Accepted by A. Minelli: 12 Jun. 2009; published: 7 Jul 2009
ISSN 1175-5326 (print edition)
1175-5334 (online edition)
Copyright © 2009 · Magnolia Press
Zootaxa 2151: 47–65 (2009)
Cave millipeds of the United States. VIII. New genera and species of
polydesmidan millipeds from caves in the southwestern United States
(Diplopoda, Polydesmida, Macrosternodesmidae)
WILLIAM A. SHEAR
, STEVEN J. TAYLOR
, J. JUDSON WYNNE
& JEAN K. KREJCA
Department of Biology, Hampden-Sydney College, Hampden-Sydney, Virginia 23943 USA. E-mail: firstname.lastname@example.org
Illinois Natural History Survey, University of Illinois, 1816 S. Oak St., Champaign, Illinois 61820 USA. E-mail: email@example.com
Merriam-Powell Center for Environmental Research, Colorado Plateau Museum of Arthropod Biodiversity, and Department of
Biological Sciences, Northern Arizona University, Flagstaff, Arizona 86011 USA. E-mail: firstname.lastname@example.org
Zara Environmental LLC, 118 Goforth Rd., Buda, TX 78610 USA. E-mail: email@example.com
Four new species of presumed troglobitic polydesmidan millipeds in two new genera are described from caves in the
states of Arizona, Nevada and California. Pratherodesmus, n. gen., is comprised of the type species, P. voylesi , n. sp., P.
ecclesia, n. sp., and P. despaini, n. sp. The genus is found in Arizona and California. Nevadesmus ophimontis, n. gen., n.
sp., is from White Pine Co., Nevada; the new genus also includes N. hubbsi (Chamberlin) 1943, new combination. All
four species were collected in or near United States National Parks, Bureau of Land Management lands, and in a private
preserve. All new taxa are authored by W. A. Shear only.
Cave habitats in the western United States are of exceptional interest, because even more than the caves of
extensive karst regions of the eastern part of the country, they represent “islands” which preserve relictual
fauna, dating back to the most recent glacial advance and retreat, or even older. While caves in the east are
likely to be surrounded by forested habitats in which the litter and soil are amenable to the continued existence
of source populations for troglobitic and troglophilic arthropods, western caves are most often located in
desert or semi-desert regions, or at high altitudes. In the former case, the surrounding environments are
inimical to soil-dwelling arthropods, and in the latter, conditions approximating those that obtained at lower
altitudes during glacial maxima or the early stages of glacial retreat still exist.
Millipeds, animals generally adapted to cool, moist conditions, are rare in deserts (there are some
exceptions) and above timberline (Golovatch 2009). During the so-called pluvial periods at glacial maxima in
North America, however, grasslands, savannah, and even forest occurred where today desert and semidesert
are seen. Conversely, during interglacials the climate may have been warmer and drier than at present, so that
the severe climates at high altitudes would have been considerably ameliorated, probably with less winter
snow and longer, warmer summers than exist today (Webb & Bartlein 1992). As these conditions changed
with the advance or retreat of the continental ice sheets, caves in both climatic regimes could provide refuges
for millipeds. During the dry, warm interglacials (such as our present time) millipeds that had colonized the
wetter, cooler environments at low altitudes could find themselves isolated in caves now surrounded by dry
grasslands, woodlands, or desert (Peck 1973, 1981, 1982). Conversely, as glaciers advanced, creating harsh
climates at high altitudes, the generally milder conditions in caves could provide a haven for millipeds that
SHEAR ET AL.48 · Zootaxa 2151 © 2009 Magnolia Press
had adapted to warmer temperatures and milder winters during an interglacial. In this model, both glacial
retreat and advance could be responsible for the isolation in caves of millipeds and other soil-dwelling
Despite this inherent interest, relatively little has been published on cave animals in the western part of the
country (apart from Texas, which has a rich literature) compared to the east (summaries in Peck 1973, 1982;
see Wynne et al. 2007, and references therein; Shear 2006, 2007, Shear & Shelley 2007, 2008). Western caves
may be more difficult to access—situated in remote desert country or high in the mountains. Caves at high
altitudes may be accessible to collectors only for relatively brief times of the year, due to snowfall and low
temperatures. Many western caves are located on public lands—national parks or monuments, national forests
or Bureau of Land Management (BLM) lands. It is important to document finds of animals in these caves for
This paper describes two new genera and four new species of polydesmidan millipeds recently collected
in caves in the states of Nevada, Arizona and California. Because the locations where three of these new taxa
were collected are on U.S. public lands and a private preserve (Sequoia Kings Canyon National Park, Great
Basin National Park, BLM -Arizona Strip Field Office lands, and Cathedral Cave Preserve), they are of
significance for the conservation of their habitats. These animals likely retain a limited ability to disperse to
other caves, microclimatic conditions of caves are critical to their survival, and their habitats are sensitive to
disturbances from local and regional threats, so all four species are candidates for potential designation as
species of concern.
