2.6 km west of Hicksville, 37.1892°N, -81.1633°W.

Nannaria tsuga sp. nov.

http://zoobank.org/96611C3A-FC68-48FB-8EA4-9B241C3C3D53

Figs 94, 95 Vernacular name: “The Hemlock Grove Twisted-Claw Millipede”

Material examined

Holotype: United States – Tennessee • ♂; Sullivan County, Bristol, Steele Creek Park, along Hemlock Hollow Trail; 36.5703°N, -82.2356°W; elev. 506 m; 11 June 2018; hand collected; D. Hennen leg.; VTEC MPE04047.

Paratypes: United States – Tennessee • 2 ♂♂; same collection data as holotype; VTEC MPE04048, 4049 • 1 ♂; same collection data as holotype; VMNH MPE04311 • 1 ♀; same collection data as holotype; VTEC MPE04043 • 1 ♀; same collection data as holotype; VMNH MPE04050. For detailed collection data see Suppl. material 7.

Diagnosis

Adult males of Nannaria tsuga sp. nov. are distinct from other Nannaria and the nearby N. aenigma, based on the following combination of characters: Gonopods. Gonopodal acropodite gently curving medially before apex, not nearly straight as in N. ambulatrix sp. nov. Distal zone short, directed dorsomedially, with large, lobed lateral flange (Fig. 94A, red arrow), not with triangular lateral flange as in N. ambulatrix sp. nov., or without flange and serpentine as in N. aenigma. Telopodite basal zone height < 1/2 length of acropodite, not enlarged, ca. 1/2 length as in N. ambulatrix sp. nov., or reduced, ca. 1/6 length as in N. aenigma. Prefemur with straight, laminate prefemoral process, separated widely from projected, with blunt prefemoral spine (Fig. 94A, red triangle), prefemoral process not arising dorsomedially from small, sharp prefemoral spine as in N. ambulatrix sp. nov. Color. Tergites with orange paranotal spots (Fig. 95). Black background. Dorsum of collum smooth with orange caudal margin.

Figure 94.

Nannaria tsuga sp. nov. holotype ♂ (VTEC, MPE04047) left gonopod A anterior view; red arrow indicates large, lobed lateral flange; red triangle indicates prefemoral process arising distally from prefemoral spine B medial view C posterior view. Scale bar: 0.5 mm.

Figure 95.

Nannaria tsuga sp. nov. holotype ♂ (VTEC, MPE04047) coloration. Scale bar: 4.0 mm.

Measurements

♂ holotype (VTEC, MPE04047): BL = 33.9, CW = 4.3, IW = 2.0, ISW = 1.0, B11W = 5.2, B11H = 3.0; ♀ paratype (VTEC, MPE04043): BL = 31.1, CW = 3.8, IW = 2.6, ISW – 1.1, B11W = 4.9, B11H = 3.8.

Variation

No known variation.

Distribution

Known only from the type locality (Tennessee: Sullivan County, Suppl. material 7; Fig. 126). Distribution area: N/A; status: MRE.

Ecology

Individuals of Nannaria tsuga sp. nov. were collected from a mesic mixed hardwood and hemlock forest, composed of, in addition to hemlock, oak, beech, and buckeye. Individuals were found under moist litter on a hillside.

Etymology

This species is named for the habitat in which it was discovered, a hemlock grove. The specific name is derived from the name Tsuga Carrière, the genus containing hemlock trees, and is a noun in apposition.

Type locality

United States, Tennessee, Sullivan County, Bristol, Steele Creek Park, along Hemlock Hollow Trail, 36.5703°N, -82.2356°W.

minor clade

Components. Nannaria cryomaia sp. nov., N. daptria sp. nov., N. hippopotamus sp. nov., N. honeytreetrailensis sp. nov., N. kassoni sp. nov., N. minor, N. rhysodesmoides (Hennen & Shelley, 2015), and multiple female specimens from northeastern Tennessee (Fig. 114; Suppl. material 2). The minor clade is fairly morphologically diverse, though all members have either a small medial flange near the acropodite tip, or a medial swelling before the acropodite apex. No specimens of N. rhysodesmoides were collected for this study; based on the close geographic proximity of N. rhysodesmoides to N. cryomaia sp. nov. and N. kassoni sp. nov., as well as the shared gonopodal characters of a sharp acropodite tip and the presence of a lateral flange on the acropodite apex (as seen in N. kassoni sp. nov.), we place N. rhysodesmoides within the minor clade. We had expected N. kassoni sp. nov., and N. cryomaia sp. nov., to form a clade, due to their close geographic proximity, and similarly expanded medial flange; however, they appear to be more closely related to other, less morphologically similar members of the minor clade. Nannaria cryomaia sp. nov. is found only ~ 30 km from N. kassoni sp. nov., and yet is ~ 170 km away from N. minor, to which it is more closely related. Phylogeographic analyses may reveal that there have been multiple distributional expansions and retractions of the minor clade, leading to the interesting patterns of relatedness observed in this group.

Distribution. the N. minor clade extends from southwestern Virginia into northeastern Tennessee (Fig. 114).

Nannaria cryomaia sp. nov.

http://zoobank.org/B5C5C054-0EAE-48AB-8422-CC516979A45E

Figs 96, 97 Vernacular name: “The Frozen Head Twisted-Claw Millipede”

Material examined

Holotype: United States – Tennessee • ♂; Morgan County, campground at Frozen Head State Park; 36.1321°N, -84.4978°W; elev. 423 m; 12 May 2017; hand collected; J. Means, D. Hennen, V. Wong leg.; VTEC MPE02642.

Paratypes: United States – Tennessee • 1 ♂; same collection data as holotype; VMNH MPE03765 • 2 ♀; same collection data as holotype; VTEC MPE02793, 94 • 2 ♀; same colletion data as holotype; VMNH MPE02795, 96.

Other material

United States – Tennessee • 1 ♂; Morgan County, Frozen Head State Park; 36.1321°N, -84.4978°W; 29 May 1980; R. Shelley, MSM leg.; NCSM NAN0452. For detailed collection data see Suppl. material 7.

Diagnosis

Adult males of Nannaria cryomaia sp. nov. are distinct from other Nannaria and the nearby Nannaria sp. nov. ‘Cratagae’ (wilsoni species group) based on the following combination of characters: Gonopods. Gonopodal acropodite curving medially before apex, not straight as in Nannaria sp. nov. ‘Cratagae.’ Apex with distinct constriction (Fig. 96A, red arrow), tip expanded distally, with small lobed lateral and medial flanges, not with acuminate, triangular lateral flange as in N. kassoni sp. nov. Apex without ventrally directed bend as in Nannaria sp. nov. ‘Cratagae.’ Acropodite with expanded medial flange (Fig. 96A, red triangle). Telopodite basal zone height > 1/3 length of acropodite, not ca. 1/3 as in N. kassoni sp. nov., and not ca. 1/4 length as in Nannaria sp. nov. ‘Cratagae.’ Prefemur with medially curving prefemoral process, paralleling curve of acropodite, not straight, acicular as in Nannaria sp. nov. ‘Cratagae.’ Prefemur with short, triangular prefemoral spine, not pronounced, curving as in N. kassoni sp. nov. Color. Tergites with faint orange stripes (Fig. 97). Dark brown background. Dorsum of collum smooth with orange margin.

Figure 96.

Nannaria cryomaia sp. nov. holotype ♂ (VTEC, MPE02642) left gonopod A anterior view; red arrow indicates apical constriction; red triangle indicates medial flange B medial view C posterior view. Scale bar: 0.5 mm.

Figure 97.

Nannaria cryomaia sp. nov. paratype ♀ (VTEC, MPE02793) coloration. Scale bar: 4.0 mm.

Measurements

♂ holotype (VTEC, MPE02642): BL = 24.9, CW = 3.9, IW = 2.4, ISW = 0.9, B11W = 4.7, B11H = 3.2; ♀ paratype (VMNH, MPE02796): BL = 26.6, CW = 3.9, IW = 2.2, ISW = 1.0, B11W = 5.2, B11H = 3.7.

Variation

No known variation.

Distribution

Known only from the type locality (Tennessee: Morgan County; Suppl. material 7; Fig. 126). Distribution area: N/A; status: MRE.

Ecology

Individuals of Nannaria cryomaia sp. nov. were collected from a mesic hardwood forest dominated by oak, ironwood, beech, and hemlock, under moist leaf litter.

Etymology

This species is named for its type locality, Frozen Head State Park in the Crab Orchard Mountains of eastern Tennessee. The specific name is a latinized noun in apposition derived from the Greek kryos-, meaning frozen, and maia- for a kind of crab.

Type locality

United States, Tennessee, Morgan County, campground at Frozen Head State Park; 36.1321°N, -84.4978°W.

