Meet the latest member of Hokie Nation, a newly discovered millipede

The millipede Nannaria hokie that was discovered on the campus of Virginia Tech
The millipede Nannaria hokie was recently found on Virginia Tech’s campus

In our new paper, we named the species, Nannaria hokie, which we discovered on Virginia Tech’s campus. The newest species in Hokie Nation lives by the Duck Pond and Stadium Woods on Virginia Tech’s campus, and a handful of forests surrounding Blacksburg. Nowhere else on Earth.

As detritivores, millipedes feed on decaying plant material, thereby fragmenting detritus into smaller pieces fostering later colonization by bacteria and fungi. This organic input aerates and conserves the soil, and the process releases nitrogen, carbon, simple sugars, and other nutrients back into the biosphere.

Despite their antiquity and important role as detritivores, known millipede species diversity tremendously lags behind estimated global diversity. For example, there are more than 50 undescribed species of Nannaria in the eastern United States alone, and globally an additional 3,000 – 80,000 species of Diplopoda when tropical locales and other poorly sampled regions are included.

The larger group that contains Nannaria exhibits fascinating biological characteristics. The millipede family Xystodesmidae includes notable taxa such as bioluminescent species in California, Motyxia spp.; Müllerian mimics in Appalachia, apheloriine spp.; the train millipede in Japan, Parafontaria laminata, whose aggregations of 311 individuals m-2 have obstructed trains; the giant 8-cm long armadillo millipede in Mexico, Rhysodesmus dasypus; and the cherry millipede (Apheloria virginiensis corrugata) able to generate hydrogen cyanide in an amount 18 times that necessary to be lethal to a pigeon-size bird.

This research was supported by a National Science Foundation Advancing Revisionary Taxonomy and Systematics grant to P. Marek (Division of Environmental Biology, Systematics and Biodiversity Sciences #1655635).

Means, J.C., D.A. Hennen, T. Tanabe, P.E. Marek. 2021. Phylogenetic systematics of the millipede family Xystodesmidae. Insect Systematics and Diversity, 5: 1-26. https://doi.org/10.1093/isd/ixab003 [Open access]

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The cherry millipede

We made this short movie about the cherry millipede, Apheloria virginiensis corrugata. Known from forests in the eastern U.S., the cherry millipede oozes cyanide and feeds on decaying leaves and other detritus.

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Fieldwork in Japan

Earlier this month, I returned from two weeks of fieldwork in Japan where I collected millipedes and visited my colleague Dr. Tsutomu Tanabe of Kumamoto University. The trip was a wonderful combination of productive fieldwork, great discussions with Tsutomu and his students, and delicious ramen. The field trip was organized to collect millipedes of the family Xystodesmidae, which is the topic of my National Science Foundation grant (DEB #1655635). I started in Sapporo (Hokkaido Prefecture), then travelled to Tsukuba (Ibaraki Prefecture), Kumamoto (Kumamoto Prefecture), and finally Mount Unzen (Nagasaki Prefecture).


The xystodesmid millipede Levizonus takakuwai

Millipedes of the family Xystodesmidae occur in the Holarctic realm (more specifically in North America, East Asia, the Russian Far East, China, Northern Vietnam, and the Mediterranean). My laboratory’s research at Virginia Tech focuses on this family. It’s always exciting to find xystodesmid millipedes because they’re brightly colored and smell like cherries. We continue to discover new species in Appalachia, and even found a new species on Virginia Tech’s campus. Notably, xystodesmids in California are nocturnal and some glow in the dark. In contrast, xystodesmids in Appalachia tend to be diurnal (day active). Because I am familiar with xystodesmids and have collected them from throughout the U.S., it’s been a goal of mine to collect them in Japan.

The first genus on the list was Levizonus, which has eight species. Five of them live in the Russian Far East, one in Korea, and two in Japan. The two Japanese species overlap in their distributions in Sapporo, and include Levizonus takakuwai (Verhoeff, 1941) and Levizonus montanus (Takakuwa, 1941). Levizonus takakuwai occurs in southwestern Hokkaido with a well-documented population at Maruyama Park in Sapporo. This is where we looked first. The previous week, I collected xystodesmids in Virginia, which are day active, so we started fieldwork for L. takakuwai at Maruyama Park in the early morning.


Sapporo from Mount Maruyama, Hokkaido Prefecture

Maruyama Park is composed of deciduous forests with maples, magnolias, and even poison ivy, and stinging nettles, which reminded me of the flora of Appalachia. We spent the entire morning, took a break at noon for ramen, and continued in the early afternoon searching for L. takakuwai. Even with geographical coordinates from Tsutomu, we could not find it. After hiking up to near the peak of Mount Maruyama, I finally found a male L. takakuwai, followed by a female specimen that was collected by Charity. The site was a deciduous forest with lots of maple, and knee-high bamboo growing in large patches. During a hour-long search, two millipedes were found beneath decaying leaves and other moist detritus. Taking note of their scarcity and remembering Tsutomu’s observation that they’re burrowed underground during the day, we planned to return at night with an ultraviolet flashlight. Xystodesmid millipedes are fluorescent and glow green when illuminated with ultraviolet light. I’ve collected xystodesmids in California where they are nocturnal and practically unfindable during the day, but easily found during the night.