Type specimens of the new species described below are deposited in the Field Museum of Natural History
(FMNH), Chicago; paratype specimens of Pratherodesmus voylesi and P. ecclesia are curated at the Colorado
Plateau Museum of Arthropod Biodiversity (CPMAB), Flagstaff, Arizona and additional paratypes of
Nevadesmus ophimontis n. sp. are deposited in the collection of the Illinois Natural History Survey (INHS).
Order Polydesmida Pocock, 1887
Suborder Polydesmidea Pocock, 1887
Infraorder Polydesmoides Pocock, 1887
Superfamily Trichopolydesmoidea Verhoeff, 1910
Family Macrosternodesmidae Brölemann, 1916
Shear and Shelley (2007, 2008) diagnosed and discussed the Family Macrosternodesmidae, but the line of
demarcation between the macrosternodesmids and nearctodesmids remains rather unclear, particularly in light
of the fact that the genera described below have smooth metaterga, rather than having them divided into
subquadrate elevated areas, as in Tidesmus Chamberlin, 1943. Smooth metaterga, with few or no setae, are
characteristic of nearctodesmids. The gonopods of at least two of the species of Pratherodesmus, new genus,
have the process arising from the mediodistal side of the prefemur, called process B by Shear and Shelley
(2007), reduced or absent, likewise the distal zone. The gonopods of macrosternodesmids and nearctodesmids
are very similar, except that nearctodesmids have been maintained as having both process A and B, and
macrosternodesmids only B, but in the present two genera, it would seem process A is indeed present, and as
mentioned above, process B is absent in two species of Pratherodesmus. This blurring of the distinction
between the families suggests to us that eventually they will need to be combined under
Macrosternodesmidae, the older name, as more taxa are discovered and described.
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A recent paper by Djursvoll (2008) on the Iberian polydesmid genus Schizomeritus Verhoeff, 1931,
makes possible, through its clear SEM photographs and labeling, some reconciliation between the gonopod
nomenclature originated by Shelley (1994) and used in previous papers on macrosternodesmids (Shear &
Shelley 2007, 2008), and that adapted by Djursvoll et al. (2001 ) and Golovatch and Wytwer (2007). It
appears that our process B is the exomere of Djursvoll (ex in our figures), process A corresponds to the
endomerite (en in our figures), and our distal zone is Djursvoll’s tibiotarsus (tt in our figures). We previously
have not used the term femorite, which Djursvoll (2008) uses for the part of the acropodite traversed by the
seminal groove, but agree this term is a useful one, and apply it here. Our solenomere (s in our figures) and
that of Djursvoll (2008) are the same. In the descriptions that follow we have decided to use the Djursvoll
terminology, because it has the advantage of using terms that have already been in the literature for a
considerable time, but now with clear definitions; application of this terminology allows for discussion of
possible homologies between genera and families of the superfamilies Polydesmoidea and
Trichopolydesmoidea. Earlier concerns expressed by Shear and Shelley (2007) now seem to us to be resolved.