Nannaria daptria sp. nov.

http://zoobank.org/78C0F60F-71A6-4C4B-BD7B-6BD036C30277

Figs 98, 99 Vernacular name: “The Picnicking Twisted-Claw Millipede”

Material examined

Holotype: United States – Tennessee • ♂; Tennessee, Greene County, Paint Creek Corridor, Overlook Picnic Area, forest across road; 35.9777°N, -82.8478°W; elev. 500 m; 16 June 2018; hand collected; D. Hennen leg.; VTEC MPE04156. For detailed collection data see Suppl. material 7.

Diagnosis

Adult males of Nannaria daptria sp. nov. are distinct from other Nannaria and the nearby N. scutellaria, based on the following combination of characters: Gonopods. Gonopodal acropodite gently curving medially throughout, not straight with 90° medial bend as in N. scutellaria. Tip blunt, with small, triangular lateral and medial flanges (Fig. 98A, red arrows), not simple as in N. minor. Acropodite swollen before apex (Fig. 98A, red triangle). Acropodite lacking tooth-like medial flange of N. minor, or acicular medial flange of N. scutellaria. Prefemur with long, thin, medially curving prefemoral process, not laterally curving as in N. minor and N. scutellaria. Prefemoral spine prominent, claw-like, curving cephalically, not small as in N. minor. Telopodite basal zone > 1/3 length of acropodite, not < 1/3 as in N. scutellaria. Color. Tergites with orange paranotal spots (Fig. 99). Light brown to black background. Dorsum of collum smooth with orange and white margin.

Figure 98.

Nannaria daptria sp. nov. holotype ♂ (VTEC, MPE04156) left gonopod A anterior view; red arrows indicate medial and lateral flanges; red triangle indicates medial swelling B medial view C posterior view. Scale bar: 0.5 mm.

Figure 99.

Nannaria daptria sp. nov. holotype ♂ (VTEC, MPE04156) coloration. Scale bar: 4.0 mm.

Measurements

♂ holotype (VTEC, MPE04156): BL = 30.6, CW = 3.7, IW = 2.0, ISW = 0.8, B11W = 4.8, B11H = 3.4.

Variation

No known variation.

Distribution

Known only from the type locality (Tennessee: Greene County, Suppl. material 7; Fig. 126). Distribution area: N/A; status: MRE.

Ecology

Nannaria daptria sp. nov. was collected from mesic, broadleaved forests composed of oak, maple, rhododendron, and hemlock, found under deciduous leaf litter in sandy soil.

Etymology

This species was named for the relaxing lunch that directly preceded its discovery. The specific name is a noun in apposition derived from the Greek daptria, meaning ‘eater.’

Type locality

United States, Tennessee, Greene County, Paint Creek Corridor, Overlook Picnic Area, forest across road; 35.9777°N, -82.8478°W.

Nannaria hippopotamus sp. nov.

http://zoobank.org/AFE8AC4E-F463-4AA3-A9F6-4D0B4C16DCC9

Figs 100, 101 Vernacular name: “The Horse Creek Twisted-Claw Millipede”

Material examined

Holotype: United States – Tennessee • ♂; Greene County, Cherokee National Forest, Horse Creek Rec Area, picnic area on east side of Horse Creek; 36.1059°N, -82.6545°W; elev. 581 m; 16 June 2018; hand collected; D. Hennen leg.; VTEC MPE04150.

Paratypes: United States – Tennessee • 3 ♀; same collection data as holotype; VTEC MPE04151–53 • 3 ♀♀; same collection data as holotype; VMNH MPE04154, 55, 4233 • 1 ♂; same collection data as holotype; 16 Oct. 1978; R. Shelley, W. Jones leg.; NCSM NAN0469.

Other material

United States – Tennessee • 1 ♂; Greene County, 8.5 air km south of Horse Creek, Cherokee National Forest, Old Forge Recreation Area; 36.0878°N, -82.6811°W; elev. 623 m; 15 June 2018; hand collected; D. Hennen leg.; VTEC MPE04141 • 1 ♀; same collection data as preceding; VTEC MPE04142. For detailed collection data see Suppl. material 7.

Diagnosis

Adult males of Nannaria hippopotamus sp. nov. are distinct from other Nannaria, and the nearby N. scutellaria, based on the following combination of characters: Gonopods. Gonopodal acropodite straight, curving medially at apex. Acropodite tip with prominent lobed lateral flange (Fig. 100A, red arrow), curving dorsomedially, not simple as in N. terricola, or with tooth-like medial flange as in N. scutellaria. Acropodite swollen medially before apex (Fig. 100A, red triangle). Telopodite basal zone ca. 1/2 length of acropodite, not ca. 1/4 as in N. scutellaria. Telopodite basal zone with very slight lateral bulge, not prominent as in N. terricola. Prefemur with straight, acicular prefemoral process, not curving ventrally as in N. scutellaria. Prefemoral spine small, sharp, not fused with prefemoral process as in N. terricola. Color. Tergites with orange paranotal spots (Fig. 101). Black background. Dorsum of collum smooth with orange margin.

Figure 100.

Nannaria hippopotamus sp. nov. paratype ♂ (NCSM, NAN0469) left gonopod A anterior view; red arrow indicates lateral flange; red triangle indicates acropodite medial swelling B medial view C posterior view. Scale bar: 0.5 mm.

Figure 101.

Nannaria hippopotamus sp. nov. non-type ♂ (VTEC, MPE04141) coloration. Scale bar: 4.0 mm.

Measurements

♂ holotype (VTEC, MPE04150): BL = 30.4, CW = 4.2, IW = 2.3, ISW = 1.0, B11W = 4.9, B11H = 3.3; ♀ paratype (VTEC, MPE04152): BL = 31.2, CW – 4.1, IW = 2.3, ISW = 1.0, B11W = 5.3, B11H = 4.0.

Variation

No known variation.

Distribution

Known only from two campsites in eastern Tennessee (Tennessee: Greene County; Suppl. material 5; Fig. 126). Distribution area: N/A; status: MRE.

Ecology

Individuals of Nannaria hippopotamus sp. nov. have been found under moist leaf litter in mesic broadleaved forests dominated by sweetgum, hickory, hemlock, oak, rhododendron, and maple.

Etymology

This species is named for its type locality, Horse Creek, a national forest recreation area hardwood cove. The specific name is a noun in apposition derived from the Greek hippos-, meaning horse, and potamós, meaning river.

Type locality

United States, Tennessee, Greene County, Cherokee National Forest, Horse Creek Rec Area, picnic area on east side, 36.1059°N, -82.6545°W.

Nannaria honeytreetrailensis sp. nov.

http://zoobank.org/90CC3A0F-C6AF-41BA-A4A7-CB7D94721204

Figs 102, 103 Vernacular name: “The Honey Tree Trail Twisted-Claw Millipede”

Material examined

Holotype: United States – Virginia • ♂; Lee County, Cumberland Gap National Historic Park, Honey Tree Trail, 0.2 km E junction with Gibson Gap Trail; 36.6078°N, -83.6322°W; 29 Sep. 2006; P. Marek leg.; VTEC MMC0334.

Diagnosis

Adult males of Nannaria honeytreetrailensis sp. nov. are distinct from other Nannaria and the nearby N. scutellaria, based on the following combination of characters: Gonopods: Gonopodal acropodite gently curving medially throughout, not straight with 90° medial bend as in N. scutellaria. Tip blunt, with small, triangular lateral flange (Fig. 102A, red arrow), lacking triangular medial flange of N. daptria sp. nov. Acropodite swollen basal to apex (Fig. 102A, red triangle), lacking acicular medial flange of N. scutellaria. Prefemur with short, acuminate, ventrally curving prefemoral process, not long, medially curving as in N. daptria sp. nov., or laterally curving as in N. scutellaria. Prefemoral process arising dorsolaterally from prominent, blunt prefemoral spine, not claw-like, as in N. daptria sp. nov. Telopodite basal zone height > 1/3 length of acropodite, not < 1/3 as in N. scutellaria. Color: Tergites with large, hot orange/pink paranotal spots (Fig. 103). Black background. Dorsum of collum smooth with hot orange/pink margin.

Figure 102.

Nannaria honeytreetrailensis sp. nov. holotype ♂ (VTEC, MMC0334) left gonopod A anterior view; red arrow indicates medial flange; red triangle indicates medial swelling B medial view C posterior view. Scale bar: 0.5 mm.

Figure 103.

Nannaria honeytreetrailensis sp. nov. holotype ♂ (VTEC, MMC0334) coloration. Scale bar: 4.0 mm.

Measurements

♂ holotype (VTEC, MMC0334): BL = 35.5, CW = 4.5, IW = 2.6, ISW = 0.9, B11W = 5.5, B11H = 3.8.

Variation

No known variation.

Distribution

Nannaria honeytreetrailensis sp. nov. is known only from the type locality (Virginia: Lee County; Suppl. material 7, Fig. 126). Distribution area: N/A; status: MRE.

