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We describe three new genera and four new species of small, litter-dwelling millipedes from the states of Oregon and Washington, USA: Miniaria ramifera, n. gen., n. sp., Miniaria richarti, n. gen., n. sp., Tigraria oregonensis, n. gen., n. sp., and Kingaria prattensis, n. gen., n. sp. Some of the unusual characters of these species are discussed, including a new type of sensory array on the third tarsus of males and a newly observed mandibular gland. 

The millipede fauna inhabiting deep soil are poorly known. They are small and threadlike, slow moving, lacking pigmentation, and rarely encountered due to their obscure underground way of life. One family, the Siphonorhinidae, encompasses four genera and 12 species in a fragmentary distribution in California, southern Africa, Madagascar, the Malay Archipelago, and Indo-Burma. The family is represented in the Western Hemisphere by a single genus, Illacme Cook & Loomis, 1928 from California, with its closest known relative, Nematozonium filum Verhoeff, 1939, from southern Africa. A new species of this family is documented from soil microhabitats in the Los Angeles metropolitan area, Illacme socal Marek & Shear, sp. nov. Based on this discovery and the recent documentation of other endogean millipede species, we show that these grossly understudied subterranean fauna represent the next frontier of discovery. However, they are threatened by encroaching human settlement and habitat loss, and conservation of this species and other subterranean fauna is of high importance. 

The following new genera and species of Striariidae are described from the states of California, Oregon and Washington: Nototrisaria ornata n. gen., n. sp., Maraplia napa n. gen., n. sp., Maraplia chico n. gen., n. sp., Maraplia schusteri n. gen., n. sp., Lamparia curryensis n. gen., n. sp., Lamparia bentonensis n. gen., n. sp., Lamparia pratensis n. gen., n. sp., Lamparia millicoma n. gen., n. sp., Rampalia cheathamensis n. gen., n. sp., Plaramia arcata n. gen., n. sp., Plaramia johnsonae n. gen., n. sp., Ralampia complexa n. gen., n. sp., and Ralampia filamentosa n. gen., n. sp.. The new species are all very small millipedes, 5.5 mm or less in length and with two or three ommatidia on each side of the head or blind. The majority have the sixth metazonital crest larger than the other crests and extended horizontally, so that the animals have the general appearance of having paranota. A new key to striariid genera is presented. 

Although most binaural organisms locate sound sources using neurological structures to amplify the sounds they hear, some animals use mechanically coupled hearing organs instead. One of these animals, the parasitoid flyOrmia ochracea(O. ochracea), has astoundingly accurate sound localization abilities. It can locate objects in the azimuthal plane with a precision of 2°, equal to that of humans, despite an intertympanal distance of only 0.5 mm, which is less than$1/100$th of the wavelength of the sound emitted by the crickets that it parasitizes.O. ochraceaaccomplishes this feat via mechanically coupled tympana that interact with incoming acoustic pressure waves to amplify differences in the signals received at the two ears. In 1995, Mileset aldeveloped a model of hearing mechanics inO. ochraceathat represents the tympana as flat, front-facing prosternal membranes, though they lie on a convex surface at an angle from the flies’ frontal and transverse planes. The model works well for incoming sound angles less than$\pm {30}^{\circ }$but suffers from reduced accuracy (up to 60% error) at higher angles compared to response data acquired fromO. ochraceaspecimens. Despite this limitation, it has been the basis for bio-inspired microphone designs for decades. Here, we present critical improvements to this classic hearing model based on information from three-dimensional reconstructions ofO. ochracea’s tympanal organ. We identified the orientation of the tympana with respect to a frontal plane and the azimuthal angle segment between the tympana as morphological features essential to the flies’ auditory acuity, and hypothesized a differentiated mechanical response to incoming sound on the ipsi- and contralateral sides that depend on these features. We incorporated spatially-varying model coefficients representing this asymmetric response, making a new quasi-two-dimensional (q2D) model. The q2D model has high accuracy (average errors of under 10%) for all incoming sound angles. This improved biomechanical model may inform the design of new microscale directional microphones and other small-scale acoustic sensor systems.