Interestingly, the gonopod structure of species of Schizomeritus is very similar to that of the new genus
Pratherodesmus, while the nonsexual characters are quite different. Species of Pratherodesmus have only
small paranota, strongly produced posteriolaterally, and with regularly arranged rows of clavate setae on an
otherwise featureless surface. Species of Schizomeritus are more typical of Polydesmidae, with large, squared,
anteriorly elevated paranota not strongly produced posteriolaterally. The metazonite surfaces are deeply
sculpted into bulging polygonal areas, and the setae are inconspicuous. On poriferous segments
Pratherodesmus species have the ozopores nearly at the posterior corner and the edges of the paranota are not
swollen, while in Schizomeritus the pores are more anterior and the edges strongly rebordered. Despite the
similarity of the gonopods in general appearance, there are differences. The course of the seminal canal
crosses over to the lateral side in Schizomeritus and ends in a vesicle and pulvillus. In Pratherodesmus, the
canal stays mesal, does not end in a vesicle, and there is no pulvillus (although a dense group of cuticular
fimbriae probably not homologous to a true pulvillus occurs). At this point it is not at all clear what the
significance of these similarities and differences may be.
As previously remarked (Shelley and Shear 2007), the millipeds of the superfamilies
Trichopolydesmoidea and Polydesmoidea need close re-examination. Several families in both superfamilies,
including Polydesmidae, remain poorly diagnosed and have been used as “wastebaskets” for enigmatic genera
and species. We believe that at least two undiagnosed polydesmoid families are to be found in the very
diverse, but largely undocumented, North American fauna, and have on hand many new generic and species-
level taxa, primarily from the Pacific northwest. In the course of describing these taxa it may be possible to
begin to make some sense of the two superfamilies, but without much more knowledge of small tropical
polydesmidans, the composition and fate of the Trichopolydesmoidea remains uncertain.
Presently four genera from southwestern North America, including the two new ones below, have been
assigned to the family Macrosternodesmidae. They may be separated by the following key:
Key to macrosternodesmid genera in southwestern North America
1a. Adults 6 mm or less in length......................................................................................................................................2.
1b. Adults 8 mm or more in length....................................................................................................................................3.
2a. Paranota obselete or lacking; metazonites with scattered, thin, acute setae; male gonopod with solenomere longer
than other gonopod processes.................................................................................... Sequoiadesmus Shear & Shelley.
2b. Paranota more obvious, toothed; metazonites with three rows of clavate setae arising from low pustules; male gono-
pod with soleomere short...............................................................................................................Nevadesmus, n. gen.
3a. Metazonital setae arising from obvious pustules, 12–16 in anterior row, posteriorlateral corners of paranota not pro-
duced; male gonopod with exomere large, U-shaped; large, folded tibotarsus................. Tidesmus Chamberlin 1943.
3b. Metazonital setae not arising from pustules, no more than 8 in anterior row, posteriolateral corners of metazonites
produced as acute processes beyond ozopore; gonopods with exomere small or absent, tibiotarsus small or lacking..
.................................................................................................................................................Pratherodesmus, n. gen.
SHEAR ET AL.50 · Zootaxa 2151 © 2009 Magnolia Press
Pratherodesmus Shear, new genus
Type species: Pratherodesmus voylesi Shear, new species.
Etymology: Named for the late John W. Prather, former lead scientist and spatial ecologist for the ForestERA
Project, and professor at Northern Arizona University, Flagstaff, Arizona.