Ecology

The N. honeytreetrailensis sp. nov. individual was collected from a mesic hardwood forest, on the side of a hiking path.

Etymology

This species is named for its type locality. The specific name is a feminine adjective derived from the type locality.

Type locality

United States, Virginia, Lee County, Cumberland Gap National Historic Park, Honey Tree Trail, 0.2 km E junction with Gibson Gap Trail, 36.6078°N, -83.6322°W.

Nannaria kassoni sp. nov.

http://zoobank.org/72902CE6-1FAF-46D7-8937-338D97E684BE

Figs 104, 105 Vernacular name: “Kasson’s Twisted-Claw Millipede”

Material examined

Holotype: United States – Tennessee • ♂; Campbell County, Caryville, along Cove Creek Trail off Bruce Gap Rd., near Route I-75 and the peak of Log Mountain; 36.3072°N, -84.2260°W; elev. 320 m; 3 June 2015; hand collected; M. Kasson leg.; VTEC MPE00544.

Other material

United States – Tennessee • 1 ♂; Campbell County, 10 miles NE of LaFollette; 36.4716°N, -83.9848°W; 11 May 1951; L. Hubricht leg.; VMNH NAN0192. For detailed collection data see Suppl. material 7.

Diagnosis

Adult males of Nannaria kassoni sp. nov. are distinct from other Nannaria and the nearby Nannaria sp. nov. ‘Cratagae’ (wilsoni species group) based on the following combination of characters: Gonopods. Gonopodal acropodite curving medially before apex, not straight as in Nannaria sp. nov. ‘Cratagae.’ Apex with distinct constriction (Fig. 104A, red arrow), tip sharp, with acuminate, triangular lateral flange, not blunt with small lobed lateral and medial flanges as in N. cryomaia sp. nov. Acropodite with expanded, laminate, medial flange (Fig. 104A, red triangle). Prefemur with medially curving prefemoral process, paralleling curve of acropodite, not straight, acicular as in Nannaria sp. nov. ‘Cratagae.’ Prefemoral process arising dorsolaterally from pronounced curving prefemoral spine, not arising from prefemur with short, triangular prefemoral spine as in N. cryomaia sp. nov. Telopodite basal zone height ca. 1/3 length of acropodite, not > 1/3 as in N. cryomaia sp. nov., and not ca. 1/4 length as in Nannaria sp. nov. ‘Cratagae.’ Color. Tergites with orange stripes (Fig. 105). Dark brown background. Dorsum of collum smooth with orange margin.

Figure 104.

Nannaria kassoni sp. nov. holotype ♂ (VTEC, MPE00544) left gonopod A anterior view; red arrow indicates apical constriction; red triangle indicates medial flange B medial view C posterior view. Scale bar: 0.5 mm.

Figure 105.

Nannaria kassoni sp. nov. holotype ♂ (VTEC, MPE00544) coloration. Scale bar: 4.0 mm.

Measurements

♂ holotype (VTEC, MPE00544): BL = 27.0, CW = 3.8, IW = 2.2, ISW = 1.0, B11W = 4.7, B11H = 2.6.

Variation

No known variation.

Distribution

Known from two locations near La Follette, Tennessee (Tennessee: Campbell County; Suppl. material 7, Fig. 126). Distribution area: N/A; status: MRE.

Ecology

Individuals of N. kassoni sp. nov. were collected from mesic broadleaved forests, ranging from ca. 300 m elevation adjacent to a floodplain to ca. 750 m elevation near the peak of Log Mountain.

Etymology

This species is named after its collector, Dr. Matt Kasson. The specific name is a genitive noun derived as a patronym.

Type locality

United States, Tennessee, Campbell County, Caryville, along Cove Creek Trail off Bruce Gap Rd., near Route I-75 and the peak of Log Mountain, 36.3072°N, -84.2260°W.

Nannaria minor Chamberlin, 1918

Figs 106, 107 Vernacular name: “The First Twisted-Claw Millipede”

Nannaria minor Chamberlin, 1918: 124. Attems 1938: 199. Brimley 1938: 499. Chamberlin and Hoffman 1958: 41. Hoffman 1964: 11. Wray 1967: 152. Hoffman 1999: 367. Shelley 2000: 196. Marek et al. 2014: 37. Means et al. 2021: 16, S71.

Material examined

United States – North Carolina • 1 ♂; Ashe County, West Jefferson, Mount Jefferson State Natural Area, lower loop of Rhododendron Trail; 36.4032°N, -81.4646°W; elev. 1388 m); 10 Oct. 2018; hand collected; D. Hennen leg.; VTEC MPE04388 • 1 ♀; same collection data as preceding; VTEC MPE04389 • 3 ♂♂; Mitchell County, Little Switzerland, Grassy Creek Falls; 35.8561°N, -82.0876°W; elev. 995 m; hand collected; D. Hennen leg.; VTEC MPE04391–93 • 2 ♀♀; same collection data as preceding; VTEC, MPE04394, 4395 • 6 ♂♂; Mitchell County, Crabtree Meadows Rec. Area, B.R.P.; 35.8996°N, -82.2147°W; elev. 1036–1097 m; 30 Oct. 1971; R. Hoffman, Knight leg.; VMNH NAN0200 • 1 ♂; Mitchell County, 1000’ S of Sam’s Gap, on U.S. Hy. 23; 35.9385°N, -82.5632°W; 2 Aug. 1962; R. Hoffman leg.; VMNH NAN0202 • 1 ♂; Mitchell County, Bakersville, around garage at home; 36.0156°N, -82.1587°W; 13 Feb. 1976; D. Terrel leg.; NCSM NAN0554 • 1 ♂; Mitchell County, summit of Roan Mtn., in spruce-fir forest; 36.1039°N, -82.1219°W; elev. 1829 m; 24 July 1975; R. Shelley, J. Clamp leg.; NCSM NAN0502 • 4 ♂♂♀♀; Watauga County, 2.8 WNW Blowing Rock, Julian Price Memorial Park, off Blue Ridge Pkwy.; 36.1422°N, -81.7456°W; 9 Sep. 1973; R. Shelley leg.; NCSM NAN0520; SCAU – Tennessee • 1 ♀; Carter County, northeast of Hampton, Watauga Point Rec Area, trail from parking lot; 36.3199°N, -82.0834°W; elev. 632 m; 12 June 2018; hand collected; D. Hennen leg.; VTEC MPE04060 • 1 ♂; Carter County, Hampton, Cherokee National Forest, Laurel Creek Falls Trail; 36.2652°N, -82.1235°W; elev. 768 m; 12 June 2018; hand collected; D. Hennen leg.; VTEC MPE04074 • 3 ♀♀; Carter County, Roan Mountain, Roan Mountain State Park, Blue 2 Trail; 36.1674°N, -82.0984°W; elev. 872 m; 12 June 2018; D. Hennen leg.; VTEC MPE04080–82 • 3 ♂♂; Carter County, The Laurels Rec Area, Cherokee National Forest; 36.2476°N, -82.2696°W; elev. 584 m; 13 June 2018; D. Hennen leg.; VTEC MPE04089, 4090, 4226 • 2 ♀♀; same collection data as preceding; VTEC, MPE04091, 4092 • 1 ♂; Carter County, south of Alizabethton, at pull off on Route 362; 36.2651°N, -82.2300°W; elev. 620 m; 25 May 2016; hand collected; J. Means, D. Hennen leg.; VTEC MPE01249 • 8 ♀♀; same collection data as preceding; VTEC MPE01252, 1253, 1296, 1302, 1303, 1313, 1314, 2061 • 1 ♂; Carter County, Doe R. Bluff, 1 mile NW of Hampton; 36.2944°N, -82.1855°W; 3 May 1951; L. Hubricht leg.; VMNH NAN0293 • 1 ♂; Loudon County, Lenoir City, basement of house; 35.7972°N, -84.2561°W; 10 Dec. 1971; W. Tolbert leg.; VMNH NAN0144 • 1 ♀; Unicoi County, Unicoi, Limestone Cove Rec Area; 36.1742°N, -82.2982°W; elev. 697 m; hand collected; D. Hennen leg.; VTEC MPE04103 • 3 ♀♀; Unicoi County, Erwin, Cherokee National Forest, Rock Creek Falls Rec Area, Rock Creek Falls Trail; 36.1389°N, -82.3468°W; elev. 704 m; 13 June 2018; hand collected; D. Hennen leg.; VTEC MPE04107, 4108, 4231 • 2 ♂♂; Unicoi County, 10.7 air km from Erwin, Cherokee National Forest, beginning of trail to Sill Creek Falls, off Clark Creek Road; 36.1281°N, -82.5340°W; elev. 559; 14 June 2018; hand collected; D. Hennen leg.; VTEC MPE04127, 4128 • 3 ♀♀; Unicoi County, Flag Pond, Rocky Fork State Park, Rocky Fork Trail; 36.0482°N, -82.5615°W; elev. 740 m; hand collected; D. Hennen leg.; VTEC MPE4133–35 • 1 ♂; Unicoi County, 3 SW Erwin, Unaka Springs Rd., 1 mi jct. Chestoa Rd.; 36.0931°N, -82.4396°W; 1 Apr. 2002; A. Gagan leg.; NCSM NAN0476 • 2 ♀♀; Unicoi County, Erwin, Cherokee National Forest, Rock Creek Falls Rec Area, Rock Creek Falls Trail; 36.1389°N, -82.3468°W; 13 June 2018; hand collected; D. Hennen leg.; VTEC MPE04108, 4231 • 1 ♀; Washington County, Johnson City, Buffalo Mountain Park, White Rock Lower Loop; 36.2770°N, -82.3461°W; elev. 717 m; 13 June 2018; D. Hennen leg.; VTEC MPE04100. For detailed collection data see Suppl. material 7.

