Chilopoda

2023 Collecting Summary

Local mixed forest habitat, showing the grandeur of winter collecting. Photo taken 16 January 2023.

Continuing my trend from 2021 and 2022, it’s time for the summary of my myriapod collecting during 2023! I made a concerted effort last year to collect less so that I could focus on other things (mainly writing, curation of my specimens, and job hunting), but I still found a number of interesting and rare species. An added bonus from last year is that a number of friends and colleagues sent me specimens from far-flung areas, which added some exceptional species to my collection. I want to give a particular thank you to Lance Andrew, Joe Girgente, Adam Haberski, Curt Harden, Brendan O'Loughlin, and Carol Tingley for sending me their specimens.

Thanks to the specimens sent to me by colleagues, the collecting map spans the continent! The Alaska locality especially sticks out, and represents some beautiful Parajulidae.

Map of 2023 collecting localities

 I totaled 37 collecting events in 2023, spanning only two states: Virginia and Ohio. All the dots on the map from outside those states are specimens sent to me by other people, and those specimens really augmented my collection last year! The numbers I’m using for the rest of this post include the specimens sent by colleagues.

Total myriapod specimens collected in 2023

All in all, my colleagues and I collected 1,450 myriapods in 2023, as shown in the above graph (blue bars, left axis). The total number of collection events are represented by the orange line, with numbers reflected on the right axis. There were a total of 81 collection events last year: I did 37 of them, while my colleagues conducted 44. Again, the value of others collecting myriapods for me is immense!

These collections included 109 species from 36 families, beating 2022’s count of 88 species from 30 families. Those 109 species include 71 genera, including a number of unique genera I didn’t have in my collection before. Last year I wrote about wanting to top 100 species in a year, and I achieved that goal!

Since this is my third year of tracking this data, I can include my previous years of data to see how my collecting habits have differed.

Total specimens collected monthly, 2021-2023

The above graph shows the total number of specimens collected during each month from 2021-2023. There’s quite a bit of variation there, mostly reflecting collecting effort. The totals from 2021 are quite low until September 2021, since that’s when I started using my homemade Berlese funnels. The main takeaway here is that collection effort is typically the most important determiner of how many specimens I collect each month, with local weather playing a part as well. Totals from November through February tend to be lower than Spring through Fall, but there are, of course, exceptions. Additionally, if you want to get the full picture of your local myriapod diversity, you’ll miss unique cold weather-active species if you don’t collect between November and February.

I’ve collected throughout the year enough now to recognize a few distinct myriapod faunal groups. There are species like Nadabius pullus (Lithobiomorpha: Lithobiidae) that you can find easily throughout the year. Other groups like the Apheloriini (Polydesmida: Xystodesmidae) are most active during the spring and fall. And then there are other species, like the genus Garibius (Lithobiomorpha: Lithobiidae) and many Chordeumatida, which are active during late fall through early spring. Figuring out which species fit these various seasonal patterns (and which ones don’t!) is one of the most fulfilling parts of collecting myriapods for me personally. The cold weather-active species in particular are some of the most exciting to find, as these are typically rare in collections, despite their ubiquity during their active season. Besides, you pretty much have the entire woods to yourself while searching for them. It’s quite pleasant to grub around in the dirt in brisk weather.

Another fun thing to check the yearly data for is the number of unique species per month, shown in the graph below:

Number of species collected monthly, 2021-2023

These monthly totals are somewhat less variable than the total specimens collected each month, but collecting effort still has a strong influence (see November 2023!). These totals can be highly influenced by colleagues sending me specimens, particularly if they’re based outside of Virginia. The standout from this chart is April 2023, with the most species I’ve ever collected in a month, at 51! That was also the month with the highest total specimens, at 297 (see previous graph).

Diving into the 2023 species-specific data, there are similar trends to my 2022 collecting. Once again, the centipede Nadabius pullus (Lithobiomorpha: Lithobiidae) was my most-collected species, represented by 154 specimens, less than last year’s count of 195 specimens. The full top ten list is:

Nadabius pullus (154), Nampabius sp 1 (90), Arctogeophilus umbraticus (70), Oxidus gracilis (70), Hanseniella sp (63), Strigamia branneri (60), Typhlobius sp (52), Ptyoiulus impressus (49), Chaetaspis sp (40), Paitobius zinus (36)

The four names in bold are species that weren’t on last year’s list. Collectively, these 10 species account for 51% of all myriapods collected in 2023. At the other end of the spectrum, 32 species are represented by 1 or 2 specimens, making up only 3% of the total specimens collected, but 29% of the total species collected. Sometimes those singletons pack a big punch diversity-wise.