Deciduous forest in Maruyama Park, Sapporo


Jack-in-the-pulpit, Maruyama Park

At night, we walked across the grassy field at the edge of Maruyama Park into the forest, turned on our UV flashlights, and immediately found a fluorescent green L. takakuwai emerging from the ground. Then, we found a second, a third, and eventually encountered about 50 millipedes. Levizonus takakuwai, like Californian xystodesmids, is nocturnal and fluorescent. The defense secretions of L. takakuwai are similar to Californian xystodesmids too, and they have less of a cherry odor and smell more like the burnt rubber aroma of Chonaphe, Harpaphe, Xystocheir, and other species from the U.S. West Coast. Having found two millipedes in six hours of collecting during the day and 50 millipedes in 15 minutes at night, we spent almost the entire trip collecting at night.

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Fieldwork in Japan (part 2)

In Tsukuba, which is about 50 km northeast of Tokyo, we met Tsutomu and his student Ryosuke Kuwahara. I really enjoyed meeting Ryosuke because he’s very enthusiastic about natural history and takes wonderful photographs of xystodesmid millipedes (here’s a link to his website). That evening, we visited Seiryuji Temple in Tsuchiuria and collected millipedes in a nearby forest. When we walked into the forest, a Luna moth flew in front of us and landed on a tree.


The xystodesmid millipede Parafontaria tonominea

Often, finding millipedes is sporadic, and we spent about 20 minutes before we encountered the first xystodesmid, Parafontaria tonominea (Attems, 1899). That evening, we also found Parafontaria ishiii Shinohara, 1986 and Xystodesmus martensii (Peters, 1864). (Notably in 1895, Cook established the genus Xystodesmus and the family Xystodesmidae based on X. martensii, a species originally described by Peters in 1864 as Polydesmus martensii). With the moth and millipedes, there were other fascinating organisms in the forest including a gray hand-sized centipede, Scolopendra japonica C.L. Koch, 1878.


The xystodesmid millipede Parafontaria ishiii

It became clear that night collecting was the most productive. For example in Sapporo, we found two millipedes in six hours during the day and 50 millipedes in 15 minutes at night. Consequently, we spent our days planning fieldwork, resting for night-collecting, looking at Tsutomu’s natural history collections, and tending to the live millipedes (and of course eating a good amount of ramen).


Xystodesmus martensii

The second night in Tsukuba, we travelled through a network of rice plantations to Mount Hōkyō-san and collected in the cherry, chinquapin, and bamboo forest that covers the mountain. We hiked a couple miles to a site about 400 meters in elevation that was just below the peak of Hōkyō-san. At 8PM, P. ishiii, X. martensii (and another millipede that Ryosuke and Tsutomu will soon share), started to emerge from their subterranean hiding places. An hour later, we encountered more. At the end of the night, we found about one hundred xystodesmid millipedes. We collected about 20 individuals to bring to Virginia Tech for DNA sequencing and molecular phylogenetics. At the moment, we have sequenced about 239 of the family’s 393 species (and nearly all of the taxon’s 62 genera). We’ve done this to estimate the family’s evolutionary history in a phylogeny. The phylogeny of Xystodesmidae is important as a guide for describing species and tracing the evolution of mimicry and features like chemical defenses.


Mount Hōkyō-san, Ibaraki Prefecture


Left to right: Tsutomu Tanabe, Paul Marek, Charity Hall, and Ryosuke Kuwahara

Walking down Mount Hōkyō-san, we enjoyed being entomologists and encountered large Carabus beetles, sawflies, and several huge S. japonica centipedes hanging out on the sides of trees. Ryosuke pointed out a handsome mamushi on the trail near the rice field, Gloydius blomhoffii (H. Boie, 1826), which is species of viper.


The xystodesmid Parafontaria erythrosoma

The third night in Tsukuba, we collected at a lower elevation (89 m) forest near Kashima Shrine in Mito. Again, as a testament to the patchiness of millipede encounters, we found little in the first 30 minutes. But once we encountered a suitable patch, we found lots of millipedes. The first xystodesmid millipede that we found was Parafontaria erythrosoma (Takakuwa, 1942). A handsome millipede, which as its name implies, has a red (“erythro”) body (“soma”). With P. erythrosoma, we found X. martensii again, which is a very widespread species with a considerable amount of variation (Tanabe and Shinohara, 1996). Ryosuke broke open a decaying log and found a species of Hyleoglomeris, which is a glomerid millipede capable of rolling into a ball. We encountered other millipedes including the haplodesmid Eutrichodesmus peculiaris (Murakami, 1966); the polydesmid Epanerchodus lacteus Shinohara, 1958; and a species of the paradoxosomatid Cawjeekelia Golovatch, 1980. Notably, E. peculiaris has maternal care of eggs, which is a rare behavior in millipedes, and an incredibly rare behavior in millipedes of the order Polydesmida (Murakami, 1972).


The haplodesmid millipede Eutrichodesmus peculiaris

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