Diagnosis

Adult males of Nannaria minor are distinct from other Nannaria, and the nearby N. aenigma, based on the following combination of characters: Gonopods. Gonopodal acropodite gently curving medially throughout with extremely short, blunt distal zone, not with serpentine distal zone as in N. aenigma. Acropodite with small, tooth-like medial flange near apex (Fig. 106A, red arrow), not with lobed medial flange as in N. mcelroyorum sp. nov. Prefemur with long, thin, acicular prefemoral process slightly curving laterally, not sinuous, cephalically directed as in N. mcelroyorum sp. nov. Prefemoral spine small, curving cephalically (Fig. 106B, red triangle), without secondary hump as in N. mcelroyorum sp. nov. Telopodite basal zone height < 1/2 length of acropodite, not ca. 1/2 length as in N. mcelroyorum sp. nov., or ca. 1/6 length as in N. aenigma. Color. Tergites with hot pink/orange paranotal spots (Fig. 107). Light brown to black background. Dorsum of collum smooth with orange margin.

Figure 106.

Nannaria minor ♂ (VTEC, MPE01249) left gonopod A anterior view; red arrow indicates tooth-like medial flange B medial view C posterior view; red triangle indicates cephalically curved prefemoral spine. Scale bar: 0.5 mm.

Figure 107.

Nannaria minor ♂ (VTEC, MPE01249) coloration. Scale bar: 4.0 mm.

Measurements

♂ (VTEC, MPE01249): BL = 31.8, CW = 3.9, IW = 2.1, ISW = 0.9, B11W = 4.6, B11H = 3.1; ♀ (VTEC, MPE01253): BL = 31.2, CW = 3.7, IW = 2.1, ISW = 1.0, B11W = 5.1, B11H = 3.5.

Variation

Notable variation found in N. minor is the length and general directionality of the prefemoral process; a specimen collected from the summit of Roan Mountain in North Carolina (NAN0502) has a short and somewhat curving prefemoral process, while specimens from Sam’s Gap in Tennessee (NAN0202) have long prefemoral processes that do not cross the acropodite in the anterior view, and the specimen illustrated by Hoffman (1964: 32) has a medially directed prefemoral tip.

Distribution

Known from the Southern Appalachians, primarily along the northern half of the North Carolina and Tennessee border (North Carolina: Ashe, Madison, Mitchell, Watauga, and Yancey counties; Tennessee: Carter, Loudon, and Unicoi counties; Virginia: Carroll County, Suppl. material 7; Fig. 126). Distribution area: 5,671 km²; status: SRE.

Ecology

Individuals of Nannaria minor have been collected from mesic deciduous forests, often dominated by maple, oak, hemlock, and rhododendron, typically under 1–2 cm soil and/or leaf litter.

Etymology

Chamberlin (1918) gave no etymology for his name, N. minor, though it is reasonable to assume that he named both the genus and the species for its small size in relation to other xystodesmids. Why Chamberlin would feel the need to reiterate the small nature of the species is unknown. One possibility is that Chamberlin may have chosen the specific name after the Latin word minor for ‘smaller’ to differentiate it from a second species he described in the same paper N. media Chamberlin, 1918 (now Boraria stricta) which was slightly larger than N. minor, but not as large as the third species he described, N. infesta (now Howellaria infesta). Thus, minor was the smallest of the three species, media (from the Latin medi- meaning middle) was the middle-sized species, and infesta was the largest. If this hypothesis is true it is a mystery to the authors why Chamberlin broke the pattern with infesta, though the fungal infestation of the infesta type may have offered up too good of a species epithet to pass up.

Type locality

United States, Tennessee, Burbank.

Notes

In the original publication, Chamberlin (1918, 125) mentions that two specimens were collected by R. Thaxter: a male, which Chamberlin designates as the type, and a female.

Nannaria rhysodesmoides (Hennen & Shelley, 2015)

Fig. 108 Vernacular name: “The Rhysodesmus-Like Twisted-Claw Millipede”

Mimuloria rhysodesmoides Hennen & Shelley, 2015: 1–16.

Nannaria rhysodesmoides: Means et al. 2021: S72.

Material examined

Holotype: United States – Virginia • ♂; Putnam County, Cookeville; [36.1628°N, -85.5016°W]; 12 Apr. 1958; H.E. Evans leg.; FSCA.

Paratypes: United States – Virginia • 6 ♂♂; same collection data as holotype; NCSM 27913. For detailed collection data see Suppl. material 7.

Diagnosis

Adult males of Nannaria rhysodesmoides are distinct from other Nannaria and the nearby N. blackmountainensis sp. nov. based on the following combination of characters: Gonopods. Gonopodal acropodite simple, gently curving dorsomedially, not with pronounced medial swelling as in N. blackmountainensis sp. nov. Acropodite tip with small, triangular lateral flange (Fig. 108A, red arrow), not large, hooked lateral flange as in N. blackmountainensis sp. nov. Apex slightly sinuate (Fig. 108B, red triangle). Acropodite tip terminating in small, sharp, dorsally directed point, not blunt, squared off as in N. blackmountainensis sp. nov. Height of telopodite basal zone < 1/2 length of prefemoral process, not ca. 1/2 length of prefemoral process as in N. blackmountainensis sp. nov. Prefemoral process straight, wide, approaching laminate, not thin and acuminate as in N. blackmountainensis sp. nov. Prefemoral spine completely reduced not forming small ridge as in N. blackmountainensis sp. nov. Color. The preserved holotype suggests that N. rhysodesmoides may have metatergal stripes connecting the paranotal spots, though living specimens have not be observed by the authors.

Figure 108.

Nannaria rhysodesmoides holotype ♂ (FSCA) left gonopod A anterior view, red arrow indicates small, pointed lateral flange B medial view; red triangle indicates slight sinuation of the acropodite apex C posterior view. Scale bar: 0.5 mm.

Measurements

♂ paratype (NCSM27913; ♂ holotype too fragmented for measurement): BL = 22.3, CW = 2.9, IW = 1.8, ISW = 0.8, B11W = 4.4, B11H = 2.4.

Variation

No known variation.

Distribution

Known only from the type locality (Tennessee: Putnam County, Suppl. material 7; Fig. 129). Distribution area: N/A; status: MRE.

Ecology

Hennen and Shelley (2015) do not mention ecological notes in their description of the species, most likely due to a lack of ecological information from the original collector.

Etymology

Hennen and Shelley (2015) chose the specific name based on the presence of a lateral flange on the gonopod (Fig. 108A, red arrow), a character also seen in the genus Rhysodesmus Cook, 1895.

Type locality

United States, Virginia, Putnam County, Cookeville.

Notes

In the original publication, Hennen and Shelley (2015: 15) designate a male holotype and five male paratypes (FSCA) collected by H. E. Evans on 12 April 1958, and note that one male paratype from this collection was retained at the NCSM.

Species with uncertain clade membership

Nannaria conservata Chamberlin, 1940

Fig. 109 Vernacular name: “The Duke Forest Twisted-Claw Millipede”

Nannaria conservata Chamberlin 1940: 56. Chamberlin and Hoffman 1958: 40. Wray 1967: 152. Shelley 1974: 111. Shelley 1975: 180, figs 1–6. Shelley 1978: 66, figs 67–69. Hoffman 1999: 366. Shelley 2000: 196. Marek et al. 2014: 36. Means et al. 2021: S69.