Diagnosis: Small (<10.0 mm length) polydesmidan millipeds with 20 trunk segments, lacking pigment;
metatergites smooth, with three transverse rows of short, acute or clavate setae, rows sometimes strongly
recurved. Collum ovoid, narrower than head and first leg-bearing segment. Paranota low, margins toothed,
posteriolateral angles sharply drawn out. Pygidium blunt, nearly hemispherical when viewed dorsally,
sparsely setose, with usual four spinnerets (Shear 2008) arranged in a square and set in individual depressions;
pygidial process blunt, decurved. Males with pregonopodal legs unmodified or encrassate. Gonopods with
coxae globular, fixed, entirely filling gonostome, tightly appressed or fused in midline; prefemora sparsely
setose, strongly transverse, articulating with coxae by process fitting into coxal notch. Exomere small or
absent, endomerite large, bulky, dominating gonopod. Acropodite short, solenomere nearly sessile, opening of
seminal canal widened, subtended by cuticular teeth and two processes, one proximal and one distal (distal
Pratherodesmus differs from Tidesmus Chamberlin 1943 (Shear & Shelley 2007) in having a much
smaller tibiotarsus of the male gonopod and in its smooth or nearly smooth metazonites. Sequoiadesmus Shear
& Shelley 2008 occurs near the type locality of P. despaini, though at a much higher elevation and in a
different cave group (see map in Shear & Shelley 2008); the gonopod of the single known species has an
extremely long solenomere and lacks an exomere. Sequoiadesmus krejcae Shear & Shelley 2008 is also much
smaller (5.8 mm long vs. 9.0 mm for P. despaini) and has densely scattered, acute metzonital setae rather than
short, clavate ones ranged in rows.
Distribution: Known from caves in northwestern Arizona and the Sierra Nevada of California.
Notes: The genus presently consists of three species of small, white, presumably troglobitic, millipeds
found only in caves in the states of Arizona and California. We surmise that the species are in effect cave-
limited (troglobionts), because the habitats surrounding the caves are inimical to small millipeds (Fig. 38).
The caves of the southwestern part of the United States have not been well-investigated for cave life, with the
exception of limited studies of bats; and some investigations of caves where arthropods were opportunistically
collected. Notable exceptions are cave-specific inventories of Carlsbad Caverns National Park, New Mexico
(Barr and Reddell 1967) and Karchner Caverns, Arizona (Welbourn 1999). Broader surveys were published
by Peck (1973, 1981, 1982). Given this, we expect more species of Pratherodesmus to be discovered in the
Key to Species
1a. First row of metazonital setae strongly recurved, lateralmost seta on each side widely separated from others in row;
posterior marginal row with 5 or 6 setae (Fig. 5); endomerite of gonopod with two small apical lobes set with regu
larly distributed pustules (Figs.21–24); Tulare Co., California..........................................................P. despaini, n. sp.
1b. First row of metazonital setae moderately recurved, lateralmost seta on each side not widely separated from others in
row (Figs. 4, 6); posterior marginal row with 4 setae; endomerite of gonopod lacking terminal processes (Figs. 33,
2a. In anterior or posterior view, endomerite of gonopod narrow, slightly sinuous (Fig. 17); Yavapai Co., Arizona .........
............................................................................................................................................................ P. ecclesia, n. sp.
2b. In anterior or posterior view, endomerite of gonopod broad, in posterior view with distinct lateral notch (Figs.
–13); Mohave Co., Arizona ..............................................................................................................P. voylesi, n. sp.
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FIGURES 1, 2. Heads, collums and first two segments of males, dorsal view. Fig. 1. Pratherodesmus despaini. Fig. 2. P.
Pratherodesmus voylesi Shear, new species
Figs. 2, 6, 9, 11–14, 19, 33, 36.
Types: Male holotype, five male and seven female paratypes from Millipede Cave, Mojave Co., Arizona,
collected 16 February 2003 by K. Voyles, J. Jasper, M. Porter and K. Dittmar de la Cruz, deposited in FMNH.
Four male and sixteen female paratypes from October Gyp Cave, collected 27 March 2004 by same
collectors. All specimens deposited in FMNH and CPMAB.