 

Petra sierwaldae, n. gen., n. sp. is described from males and females collected at four localities in Kootenai County, Idaho, USA. The new genus is defined by its unique gonopod and ninth leg anatomy, a notched collum and single ommatidium on each side of the head, and lacking third coxal flasks as well as having modified fifth coxae. 

The millipede genus Cherokia Chamberlin, 1949 is a monospecific taxon, with the type species Cherokia georgiana (Bollman, 1889). The last revision of the genus was made by Hoffman (1960) where he established three subspecies. Here we used molecular phylogenetics to assess the genus and evaluate whether it is a monophyletic group, and if the subspecies are each monophyletic. We included material from literature records and three natural history collections. Newly collected samples were obtained through a citizen science project. Morphological characters underlying subspecies groups—the shape of the paranota, body size, and coloration—were evaluated. A molecular phylogeny of the genus was estimated based on DNA sequences for seven gene loci, and a species delimitation analysis was used to evaluate the status of the subspecies. The documented geographical range of Cherokia in the United States was expanded to include a newly reported state record (Virginia) and about 160 new localities compared to the previously known range. Morphological characters, which included the shape of the paranota and body size that had been historically used to establish subspecies, showed clinal variation with a direct relationship with geographical distribution and elevation, but not with phylogeny. Coloration was highly variable and did not accord with geography or phylogeny. The phylogeny recovered Cherokia as a monophyletic lineage, and the species delimitation test supported the existence of a single species. The subspecies Cherokia georgiana ducilla (Chamberlin, 1939) and Cherokia georgiana latassa Hoffman, 1960 have been synonymized with Cherokia georgiana . The molecular and morphological evidence showed that Cherokia is a monospecific genus with the sole species, Cherokia georgiana , being geographically widespread and highly variable in its morphology. 

Although many new species of the millipede genus Nannaria Chamberlin, 1918 have been known from museum collections for over half a century, a systematic revision has not been undertaken until recently. There are two species groups in the genus: the minor species group and the wilsoni species group. In this study, the wilsoni species group was investigated. Specimens were collected from throughout its distribution in the Appalachian Mountains of the eastern United States and used for a multi-gene molecular phylogeny. The phylogenetic tree recovered Nannaria and the two species groups as monophyletic, with Oenomaea pulchella as its sister group. Seventeen new species were described, bringing the composition of the wilsoni species group to 24 species, more than tripling its known diversity, and increasing the total number of described Nannaria species to 78. The genus now has the greatest number of species in the family Xystodesmidae. Museum holdings of Nannaria were catalogued, and a total of 1,835 records used to produce a distribution map of the species group. Live photographs, illustrations of diagnostic characters, ecological notes, and conservation statuses are given. The wilsoni species group is restricted to the Appalachian region, unlike the widely-distributed minor species group (known throughout eastern North America), and has a distinct gap in its distribution in northeastern Tennessee and adjacent northwestern North Carolina. The wilsoni species group seems to be adapted to mesic microhabitats in middle to high elevation forests in eastern North America. New species are expected to be discovered in the southern Appalachian Mountains. 

Two new genera and species, Stegostriaria dulcidormus (Trisariinae) and Kentrostriaria ohara (Striariinae), are described from Oregon and Idaho, respectively. The new species are distinct from any other striariids, and indeed from any other chordeumatidan millipedes, in having the second dorsal crests greatly enlarged, giving them a fanciful resemblance to stegosaurid dinosaurs. In spite of this similarity, the two species are so distinct they cannot be accommodated in the same genus or the same subfamily. The exaggerated metazonital crests are therefore attributed to convergent evolution. We present a key to the genera of the family Striariidae. 

A new species, Amplaria oedipus, is described from Oregon, USA. Males of the new species have greatly inflated pyriform tarsi on the 5th and 6th legpairs, which is a secondary sexual modification previously not known from any other millipede.