Material examined

United States – North Carolina • 1 ♂; Franklin County, near Moccasin Creek, 1.5 miles SW of Pilot; 35.8625°N, -78.2683°W; 2 May 1959; L. Hubricht leg.; VMNH, NAN0336 • 7 ♂♂♀♀; Person County, 2 mi SSE Surl, off end of SR 1718 in window wells, Sheets residence; 36.3157°N, -78.8887°W; 3 Apr. 1999; A. & S. Braswell, R. Sheets leg.; NCSM NAN0547 • 1 ♀; Wake County, pond near William B Umstead State Park; 35.8366°N, -78.7623°W; elev. 118 m; 17 Apr. 2016; hand collected; J. Means leg.; VTEC MPE02117 • 8 ♂♂; Wake County, Raleigh, 209 Lynwood Lane; 35.8404°N, -78.6376°W; 15 Mar. 1975; M. Cooper leg.; VMNH NAN0337, 338 • 12 ♂♂♀♀; same collection data as preceding; 29 Jan. 1975; M. Cooper leg.; NCSM, NAN0549, 550, 552 • 2 ♂; Wake County, Raleigh, 1611 Oberlin Rd.; 35.8036°N, -78.6582°W; 23 Jan. 1990; D. Stephan leg.; NCSM NAN0546 • 1 ♂; Wake County, Raleigh, jct. Athens Dr. & Avent Ferry Rd.; 35.7674°N, -78.7066°W; 10 Mar. 1990; R. Shelley leg.; NCSM NAN0550. For detailed collection data see Suppl. material 7.

Diagnosis

Adult males of Nannaria conservata are distinct from other Nannaria and the nearby N. hardeni sp. nov. based on the following combination of characters: Gonopods. Gonopodal acropodite gently curving dorsomedially before apex, not medially as in N. hardeni sp. nov.; tip simple, blunt. Telepodite basal zone height < 1/2 length of acropodite, not > 1/2 as in N. hardeni sp. nov. Prefemoral process acicular, reduced, not subequal to height of telopodite basal zone and crossing acropodite as in N. hardeni sp. nov. Prefemoral spine acicular and paralleling prefemoral process (Fig. 109B, red arrow), not blunt and widely separated from prefemoral process as in N. hardeni sp. nov. Color. In his redescription of N. conservata, Shelley (1975) describes its color as “olive-brown with a pink epiproct and paranota…” The single female individual collected for this revision conformed to Shelley’s description, but was placed in 100% EtOH in the field not photographed alive.

Figure 109.

Nannaria conservata ♂ (VMNH, NAN0338) left gonopod A anterior view B medial view; red arrow indicates reduced prefemoral spine paralleling prefemoral process C posterior view. Scale bar: 0.5 mm.

Measurements

♂ (VMNH, NAN0388): BL = 21.2, CW = 2.8, IW = 1.6, ISW = 0.8, B11W = 3.2, B11H = 2.3.

Variation

There is no notable variation between individuals of N. conservata, with the exception of an abnormal male with three gonopods which Shelley (1975) illustrated and discussed.

Distribution

Known only from a small area around Raleigh and Durham, North Carolina (North Carolina: Person, Orange, Wake and Franklin counties; Suppl. material 7; Fig. 128). Distribution area: 1,820 km²; status: SRE.

Ecology

As no fresh identifiable material was collected for this revision, the authors cannot comment on habitat preference. However, Shelley (1975) noted that specimens were never found in moist areas, and instead were discovered under pine litter and in sandy, dry soils. As Shelley (1975) noted, this is unusual for a majority of xystodesmid species.

Etymology

Chamberlin (1940) gives no etymology for N. conservata, but the specific name is seemingly a femininization of the Latin conservo, meaning conserve. This may be in reference to the conservation efforts which take place within its type locality, Duke Forest.

Note on clade membership

The authors were unsuccessful at collecting specimens of Nannaria conservata for the present study, so DNA sequences, molecular phylogenetic relationship, and taxonomic affinities between N. conservata and the rest of the minor species group remain unknown. A female specimen from Raleigh, Wake County, North Carolina was collected and sequenced, and was placed within the paupertas clade. The only species of Nannaria known from Wake County is N. conservata. However, as N. conservata shares none of the gonopodal characters that unite the paupertas clade, and in light of the possibility that the female from Wake County represents an undescribed species, the authors refrain from placing N. conservata within the paupertas clade and leave the taxonomic affinity of N. conservata to be addressed in a future study.

Type locality

United States, North Carolina, Durham County, Durham.

Notes

In the original publication, Chamberlin (1940: 57) did not designate a type, but he describes only a male specimen which is therefore the holotype by monotypy. Chamberlin did not mention the collector of this specimen, but Shelley (1975: 180) reported that the type specimen was collected by N. B. Causey on November 12, 1939.

Nannaria oblonga (Koch, 1847)

Fig. 110 Vernacular Name: “The Pennsylvania Twisted-Claw Millipede”

Fontaria oblonga C.L. Koch, 1847: 142. C.L. Koch 1863: 73–74, fig. 64. Attems 1898: 263. Attems 1938: 167. Chamberlin and Hoffman 1958: 55.

Nannaria oblonga: Hoffman 1999: 367. Marek et al. 2014: 37. Means et al. 2021: S71.

Material examined

Syntype: United States – Pennsylvania • ♂; from an unknown locality in “Pennsylvania”; MFN.

Diagnosis

Adult males of Nannaria oblonga are distinct from other Nannaria, the possibly sympatric N. fowleri, and the nearby N. shenandoa, based on the following combination of characters: Gonopods. Gonopodal acropodite gently curving medially before apex, not strongly curved as in N. shenandoa. Distal zone short, simple, curving smoothly posteriorly, with a crochet-like appearance (Fig. 110A, red triangle). Distal zone not directed medially as in N. fowleri, or elongated, with flanges as in N. shenandoa. Telopodite basal zone ca. 1/3 length of acropodite, not ca. 1/4 as in N. shenandoa. Prefemur with prefemoral process thin, ca. 1/2 length of acropodite, curving medially to parallel acropodite near tip, not elongated, 1/3 length of acropodite and paralleling acropodite majority of length as in N. fowleri, or curving laterally as in N. shenandoa. Prefemoral spine lacking, not pronounced as tooth-like structure as in N. fowleri. Color. Koch (1847) describes N. oblonga as being dark brown with white paranota.

Figure 110.

Nannaria oblonga syntype ♂ (MFN) left gonopod A anterior view; red triangle indicates crochet-like acropodite tip B medial view C posterior view. Scale bar: 0.5 mm.

Measurements

The type is too fragmented for accurate measurement.

Variation

No known variation.

Distribution

Known only from the type locality (Pennsylvania, Fig. 127). Distribution area: N/A; status: MRE.

Ecology

Koch (1847) does not mention any ecological notes in his description of the species.

Etymology

Koch (1847) does not give an etymology for the name oblonga in his 1863 publication, he goes into more detail about the syntype, and describes the species as “länglich,” meaning elongated or oblong in German. It is therefore reasonable to assume that the name is in reference to the general oblong shape of Nannaria.

Note on clade membership

No specimens of Nannaria oblonga were collected for this study, and while N. oblonga shares some gonopodal characters with N. tenuis sp. nov. (such as the caudally-directed acropodite tip and a lack of a prefemoral spine) N. oblonga is geographically disjunct from the ignis clade, to which N. tenuis sp. nov. belongs. Due to a lack of molecular data and a paucity of shared morphological characters, clade membership is uncertain.

Type locality

United States, Pennsylvania; MFN.

Notes

In the original publication, Koch (1847: 142) did not designate a type or mention the number of specimens he examined and later on, Koch (1863) described both a male and female specimen in greater detail. The specimen in the MFN collection was labeled “Holotypus.” Without knowing the number of specimens Koch examined for his original description of the species, we cannot be sure if it is a holotype by monotypy, and hence refrain from designating a lectotype.

Nannaria rutherfordensis Shelley, 1975

Fig. 111 Vernacular name: “The Rutherfordton Twisted-Claw Millipede”

Nannaria rutherfordensis Shelley, 1975: 184, figs 7–9. Hoffman 1999: 367. Shelley 2000, 197. Marek et al. 2014: 38. Means et al. 2021: S72.

Material examined

Holotype: United States – North Carolina • ♂; Rutherford County, Rutherfordton, Bus. 74, 0.2 W Byp. 74; [35.3108°N, -81.9303°W]; 15 Oct. 1973; R. M. Shelley leg.; NMNH USNMENT 01491821. For detailed collection data see Suppl. material 7.

Diagnosis

Adult males of Nannaria rutherfordensis are distinct from other Nannaria and the nearby N. minor based on the following combination of characters: Gonopods. Gonopodal acropodite simple, gently curving medially with extremely short, blunt distal zone, not with small, tooth-like medial flange near apex as in N. minor. Prefemur with long, thin, medially curving prefemoral process, not acicular slightly curving laterally as in N. minor. Prefemoral process arising from sharp, projected prefemoral spine (Fig. 111A, red arrow), not from small prefemoral spine which curves cephalically as in N. minor. Telopodite basal zone height < 1/2 length of acropodite. Color. Shelley (1975) described N. rutherfordensis as having white paranota with a concolorous stripe along the anterior edge of the collum.

Figure 111.

Nannaria rutherfordensis holotype ♂ (NMNH, USNMENT 01491821) left gonopod A anterior view, red arrow indicates pronounced, sharp prefemoral spine B medial view C posterior view. Scale bar: 0.5 mm.