Description: Male. Length, 7.5 mm, width 0.75 mm. Antennae long, extended backward reaching to
posterior border of sixth segment. Head 20–25% wider than collum, anterior margin of collum (Fig. 2) nearly
semicircular, with 10 marginal setae, posterior margin with six marginal setae, nearly straight; middle row
with four setae. Typical midbody segment (segment 10; Fig. 6) with metazonite as broad as long, three
prominent lateral marginal teeth, posteriolateral angles moderately extended beyond ozopores; marginal teeth
each with seta; anterior setal row recurved, with six setae, posterior row nearly straight, with four setae,
posterior marginal row with four setae; three setae arranged around ozopore. Posteriorly, setation of anterior
row increases to eight. Pygidium (Fig. 9) nearly hemispherical in dorsal view, with about 10 setae visible
dorsally, pygidial process short, with four spinnerets arranged in square, typical of polydesmideans. Legs
Pregonopodal legs slender, unmodified. Gonopods (Figs.11–14, 33) with hemispherical coxae filling
gonostome, tightly appressed, probably immovable, anteriorly excavate to receive telopodites. Prefemora
strongly transverse, sparsely setose, distally narrowed to short stem connecting to acropodite. Exomere
absent. Endomerite large, blunt, complex, with prominent lateral notch. Course of seminal canal along
femorite straight. Solenomere with fine cuticular scales near seminal opening, subtended by short process;
apical zone small, flattened, curved, ridged.
Female. Length, 8.2 mm, width, 0.9 mm. Nonsexual characters as in male, but segmental setae longer,
more acute, body less slender. Cyphopods as single fused organ (Fig. 19), evidently permanently extruded,
with two anterior pores.
SHEAR ET AL.52 · Zootaxa 2151 © 2009 Magnolia Press
Distribution and habitat: Known only from the two caves named above, located in the northeastern
corner of Mohave Co., Arizona, on BLM-Arizona Strip Field Office lands. The exact location of the caves is
not given because of conservation concerns.
FIGURES 3–6. Segments 10, dorsal view. Fig. 3. Nevadesmus ophimontis. Fig. 2. Pratherodesmus ecclesia. Fig. 3. P.
despaini. Fig. 6. P. voylesi.
Millipede Cave (elevation 1371 m [4497 ft.]) is a limestone and gypsum cave with 274 m (900 ft.) of
mapped passage. This cave has a vertical entrance approximately 2 by 4m. Average passage height is < 2m.
The cave floor is characterized by areas of medium to small breakdown, with unconsolidated silts and sands,
stream gravels and exposed bedrock. A single trunk passage is characterized by medium-sized rooms; cave
roof height varies from walkable to crawling passage. At the back of the cave, there are three large pools
which create ~100% rH (relative humidity) at this depth, where the accessible cave ends in a sump. Floods
periodically transport significant amounts of vegetation into this cave, including various types of weeds e.g.,
Russian thistle (Salsola kali), black brush (Coleogyne ramosissima), rabbit brush (Chrysothamnus molestus)
and creosote (Larrea tridentata). Logs and branches have also been washed in; however, the source of this
larger material is unknown. Detritus often covers portions of the cave floor and forms mounds towards the
back of the cave. P. voylesi is commonly observed on this material; we suggest this animal is either feeding on
detritus, or on fungus or bacteria growing on detritus. Individuals of P. voylesi occur from approximately 60 m
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(200 ft.) within the cave to ~121 m (400 ft.) up to the sump. Unflooded passage does occur beyond the sump;
however, it is unknown whether this species occurs in that part of the cave. Other arthropods detected in the
cave include spiders (spp. not known) and pillbugs (Porcellio laevis, det. Stefano Taiti). Surface vegetation is
inter-mountain basins mat saltbush shrubland (SWReGAP land cover type; Lowery et al. 2006).
FIGURES 7–10. Figs. 7–9. Pygidia, dorsal view. Fig. 7. Nevadesmus ophimontis. Fig. 8. Pratherodesmus despaini. Fig.