I was able to find a number of lifer species in 2023, including some I had been after for years! One of my favorites was the centipede Agathothus gracilis (Geophilidae: Linotaeniinae), which has only been reported a few times from Tennessee and southwest Virginia. It’s one of only two genera of Linotaeniinae in eastern North America, and is far rarer than the common Strigamia. I wasn’t sure I’d be able to recognize a specimen of A. gracilis, but after seeing the forcipules, it’s unmistakeable—they’re flattened and sharp, and lack the tarsungular tooth of Strigamia, making A. gracilis immediately recognizable.

Agathothus gracilis, head and anterior segments, ventral view.

By far my most surprising find of 2023 was a liliputian geophilomorph, the enigmatic Nannarrup hoffmani. Funnily enough, 2023 marks the species’ 20th birthday, having originally been described from Central Park in New York City in 2003. I found half a dozen specimens in southwest Virginia, while looking for a different species entirely! The species itself is native to an unknown part of east Asia, and only recently has the genus been found in its native range (see Tsukamoto et al. 2022). This is a new state record for the species (and the family) and makes it likely that N. hoffmani will turn up in additional states, if searched for.

Nannarrup hoffmani: live specimen in soil matrix (left) and ventral view of forcipules (right).

Another geophilomorph I was happy to find was Strigamia hoffmani, a species only described in 2009 in honor of Richard Hoffman’s 80th birthday. It’s a small species that can be difficult to separate from Virginia’s other species of Strigamia, and is only known from Tazewell and Roanoke counties. In 2023, I found it in two additional counties: Grayson and Giles. The key to finding this one may be searching areas over 1,000 m in elevation, suggesting a more boreal-adapted species. It still hasn’t been found outside of Virginia, so get searching, those of you in surrounding states!

Last year brought me my first specimen of Lamyctes pius (Lithobiomorpha: Henicopidae), likely a synonym of L. emarginatus (see Shear 2018). Chamberlin described a number of species within the genus Lamyctes, but the evidence points to them being an introduced species in North America. Native henicopids are rare here, as the family is mainly confined to the southern hemisphere.

Lamyctes pius (Lithobiomorpha: Henicopidae) head, dorsal view (left) and ventral view (right)

Moving on to the millipedes, there were a number of surprises this year. Down in Wise County, I found my first sphaeriodesmid in Virginia: Desmonus earlei!

Desmonus earlei, lateral view

This is a slight range extension and only the second record for Virginia that I know of; previously it had been recorded from Lee County. There’s another Desmonus species that ranges from Missouri south to Texas and adjacent Mexico, Desmonus pudicus. Both species are quite small, only about 5 mm across when curled into the loose spiral pictured above, and maybe a centimeter fully extended. Despite their small size, they’re immediately recognizable by the large dorsal tubercles on their rings, and by their coating of dirt. While Desmonus is the only representative that occurs in the US, the family Sphaeriodesmidae is more diverse in Mexico and down through South America, and it’s always a treat to see one of these species.

Desmonus earlei gonopods, posterior view (left) and lateral view (right)

There’s one more polydesmidan millipede I want to highlight, and it was by far the biggest millipede find of my year. I was meeting a friend at a park in Wythe County, and while I was waiting, I scratched around in the leaf litter near a stream to see what I could find. Within five minutes, I found some small cherry millipedes (Xystodesmidae) and did a double take:

Rhysodesmus restans, dorsal view of live specimen

“No way,” I whispered to myself. This was Rhysodesmus restans, something of an artifact in Appalachia. Almost all species of Rhysodesmus are further south, in Texas, Mexico, and Central America. But there are a few still around, left behind in the Appalachian Mountains. I’d been searching for this species for years at this point, and always much further away, down in Washington County (the type locality). Even better: they were abundant here! I collected a number of specimens and left more behind, still in disbelief at my luck.