Measurements

♂ holotype (NMNH, USNMENT 01491821): BL = 28.0, CW = 3.2, IW = 2.2, ISW = 0.9, B11W = 4.3, B11H = 2.8.

Variation

No known variation.

Distribution

Known only from the type locality (North Carolina: Rutherford County, Suppl. material 7, Fig. 126). Distribution area: N/A; status: MRE.

Ecology

Shelley (1975, pp. 186) notes that the holotype was found under leaves by the side of a log in an “urban patch of hardwoods, a few feet from the pavement...” The authors JCM and DAH were unable to recollect N. rutherfordensis on multiple trips to the type locality and surrounding area. The type locality had unfortunately recently been cleared for development and was found dry and lacking vegetation on each visit. The authors infer this species may be locally extirpated.

Etymology

Shelley (1975) provides no etymology for his naming in his description of the species; however, it is reasonable to assume that N. rutherfordensis is named after its type locality.

Note on clade membership

No specimens of Nannaria rutherfordensis were collected for this study. While it shares the gonopodal character of a prefemoral process that parallels the curved acropodite with N. kassoni sp. nov. (minor clade), this character is also found in N. fowleri (fowleri clade) and N. tasskelsoae sp. nov. (tasskelsoae clade), which are 352 km and 266 km north, respectively. In combination with the relative geographic isolation of the species, being the most southern species in North Carolina, clade membership is uncertain.

Type locality

United States, North Carolina, Rutherford County, Rutherfordton, Bus. 74, 0.2 W Byp. 74; [35.3108°N, -81.9303°W].

Notes

In the original publication, Shelley (1975: 184) designates a male holotype (NCSM 2053) which he collected on October 15, 1973. Interestingly, despite the type being deposited by Shelley at the NCSM, at some point before the writing of Hoffman’s (1999) checklist the specimen was moved to the NMNH and relabeled with the code USNMENT 01491821 by an unknown party.

Nannaria sigmoidea (Hennen & Shelley, 2015)

Vernacular name: “The Sigmoid Gonopod Twisted-Claw Millipede” Fig. 112

Mimuloria dilatata sigmoidea Hennen & Shelley, 2015: 1–15, figs 20, 21.

Nannaria sigmoidea: Means et al. 2021: S72.

Material examined

Holotype: United States – Tennessee • ♂; Meigs County, c.a. 7.2 km S Decatur, along country road 4274, 0.8 km from junction of Tennessee highway 58; [35.4378°N, -84.9140°W]; 14 Oct. 1978; R. M. Shelley, W. B. Jones leg.; NCSM NCSM27946. For detailed collection data see Suppl. material 7.

Diagnosis

Adult males of N. sigmoidea are distinct from other Nannaria, including the nearby N. blackmountainensis sp. nov. and Nannaria sp. nov. ‘Cratagae’ (wilsoni species group), based on the following combination of characters: Gonopods. Gonopodal acropodite simple and gently curving anteromedially, not straight as in Nannaria sp. nov. ‘Cratagae’ or with medial swelling as in N. blackmountainensis sp. nov. Acropodite with prominent, rounded lateral flange near tip (Fig. 112A, red arrow), not hooked as in N. blackmountainensis sp. nov. or lacking as in Nannaria sp. nov. ‘Cratagae.’ Acropodite tip directed posteriorly, not dorsally as in N. blackmountainensis sp. nov. Telopodite basal zone subequal to length of prefemoral process, not ca. 1/2 length as in N. blackmountainensis sp. nov. or ca. 1/3 length as in Nannaria sp. nov. ‘Cratagae.’ Prefemur with prefemoral process bent at 90° angle at halfway point, directed medioventrally, not straight, acicular as in N. blackmountainensis sp. nov. and Nannaria sp. nov. ‘Cratagae.’ Prefemoral spine reduced to small, blunt lobe at base of prefemoral process (Fig. 112A, red triangle), not fused as a small ridge as in N. blackmountainensis sp. nov. or lacking as in Nannaria sp. nov. ‘Cratagae.’ Color. Color in life unknown.

Figure 112.

Nannaria sigmoidea holotype ♂ (NCSM NCSM27946) left gonopod A anterior view; red arrow indicates lateral flange; red arrow indicates reduced prefemoral spine B medial view C posterior view. Scale bar: 0.5 mm.

Measurements

♂ holotype (NCSM, NCSM27946): BL = 36.1, CW = 5.0, IW = 3.0, ISW = 1.1, B11W = 6.0, B11H = 3.8.

Variation

No known variation.

Distribution

Known only from the type locality (Tennessee: Meigs County; Suppl. material 7; Fig. 129). Distribution area: N/A; status: MRE.

Ecology

Hennen and Shelley (2015) note that only that the type specimens were collected from the east side of the Tennessee River.

Etymology

Hennen and Shelley (2015: 15) stated that “[t]he subspecific name denotes the overall sigmoidal curvature of the prefemoral process, which also passes through numerous vertical planes.”

Note on clade membership

No specimens of Nannaria sigmoidea were collected for this study. Nannaria sigmoidea shares gonopodal characters with members of the ambulatrix clade, such as a prominent lateral flange on the acropodite tip, a prefemoral process bent at 90°, and crossing of the acropodites in the medial view (found in N. ambulatrix sp. nov. and N. botrydium sp. nov.). However, due to the wide geographic separation between N. sigmoidea and the ambulatrix clade (> 270 km), and the presence of these characters in a member of the minor clade (N. kassoni sp. nov.), the phylogenetic placement of N. sigmoidea in a clade is uncertain.

Type locality

United States, Tennessee, Meigs County, c.a. 7.2 km S Decatur, along country road 4274, 0.8 km from junction of Tennessee highway 58.

Notes

In the original publication, Hennen and Shelley (2015: 15) designate a male holotype collected by R. M. Shelley and W. B. Jones on October 14, 1978.

Figure 113.

The 17 minor species group clades represented by males. Black star: Nannaria scholastica, sky blue: N. domestica clade, orange: N. fowleri clade, burnt orange star: N. hardeni, brown: N. serpens clade, light yellow: N. paupertas clade, green: N. mcelroyorum clade, cranberry pink: N. tasskelsoae clade, turquoise: N. castanea clade, lime green: N. laminata clade, purple: N. ambulatrix clade, yellow: N. ignis clade, camo green: N. tennesseensis clade, blue: N. ohionis clade, light pink: N. blackmountainensis clade, forest green: N. terricola clade, red: N. minor clade.

Figure 114.

Geographic distributions of the 18 minor group clades. Black dot: Nannaria scholastica, sky blue: N. domestica clade, orange: N. fowleri clade, white: N. paupertas clade, dark green: Nannaria ‘Murphy’ clade, green: N. mcelroyorum clade, burnt orange: N. hardeni, brown: N. serpens clade, turquoise: N. castanea clade, neon green: N. laminata clade, cranberry pink: N. tasskelsoae clade, camo green: N. tennesseensis clade, yellow: N. ignis clade, blue: N. ohionis clade, pink: N. blackmountainensis clade, light green: N. terricola clade, purple: N. ambulatrix clade, red: N. minor clade.

Figure 125.

Nannaria “Sagittata” incertae sedis ♂ (VMNH, NAN0316) left gonopod A anterior view, red triangle indicates prefemoral process arising medially from prefemoral spine B medial view; red arrow indicates ventrally directed prefemoral process C posterior view. Scale bar: 0.5 mm.

Figure 126.

Distribution map of Nannaria minor species group collection localities in the central Appalachians.

Figure 127.

Distribution map of Nannaria minor species group collection localities in the northern Appalachians.

Figure 128.

Distribution map of Nannaria minor species group collection localities in the Eastern Piedmont and coast.

Figure 129.

Distribution map of Nannaria minor species group collection localities west of the Appalachians.

Discussion

We revised the genus Nannaria through a combination of molecular and morphological data, and provided the first molecular framework for this alpha-taxonomically poorly known and widespread group. Our molecular phylogeny supported previous molecular phylogenetic evidence that the tribe Nannariini is monophyletic within the Xystodesmidae, with O. pulchella sister to the remaining members of the tribe in the genus Nannaria (Suppl. material 3, Means et al. 2021). We described 35 new species, and confirmed that Nannaria contains two clades: the wilsoni and the minor species groups. The wilsoni species group occurs primarily in the central and southern Appalachian Mountains, while the minor species group is more widespread and occurs throughout the eastern and central U.S. and Canada. Within the minor species group, we found a considerable degree of morphological variation, even between closely related species, and a paucity of characters uniting species within clades (Fig. 113). We found several instances of close morphological similarity between distantly related species that are likely a result of convergence (e.g., N. ignis sp. nov. and N. minor). Means and Marek (2017) showed that morphological convergence between distantly related species within the family Xystodesmidae repeatedly evolved and was common, and could confuse taxonomy. Our findings suggest that morphology should not be used as the sole criteria for the taxonomic placement of species within Nannaria, and instead molecular phylogenetics ideally utilizing multiple genes should be used in combination with morphological and geographical data.