9. P. voylesi. Fig. 10. Metazonital setae anterior to ozopore of segment 10, Nevadesmus ophimontis.
October Gyp Cave (elevation 1392 m [4566 ft.]) is a linear phreatic tube with two levels, and consists of
269 m (882 ft.) of passage. This cave has a vertical entrance ~ 2m in diameter. The upper extent is
characterized by dry limestone passage, while the lower passageway is moist to wet gypsum. The lower
passage contains 30m (100 ft.) of walkable passage tapering to a series of belly crawls to hands-and-knees
crawls. The upper level is exposed hardpan with no millipeds, while the lower level is silt to silty clay; this is
where the millipeds occur. This cave receives little to no flood detritus. Millipeds are observed from ~134 m
(440 ft.) onward to the terminus of the cave. Other arthropods observed in the cave include crickets
(Ceuthophilus sp.) and beetles (Family Tenebrionidae). Surface vegetation is inter-mountain basins mixed salt
desert scrub (SWReGAP land cover type; Lowery et al. 2006).
There is one cave within two miles of Millipede and October Gyp caves that contains polydesmid
milllipeds. While unconfirmed, this likely represents an additional locality of this new species. In the course
SHEAR ET AL.54 · Zootaxa 2151 © 2009 Magnolia Press
of the survey that produced this species, about 300 individual caves were visited, and only the three
mentioned here were found to support milliped populations. This does not prove the millipeds are absent from
these caves, but they were not found during the time available for exploration.
Etymology: Named for Kyle Voyles, Arizona BLM Cave Coordinator, who organized and led the
collecting effort that resulted in the type specimens.
FIGURES 11–14. Gonopods of Pratherodesmus voylesi. Fig. 11. Gonopods in gonostome, ventral view. Fig. 12.
Posterior view. Fig. 13. Lateral view. Fig. 14. Solenomere, distal zone, and subtending process, posterior view.
Pratherodesmus ecclesia Shear, new species
Figs. 4, 15–18, 34, 36.
Types: Male holotype, two male, and four female paratypes from Cathedral Cave (part of Cathedral Cave
Preserve), 10 miles south-southwest of Ash Fork, Yavapai Co., Arizona, collected 11 January 2007 by J. J.
Wynne, deposited in FMNH; one male and one female paratype in CPMAB.
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Description: Male. Length, 8.0 mm, width 0.75 mm. Nonsexual characters as described for P. voylesi, but
metazonital setae are significantly longer and more acute (Fig 4). Pregonopodal legs slightly more crassate
than postgonopodal legs. Gonopods (Figs. 15–18, 34, 35) large, gonostome walls bulging, prozonite of
seventh segment notably swollen when seen dorsally. Gonocoxae hemispherical, filling gonostome, tightely
appressed in midline, excavated to receive telopodites. Prefemora as in P. voylesi, but with lateral articulation
less narrow, subtriangular; not obviously distally narrowed. Exomere absent, endomerite long, sinuous,
appearing narrow in either anterior or posterior view, in lateral view, acute-triangular, with small basal tooth.
Solenomere similar to P. voylesi, tibiotarsus reduced to absent, subtending process, large, lamellate.
Female. Length, 8.0 mm, width 0.83 mm. Nonsexual characters as male, cyphopods as described for P.
FIGURES 15–18. Gonopods of Pratherodesmus ecclesia. Fig. 15. Gonopods in gonostome, ventral view. Fig. 16.
Lateral view. Fig. 17. Anterior view. Fig. 18. Posterior view.
SHEAR ET AL.56 · Zootaxa 2151 © 2009 Magnolia Press
FIGS. 19, 20. Fig. 19. Cyphopods of Pratherodesmus voylesi, ventral view. Fig. 20. Solenomere, distal zone, and
subtending process of gonopod of P. despaini, posterior view.