The final species I’ll highlight is a hidden gem that I’d actually collected multiple times in the past few years, but which I had misidentified. This was pointed out to me by my colleague Pierre-Marc Brousseau, to whom I’m grateful! A small millipede in the order Julida, the species Okeanobates americanus is one I had read about, but that was the limit of my experience with it.

Okeanobates americanus, lateral view

It turns out that I had unknowingly collected this species almost yearly since 2020! I had it misidentified as an unknown species of Blaniulidae, but Pierre-Marc noticed the rectangular eye patch in one of my iNaturalist photos and corrected me. The species ranges from North Carolina up through Quebec, Canada, but is uncommon. It seems to favor life within logs, as I’ve never found it in leaf litter. I included more information about the species and the Okeanobatidae in a blog post back in November, so check that out to learn more. It’s not often that I get to add a new family to my collection, what a thrill!

That about wraps up my 2023 review. I found some superb species last year, and much of that was thanks to friends and colleagues, to whom I’m indebted for sending me specimens.

Possibly the strangest thing I did last year was that I accidentally ended up writing a key to all eastern geophilomorph centipedes. I meant to focus on Lithobiomorpha in 2023, but got hooked on those weird little soil centipedes. If you’re interested in using the key or sending me geophilomorphs from your region, please get in touch. It will be a little while before I publish it, but I’m aiming to write up centipede resources in the next few years to focus attention on the North American fauna, especially in the east.

This post has gone on long enough, so I’ll end it here with a photo comparison of the new litter sifter I bought in 2023. It replaces my previous sifter, which I’d been using for almost a decade. Can you tell which is which?

Further reading:

Chamberlin RV (1912) The Geophiloidea of the southeastern states. Bulletin of the Museum of Comparative Zoology at Harvard College 54: 405–436. https://www.biodiversitylibrary.org/page/2815547

Enghoff H (1979) The millipede genus Okeanobates (Diplopoda, Julida: Nemasomatidae). Steenstrupia 5: 161–178.

Hoffman RL (1998) An Appalachian species of Rhysodesmus (Polydesmida: Xystodesmidae: Rhysodesmini). Myriapodologica 5: 77–83. https://www.biodiversitylibrary.org/page/52243216

Pereira LA (2009) A new dwarf species of the genus Strigamia Gray, 1843 from the southern Appalachian Mountains of western Virginia (Chilopoda: Geophilomorpha: Linotaeniidae). Virginia Museum of Natural History Special Publication 16: 209–222.

Shear WA (2018) The centipede family Anopsobiidae new to North America, with the description of a new genus and species and notes on the Henicopidae of North America and the Anopsobiidae of the Northern Hemisphere (Chilopoda, Lithobiomorpha). Zootaxa 4422: 259–283. https://doi.org/10.11646/zootaxa.4422.2.6

Shelley RM (2000) Revision of the milliped subfamily Desmoninae (Polydesmida: Spheriodesmidae). Myriapodologica 6: 27–54. https://www.vmnh.net/content/vmnh/uploads/PDFs/research_and_collections/myriapodologica/myriapodologica_v6_5.pdf

Tsukamoto S, Shimano S, Eguchi K (2022) Two new species of the dwarf centipede genus Nannarrup Foddai, Bonato, Pereira & Minelli, 2003 (Chilopoda, Geophilomorpha, Mecistocephalidae) from Japan. ZooKeys 1115: 117–150. https://doi.org/10.3897/zookeys.1115.83946

The tergital projections of Bothropolys multidentatus (Lithobiidae: Ethopolyinae)

The most imposing stone centipede you’ll find in the eastern US is the charismatic Bothropolys multidentatus (Lithobiidae: Ethopolyinae). Exclusively found in dead wood microhabitats, it’s a gorgeous chestnut-brown to orange centipede that reaches lengths of up to 30 millimeters—a true beast! There are only a few other lithobiomorphs of such size in our area (really just limited to Lithobius, Eulithobius, Neolithobius, and Zygethopolys).

Live Bothropolys multidentatus under my microscope.

It’s one of our two eastern Ethopolyinae, the other being Zygethopolys atrox. Both are immediately separable from other Lithobiidae and Henicopidae by the ventral pore fields on their posterior coxae: species in the Ethopolyinae have multiple rows of pores, rather than a single row.