Nannaria minor and wilsoni species groups

Within Nannaria there are two primary stem group clades, the minor and wilsoni species groups, both containing considerable undescribed species diversity. Within the minor species group, there are 18 distinct clades (Suppl. material 3), which vary in terms of the morphological similarity between their member species. Both the minor and wilsoni species groups were recovered as monophyletic (Suppl. material 3, pp = 1), suggesting an evolutionary divide between the two groups. This divide has been unrecognized in the past literature, perhaps due to the lack of an obvious geographical separation and/or morphological synapomorphies which could be used to consistently differentiate species of either group. Upon examination of morphological characters in context of the molecular phylogeny, however, we have found characters that are useful for distinguishing wilsoni versus minor species groups. First, the minor species group exhibit a prefemoral spine (Fig. 5, ps), spare N. scholastica sp. nov., and a small subset of species (N. castra sp. nov., N. missouriensis, N. oblonga, and N. rhysodesmoides). None of the wilsoni species group taxa exhibit this character. Second, the majority of minor species group taxa have large basal zones, in some species reaching over half of the length of the telopodite, while wilsoni species group taxa have reduced basal zones, rarely reaching a third of the length of the acropodite. Third, minor species group taxa have uniformly unmodified cyphopod receptacles, while many wilsoni species group taxa have highly modified cyphopod receptacles (discussed in an upcoming revision of the wilsoni species group). Hoffman (1949) suggested that the distance between the bases of the last pair of legs and the shape of the “preanal scale” (= hypoproct) could be used to separate N. simplex (minor species group) and N. ericacea (wilsoni species group); however, these differences are not found consistently throughout either group and are therefore not useful for differentiating the minor versus wilsoni groups. Future species-level identifications of the genus should use the morphology of the gonopods or DNA barcoding with COI (Suppl. material 4C).

The utility of morphology in the minor species group

Previous descriptions of species in the genus Nannaria have been based solely on morphological characters, almost exclusively consisting of the male genitalia; however, morphological character scoring (Suppl. material 5) revealed little variation in non-genitalic somatic characters. Through molecular phylogenetic analysis of nearly 100 minor species group specimens, we have revealed the existence of 18 clades within the group, and demonstrated that morphological characters, while useful for species level diagnosis, are inadequate for determining evolutionary relationships among species. For example, the distinctive prefemoral process which was separated from the prefemoral spine, and arising from the prefemur and overlapping the acropodite in the anterior view is present in the species N. monsdomia sp. nov., N. hardeni sp. nov. and N. tsuga sp. nov. (Fig. 130). However, phylogenetic evidence indicated that N. monsdomia sp. nov., N. hardeni sp. nov. and N. tsuga sp. nov. all belong to separate clades, spread throughout the tree (Fig. 7), which suggest that this prefemoral process configuration evolved multiple times in the minor species group. The same is true for the telopodite basal zone, prefemoral spine, prefemur, and acropodite flange. We presumed that species with tall, thin basal zones would be closely related, such as in the case of N. ambulatrix sp. nov. and N. tenuis sp. nov.; however, these species are distantly related. Likewise, we expected that the prefemoral process and spine had diagnostic potential, yet both structures seem to vary considerably between closely related species, such as the prominent prefemoral spine in N. paupertas sp. nov., and the completely absent/reduced spine of its sister species, N. castra sp. nov. The prefemoral process and spine may aid in stabilizing the acropodite during mating, and perhaps allow for the repositioning of a spermatophore during copulation, as in the Parajulidae (Mathews and Bultman 1993); however, the function of the prefemoral structures have never been directly evaluated in Nannaria and therefore remain unknown. The position and shape of acropodite flanges, such as the can opener-like medial flange of N. minor (Fig. 106A) and N. fritzae sp. nov. (Fig. 55A) also seem to be inadequate diagnostic characters. Even multiple combinations of the basal zone, prefemoral spine, prefemur, and acropodite flange taken together do not indicate close evolutionary relationships: for example, the distinctive combination of a prominent lateral flange and thin acropodite basal zone found in the N. ambulatrix clade can also be found in the N. minor (e.g., N. hippopotamus sp. nov., Fig. 100), N. ohionis (e.g., N. suprema sp. nov., Fig. 69), N. ignis (e.g., N. tenuis sp. nov., Fig. 63), and N. paupertas clades (e.g., N. castra sp. nov., Fig. 20). In contrast, the N. castanea clade is the most morphologically definable group, with all members possessing long, thin acropodites, pronounced prefemoral spines and acropodite tip lateral flanges. There are no known species with long, thin, N. castanea-like acropodites that are not members of this clade.

Figure 130.

Example of morphological convergence between three distantly related species of Nannaria. In all three species, the prefemoral process (pp) arises separate from the prefemoral spine (ps), and crosses the acropodite (ap), despite the three species belonging to separate clades.

Although sufficient for species-level identification within the Nannaria, morphology is a poor indicator of evolutionary relationship above the species level. These results support previous work by Means and Marek (2017) and Means et al. (2021), which showed that morphological homoplasy in gonopod morphology is common within the Xystodesmidae, and suggested that future investigations of evolutionary relationships between millipede taxa should include molecular evidence as an independent dataset.

The phylogeographic history of the minor species group

From the molecular analysis presented here, the Appalachian Mountains—and more specifically, the Blue Ridge Mountains and Valley and Ridge physiographic regions—contain the greatest diversity of Nannaria species (Figs 125–127). Sixteen of the eighteen clades recovered in the phylogeny have members in Virginia, making this state the center of Nannariini diversity (Figs 126–129). It should be noted, however, that collecting efforts have been most intense in Virginia, and it is likely that many species of Nannaria remain undiscovered in other central Appalachian states, especially West Virginia, Kentucky, and Tennessee. Though it is difficult, without fossil evidence, to make statements about the age or movements of Nannaria species over evolutionary time, the placement of the putative relictual N. scholastica sp. nov. as the sister to the rest of the minor species group would suggest that the group originated in western Virginia before spreading throughout the eastern United States. With members of the N. castanea clade found west of the Mississippi, it is likely that Nannaria originated before the last ice age, and may have experienced multiple range contractions and expansions due to shifting environmental conditions. Although there seem to be congruent distributions between unrelated floral and faunal taxa, such as with plethodontid salamanders (Kozak 2017), the seemingly repeated patterns shown by phylogenetic analyses indicate continuous abiotic processes during the Paleogene (splitting, expansion, and retraction) mediated by cooling and warming that may have led to “pseudocongruent” distributions seen in other co-distributed taxa (Manos and Meireiles 2015).

Insight into the abiotic processes that led to the morphological and genetic patterns observed in Nannaria are necessary to help understand the diversification of the group. As a highly dispersal-limited group, environmental conditions would need to be relatively stable for long periods of time for populations to move great distances rapidly, for example in the N. castanea clade, where genetic variation between the eastern and western lineages is less than may be expected based on the nearly 800 km distance between the two lineages (Figs 126, 129). It is unclear why the wilsoni species group is limited to the Appalachian Mountains and does not occur west of the Mississippi River as in the minor species group (Fig. 1). The minor species group taxa are found in a wider variety of habitats than the wilsoni species group taxa, which appear to be tightly constrained to mesic forested areas. Whether the wider distribution of the minor species group is due to an apomorphic physiological adaptation, such as a resistance to desiccation, or some other means is unknown. Alternatively, the wilsoni species group may be a younger clade and had less time to expand its range; that the wilsoni species group is an older clade and experienced greater extinctions is a possibility as well but unlikely. Future studies of Nannaria should seek to understand abiotic factors of their habitat to illuminate the patterns of diversification in the group, and help clarify to what extent ecological factors influence speciation over temporal or other factors influencing evolutionary history.

Nannaria habitat preferences and crypsis

The majority of Nannaria specimens were collected in mesic deciduous forests, often near streams, rivers, or other lotic habitats. In his description of N. ericacea, a member of the wilsoni species group, Hoffman (1949) concluded that both N. ericacea and N. wilsoni were often found in ericaceous habitats, i.e., habitats dominated by an abundance of plants in the family Ericaceae. Rhododendron groves seem to be an excellent indicator of suitable Nannaria habitat, perhaps due to their high moisture requirements and tolerance of water-saturated soils. Adult Nannaria are rarely encountered under the Rhododendron canopy, preferring instead the oak and maple groves interspersing or abutting stands of Rhododendron; while immature Nannaria are frequently encountered directly under Rhododendron. This suggests that the dark and impenetrable understory created by Rhododendron may be an ideal area for Nannaria to molt safely and avoid desiccation during the process of tanning and hardening the cuticle; though the thick, waxy leaves are likely inedible, thereby forcing adults to move outward from Rhododendron coves. Hemlock stands (Tsuga spp.) are a second good indicator of Nannaria presence, likely due to a similar need for a cool, wet environment. Within forested habitats, Nannaria were frequently found beneath the soil at the edges of rocks, tree basal zones, and roots, which may provide shelter and a stable, moist environment. Additionally, Nannaria appeared to prefer sloped hillsides, with little understory, and sandy or loamy soil that drains well and most likely facilitates burrowing.