Distribution and habitat: Cathedral Cave (elevation 1621 m [5317 ft.]) is a limestone cave characterized
by a large borehole passage, with two narrow side passageways. This cave has not been mapped. The vertical
entrance is 1 x 3 m. During the winter months, Cathedral Cave is wet, with standing pools and high humidity,
but prior to the summer monsoon it is quite dry. P. ecclesia is often observed in the deepest parts of this cave
in association with tree branches, likely transported into the cave by human visitors (perhaps for use as
torches); they also have been observed in association with candle wax. We suggest this animal is likely
feeding directly on the wood or perhaps on the fungi and bacteria growing on the wood; there are often
accumulations of dark brown milliped scat where these animals are observed. This species is commonly
observed in association with two collembolans (Sinella sp., Drepanura sp.). Other arthropods occurring in this
cave include two spider species (Cicurina sp., Metellina curtisi, det. Pierre Paquin), three beetles (Bembidion
rupicola, det. Rolf Aalbu; Nicrophorus sp.; Rhadine n. sp. Thomas Barr), crickets (Ceuthophilus utahensis,
det. Theodore Cohn) and ants (Camponotus ocreatus, det. Robert Johnson). Surface vegetation is Colorado
Plateau pinyon-juniper woodland (SWReGAP land cover type; Lowery et al. 2006).
Etymology: The name ecclesia is a noun in apposition, Latin for “church.”
Pratherodesmus despaini Shear, new species
Figs. 1, 5, 8, 20–24, 32, 35, 36, 39
Types: Male holotype, two male and one female paratype from Kaweah Cave, Tulare Co., California,
collected 28 April 2004 by J. Krejca, P. Sprouse, S. Fryer, D. Ubick, P. Paquin and W. Savary; one male and
two female paratypes from the same locality, collected 10 August 2006 by J. Krejca and J. Despain, deposited
Description: Male. Length 9.0 mm, width 0.6 mm (Fig. 39). Head about 50% wider than collum (Fig. 1).
Antennae long, extending back to posterior border of fifth segment. Collum with anterior margin arcuate, with
12 marginal setae, posterior margin straight to slightly sinuate, with eight setae, middle row with six setae;
posteriolateral angles slightly produced (Fig. 1). Typical midbody segment (Fig. 5, segment 10) with three
lateral marginal teeth, each subtended by a seta; anterior row of six setae very strongly procurved, lateralmost
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seta of row widely separated, not appearing to be part of row; middle row of six setae slightly recurved;
posterior marginal row with six setae. Posteriolateral metazonital corners strongly produced beyond ozopores,
ozopores subtended by usual three setae. Pygidium as described for P. voylesi, but with 12 long setae (Fig. 8).
Pregonopodal legs markedly encrassate (Fig. 1). Gonopods (Figs. 20–24, 32, 35) with gonocoxae and
gonostome as described for P. voylesi. Prefemora strongly transverse, articulating process narrow, pointed;
prefemoral stem not marked. Exomere present, small, slightly curved, acute. Endomerite large, trullate,
apically with two small lobes set with prominent, regular, small warts. Solenomere longer than in other
species, with fewer cuticular scales; subtending process short, acute. tibiotarsus not flattened, as acute process
similar in length to solenomere.
Female: Length, 9.0 mm, width 0.70 mm. Nonsexual characters similar to male. Cyphopods not observed,
not extruded in available female specimens.
FIGURES 21–24. Gonopods of Pratherodesmus despaini. Fig. 21. Gonopods in gonostome, ventral view. Fig. 22.
Lateral view. Fig. 23. Anterior view. Fig. 24. Posterior view.
SHEAR ET AL.58 · Zootaxa 2151 © 2009 Magnolia Press
Distribution and habitat: Known only from the type locality. Kaweah Cave is located close to the
western boundary of Sequoia Kings Canyon National Park, and is unique among the caves in the park because
it occurs at low elevation and has greater biodiversity than is found among the higher altitude caves (for
locations of caves in the park, see the map in Shear & Shelley 2008). Four troglobitic species, including P.
despaini, and two other endemic troglophiles are to be found there. The other troglobites are an unidentified
trichoniscid isopod, a cambalid millipede, and a pseudoscorpion, probably a species of Tuberochernes. The
endemic troglophiles include a harvestmen in the genus Calicina and a spider in the genus Usofila. These may
all represent new unnamed species. The cave’s accessible portion is a sinuous crawl less than 100 m long,
ending in a large room where the animals were collected. One male was collected from a root, and the other
specimens were associated with unidentified guano (probably rodent or bat) on the floor. The temperature was
not measured on the day of collections, but during an earlier visit to the same room of the cave the temperature
was recorded at 11 degrees Celsius. The surrounding vegetation is California lower montane blue oak-foothill
pine woodland and savanna (SWReGAP land cover type; Lowery et al. 2006).