Within eastern Ethopolyiinae, B. multidentatus is widespread, while Zygethopolys atrox is only known from Cumberland Falls State Park in Kentucky. So if you find a stone centipede with multiple rows of coxal pores, it’s likely to be B. multidentatus. The map below shows the distribution of the genus Bothropolys, and all the eastern dots represent B. multidentatus.

As the species name indicates, B. multidentatus has a large number of prosternal teeth. The exact number is variable, ranging between 6 and 9 teeth in adults. At the lateral edges of the prosternal teeth, there’s a stout seta called the porodont. In Z. atrox, the porodont is inserted in the line of prosternal teeth, separating an outer tooth from the inner line of teeth.

Forcipular coxosternum of Bothropolys multidentatus,. Notice the stout porodont at the outer edge of the prosternal teeth. This specimen has 6+6 prosternal teeth.

Something that caught my eye recently is the production of the tergites in B. multidentatus. In the Lithobiomorpha, some species have triangular projections at the posterior corners of the tergites. When those projections are present, the tergites are said to be produced. When they’re absent, the tergites are not produced. In the first photo on this post, you can see that tergites 6, 7, 9, 11, and 13 are produced. This matches how Chamberlin (1925) describes the tergites. However, if you look at tergite 4 in my photo, you can see it’s also produced. I checked a few of my other photos and noticed some variation in the production of tergite 4, so I went on to check all the specimens of B. multidentatus in my collection. All my specimens are from the northeastern United States, so I was only able to check specimens from a small part of the range (Ohio, Pennsylvania, West Virginia, and Virginia).

I noticed three different character states for the 4th tergite: not produced, slightly produced (small triangular projections), and fully produced (strong triangular projections). As I checked my specimens, I made notes and took photos to directly compare later. I examined 29 specimens in total; here’s a map of the character states for all my specimens:

Distribution of Bothropolys multidentatus specimens from my collection. Dots are color coded by the character state of the 4th tergite: not produced, slightly produced, and produced.

After making the map, I organized the photos that I took and compared them to my written notes. I wasn’t completely satisfied with how I classified some specimens as having only slightly produced 4th tergites vs. produced, and concluded that I should simply categorize the specimens as having 4th tergites not produced or produced (including slightly produced). Here’s a comparison between the three states to illustrate what I mean.

The difference between slightly produced and produced is small and subjective. Even more subjective is that sometimes, a specimen will have one side of the 4th tergite produced and the other side not! Eagle-eyed readers will notice that is the case in the photo used for the not produced category: the left side is produced, while the right side isn’t. Other specimens in the not produced category didn’t have this mutation, but those specimens didn’t show the character quite as nicely. At any rate, combining the slightly produced and produced categories gives this map:

Distribution of Bothropolys multidentatus specimens from my collection. Dots are color coded by the character state of the 4th tergite: not produced and produced.

There doesn’t appear to be a geographical cline in the production of the 4th tergite; specimens with both produced and unproduced corners occur throughout the distribution of my specimens. I didn’t find any differences by sex or life stage either, though most of my specimens were females. So, no big surprises from this quick look, but it’s useful to know that the 4th tergite is sometimes produced in B. multidentatus. That should also make it easier to identify the species from photographs.

My sample size and area was limited considering the large distribution of the species, so it would be interesting to examine specimens from the rest of the range. The species could also use a survey of the genetic variation among populations as well, given that it’s so widespread. It has the distinct advantage of being easy to collect: if you focus on downed wood, you’re likely to find specimens pretty quickly.

References:

  • Chamberlin RV (1925) The Ethopolidae of America north of Mexico. Bulletin of the Museum of Comparative Zoology at Harvard College 57: 385–437. http://www.biodiversitylibrary.org/page/2810159

  • Crabill RE (1953) A new Zygethopolys from Kentucky and a key to the members of the genus. (Chilopoda: Lithobiomorpha: Lithobiidae: Ethopolyinae). The Canadian Entomologist 85: 119–120. https://doi.org/10.4039/Ent85119-3

A mysterious centipede parasite

I’ve been working through the backlog of unidentified centipedes in the Virginia Natural History Museum collection over the past year, and I found something very strange and cool yesterday: an internal parasite in a centipede collected in Virginia!

Dorsolateral view of the centipede (Nadabius pullus), with internal parasite visible through exoskeleton between segments 8-12.