Unlike the majority of xystodesmid species that are brightly colored and active on the surface (Marek and Bond 2009), Nannaria were often found burrowed 1–2 cm beneath the soil and are drab in coloration, with a lack of strongly contrasting pattern elements. Burrowing, and behavioral concealment, is likely related to their drab coloration as part of an overall cryptic strategy (Ruxton et al. 2018). Dominant color patterns of Nannaria, e.g., small red or white spots on either paranota with a dark background and lack of additional pattern elements such as spots on the prozonite (as in some Apheloria polychroma with four large yellow spots [Marek et al. 2018]), lend themselves to a cryptic appearance. Red light has been shown to be a dominant color in forest sub-canopies and red body coloration appears to be camouflaged in these habitats (Wilson et al. 2007). In addition, the dominant color pattern of Nannaria (red, white, rarely orange) may be non-functional and an ancestral polymorphism that was retained, and perhaps later minimized to small spots. Twenty-three specimens had metatergal stripes, but these were often faint and rarely found on fully tanned adults, suggesting that the stripes may disappear with tanning and hardening of the cuticle in most individuals. Whitehead and Shelley (1992) speculated that the general lack of pigmentation in Nannaria may constitute a form of mimicry with immature apheloriines, which as a result of a lack of a tanned fully sclerotized cuticle appear white, though this hypothesis remains untested. We suggest that the lack of pigmentation is due to their small size, perhaps retention of ancestral color pattern elements, and cryptic lifestyle, where bright coloration would be less beneficial. The similarly-sized Oenomaea pulchella has stripes and not small paranotal spots as in Nannaria (Fig. 3). Specimens of Oenomaea pulchella, like Nannaria species, do not generally produce a noticeable odor of benzaldehyde advertising their production of hydrogen cyanide. A larger comparative analysis of cyanide glands and defensive chemicals of O. pulchella, Nannaria species, and related members of the Xystodesmidae may illuminate the role that color and associated chemical defense plays in the Nannariini.

Conclusions

Since the description of the genus nearly a century ago, Nannaria has received little taxonomic attention beyond the occasional species description. Here, we provide the first revision of the Nannariini using molecular evidence from both genera (Oenomaea and Nannaria). We reveal the existence of two distinct clades—the minor and wilsoni species groups, and describe 35 new species within the highly species diverse and widespread minor species group. In an upcoming revision of the wilsoni species group, led by DAH, 17 new species will be described. With 52 new species of these cryptically colored and often difficult-to-find polydesmidan millipedes, the Nannaria represent a taxon that is among the least known α-taxonomically of any animal group in North America. From this analysis, we provide evidence of 35 undescribed species within the minor species group and 11 additional putative species. We have shown that Nannaria has many instances of morphological convergence confounding morphology-based species concepts. This study shows a 10% mean difference amongst species in the COI barcoding region, and establishes a large barcoding database of the genus that provides groundwork for future investigations of speciation and diversity within the Nannaria. Over 70% of species within the Nannaria minor species group are micro-range endemics, meaning they have distributions of less than 1000 km² (Harvey 2002; Means and Marek 2017). The highest density of known species diversity occurs in Virginia, primarily in the Ridge and Valley ecoregion. This topographically complex area is an exceptional repository of high species richness and relative rarity of taxa (Stein et al. 2000). Deforestation, surface mining, and habitat loss disproportionately threaten these groups with narrow endemism, strict habitat requirements, and low dispersal rates such as Nannaria. The loss of micro-range endemics and other rare species permanently hinders our ability to conserve biodiversity, reconstruct phylogenetic history, and study diversity of the life on the planet.

Acknowledgements

This research was supported by a National Science Foundation Advancing Revisionary Taxonomy and Systematics grant to PEM (DEB # 1655635). Appalachian Energy and Power provided resources for a survey of forest sensitive millipedes in Jefferson National Forest in Bland and Wythe counties, Virginia (2016, 2017) that lead to the discovery of N. ignis, N. botrydium, and N. tenuis. We thank natural history institutions and curators who sent us material for this study, including Kal Ivanov and Liberty Hightower (VMNH), Bronwyn Williams and Megan McCuller (NCSM), Hannah Wood and Dana DeRoche (NMNH), Jason Dunlop and Anja Friederichs (MFN), and Paul Skelley and Felipe Soto-Adames (FSCA). We are grateful to Bruce Snyder for his contribution of Nannaria tennesseensis specimens, and to Curt Harden, Matt Kasson, and Bill Shear for their contributions of specimens which led to the discovery of multiple undescribed species. We thank Kathlyn Means, Charity Hall, Patricia Shorter, Derek Krall, Anne Chazal, and Katy Lawler for their assistance with fieldwork. We also thank Bill Shear, Sergei Golovatch, and Dragan Antić for their revisions to previous versions of the manuscript. We thank Alessandro Minelli for reviewing our species names and for insights into the Latin language. We are grateful to the various organizations and agencies which issued scientific collection permits for this study, including: Robert Cherry at the U.S. National Park Service (permits #BLRI-2014-SCI-0033, GRSM-2016-SCI-2369, GRSM-2019-SCI2369, SHEN-2016-SCI-0018); Jesse Overcash, Jefferson National Forest, Virginia; Cincinnati Museum Center, Edge of Appalachia Preserve; Crane Hollow Preserve; Indiana University, Griffy Woods Preserve; Maryland Department of Natural Resources (Project# 2016DNR119); Maryland State Parks; Georgia State Parks (permits #132016, 142018); Kentucky Department of Parks (permit #1708); Kentucky State Nature Preserves Commission (permit #2018_0234); Lexington-Fayette Urban County Government Division of Parks and Recreation; Duke University, Office of Duke Forest (project# R1516-413); North Carolina Division of Parks and Recreation (permit# 2018_0234, 2017_0169); South Carolina Department of Parks, Recreation & Tourism, State Park Service (permit# N-26-17); Tennessee Wildlife Resources Agency (permit #3914, 1702); Tennessee Department of Environment and Conservation (permit #2017-008); West Pennsylvania State Parks (permit# 2017-64); Pennsylvania Department of Conservation and Natural Resources, Bureau of Forestry (permit# SFRA-1724); Commonwealth of Virginia Department of Forestry; Virginia Department of Game and Inland Fisheries (permit# 056958); and the West Virginia Division of Natural Resources (permits # 2018.277, 2016.238). Finally, we are eternally grateful to the late Richard Hoffman (VMNH, Radford University) for his preliminary work on Nannaria and his tireless efforts to advance our understanding of myriapodology.

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Supplementary materials

Supplementary material 1

Suppl. material 1

Means JC, Hennen DA, Marek PM

Data type: List of genes

Explanation note: List of gene regions explored for use in taxonomic analyses of Nannaria.

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.

Download file (15.17 kb)

Supplementary material 2

Suppl. material 2

Means JC, Hennen DA, Marek PM

Data type: List of taxa & NCBI accession numbers

Explanation note: List of taxa used in molecular analyses, organized alphabetically by genus and then species. Acc# refer to the NCBI Genbank database. All specimens available from the corresponding author by request and stored in the Virginia Tech Insect Collection, Blacksburg, Virginia, USA.

Download file (58.80 kb)

Supplementary material 3

Suppl. material 3

Means JC, Hennen DA, Marek PM

Data type: Phylogenetic

Explanation note: Molecular phylogeny of Nannaria.

Download file (2.05 MB)

Supplementary material 4

Suppl. material 4

Means JC, Hennen DA, Marek PM

Data type: Phylogenetic

Explanation note: Individual gene phylogenies of Nannaria.

Download file (11.36 MB)

Supplementary material 5

Suppl. material 5

Means JC, Hennen DA, Marek PM

Data type: Morphological scoring matrix

Explanation note: Morphological scoring matrix for Nannaria minor species group.

Download file (58.61 kb)

Supplementary material 6

Suppl. material 6

Means JC, Hennen DA, Marek PM

Data type: Key to morphological matrix

Explanation note: Key to morphological characters used to score Nannaria.

Download file (20.20 kb)

Supplementary material 7

Suppl. material 7

Means JC, Hennen DA, Marek PM

Data type: occurrences

Explanation note: Materials examined.

Download file (161.68 kb)

Supplementary material 8

Suppl. material 8

Means JC, Hennen DA, Marek PM

Data type: Species group distributions

Explanation note: A .kmz file of Nannaria minor species group distributions for use in programs such as Google Earth (Alphabet Inc., Mountain View, California).

Download file (42.92 kb)