The centipede in question was collected on May 22, 2005 by Richard Hoffman in Appomattox County, Virginia and is the common lithobiid Nadabius pullus. Under the scope, it was readily apparently there was something strange about this specimen: it was noticeably bulging in the middle of the body and was very tubular in that region, rather than flattened. So, I started dissecting the mass from the body.

The parasite peeks out of the body cavity (is the black structure an eye? I’ve no idea). Above the centipede is a portion of the parasite already dissected out.

Parasite fully dissected out of the centipede’s body.

After dissecting it out of the centipede, I was greeted with some type of parasite (parasitoid). I damaged it during the dissection, so it’s in two pieces now. The above photo is what I think is the dorsal view.

Ventral view of parasite

This thing is irritatingly non-descript. It’s about 5 mm long and 1.5 mm wide. It’s somewhat leathery, with various wrinkles and furrows. It’s mostly smooth, but does have various small pits and weak striae along the body. The main dorsal structure is a circular dark structure (see bottom of first full body image). Ventrally it has two large, cylindrical black structures that are smooth and widely separated. At the opposite end, it has a pair of dark structures which were sticking out of one of the posterior spiracles of the centipedes, but these were damaged during the dissection.

Close up of the paired black cylindrical structures, ventral side of parasite.

Dorsal view, showing structures at tip of body. I’ve tried to clean off the gunk around the structures, but gave up to avoid damaging them.

I’ve asked around and sent photos to my colleagues to see if anyone recognizes this thing, but no dice so far (from entomologists at least). Additionally, the search is hobbled by the lack of data we have on centipede parasites. We only know of a few arthropod parasites of lithobiomorph centipedes, which are limited to the Diptera and Hymenoptera (see Lewis 1981 for a summary).

Tachinid flies are the most well-documented lithobiid parasites, with two species known to parasitize stone centipedes: Eloceria delecta (Meigen, 1824) and Loewia foeda (Meigen, 1824). Both of these species are Palearctic and specialize on the European native centipede Lithobius forficatus (Linnaeus, 1758). Thompson (1939) conducted a study of 300 L. forficatus specimens from southern England and Paris, France, and found a parasitism rate of 7.5%. He reported two species of Tachinids, an unknown one and L. foeda.

In North America, Lithobius forficatus is a common introduced species in urban environments and disturbed woodlands. Crossing an ocean hasn’t removed the threat of parasitism, however, as Wood & Wheeler (1972) documented the presence of L. foeda in North America for the first time. They collected two specimens of the fly from Long Island and Ithaca, New York. Recently, Haraldseide & Tschorsnig (2014) described additional facets of the biology of this species and included descriptive notes of the puparium for the first time.

Less is known about the Hymenopteran parasites of stone centipedes. The only documented example I could find was Phaneroserphus calcar, a European Proctotrupid wasp. Newman (1867) colorfully described 14 larval P. calcar pupating and emerging from a Lithobius forficatus specimen brought to him by a friend. Apparently this is a common European species and it also parasitizes Staphylinids (Townes & Towes, 1981).

I don’t think it’s a Proctotrupid, based on the presence of only one parasite inside this centipede. My best guess is one of the Dipteran parasites, but that’s still a gamble. At any rate, this is the first documented case of parasitism in a native North American lithobiid that I’m aware of, which is pretty neat. I only live a few hours away from where this specimen was collected, so I’m going to take a trip or two out there in the future to see if I can find anymore. Maybe a fresh specimen will look more recognizable. Stay tuned for updates, and check your centipedes for weird roommates.

References:

Haraldseide, H. & Tschorsnig, H.P. 2014. On the biology of Loewia foeda (Meigen) (Diptera: Tachinidae). The Tachinid Times, 27: 15-19. https://www.uoguelph.ca/nadsfly/Tach/WorldTachs/TTimes/Tach27.html

Lewis, John G.E. 1981. The biology of centipedes. - Cambridge University Press, Cambridge: 476 pp.

Newman, E. 1867. A Proctotrupes parasitic on a myriapod. The Entomologist, 46: 342-344. https://www.biodiversitylibrary.org/page/11931840

Thompson, W.R. 1939. Biological control and the theories of the interactions of populations. Parasitology, 31: 299-388.

Townes, H. & Townes, M., 1981. A revision of the Serphidae (Hymenoptera). Memoirs of the American Entomological Institute. 32: 1-541.