Etymology: Named for Joel Despain, Cave Management Specialist at Sequoia and Kings Canyon
National Parks, who organized and performed much of the sampling that resulted in the discovery of many
new cave species.
Notes: This species is more distant from the closely related P. ecclesia and P. voylesi. While the gonopods
are built along the same basic plan, the presence of a small exomere in P. despaini suggests, in comparison
with Nevadesmus and Tidesmus, that it would occupy a more basal phylogenetic position within
Nevadesmus Shear, new genus
Type species: Nevadesmus ophimontis Shear, new species.
Etymology: Named for the state of Nevada, with the combining stem –desmus, traditionally used for the
names of polydesmidan millipeds.
Diagnosis: Very small (<5.0 mm length) polydesmidan millipeds with 20 trunk segments, lacking
pigment; metatergites (Fig. 3) with three transverse rows of short, brushlike or clavate setae (Fig. 10), rows
straight, setae on low pustules. Collum ovoid, narrower than head and slightly narrower than first leg-bearing
segment. Paranota low, margins not strongly toothed, posteriolateral angles right-angled to acute, but not
produced into processes beyond ozopores (Figs. 3, 10). Pygidium blunt, nearly hemispherical when viewed
dorsally, sparsely setose, with usual four spinnerets (Shear 2008) arranged in a square and set in individual
depressions; pygidial process very short, continuing line of pygidium. Males with pregonopodal legs
unmodified. Gonopods with coxae globular, fixed, entirely filling gonostome, tightly appressed or fused in
midline; prefemora sparsely setose, strongly transverse, articulating with coxae by process fitting into coxal
notch. Exomere long, sinuous, arising from distomesal margin of prefemur, endomerite large, distally with
three sinuous processes, exceeding other parts of gonopod. Acropodite bulky, solenomere incurved, nearly
sessile, opening of seminal canal widened, proximal subtending process widely separated from solenomere,
thin, curved, acute; no obvious tibiotarsus.
Nevadesmus differs from all previously known southwestern macrosternodesmids in its small size. It may
be distinguished from Sequoiadesmus by its much shorter solenomere.
Distribution: Known from caves in White Pine and Lincoln counties, Nevada.
Notes: This genus is closer in its gonopod to Tidesmus than to Pratherodesmus, particularly in the
proportions of prefemoral processes. However, the gonopods most closely resemble those of the much larger
Harpogonopus Loomis 1960, generally accepted as a nearctodesmid (Shelley 1993, 1994; Shelley & Shear
2006). Though having the general somatic appearance of a macrosternodesmid, the gonopods of species of
this genus are quite nearctodesmid-like, with both endomerite and exomere well-developed, but without an
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It now seems likely, based on size, geographical proximity, and nonsexual characters that “Tidesmus”
hubbsi Chamberlin 1943 (see Shear & Shelley 2007) is a member of this genus, and it is included here, though
males still have not been collected. A female of hubbsi was illustrated by Shear & Shelley 2007, and the
history and provenance of the species was discussed therein.
Nevadesmus ophimontis Shear, new species
Figs. 3, 7, 10, 25–31, 36, 37.
Types: Male holotype, male and two female paratypes from Model Cave, White Pine Co., Nevada, collected
23 May 2003 by S. J. Taylor, J. K. Krejca, K. Patel, M. Porter, K. Dittmar de la Cruz, deposited in FMNH. The
following specimens (deposited at FMNH and INHS) are also paratypes: NEVADA: White Pine Co., same
data as holotype, but collected 22 May 2006 by J. K. Krejca, M. E. Slay, G. Baker, 2 females; Snake Creek
Cave, 29 May 2003, S. J. Taylor, J. K. Krejca, K. Patel, L. D. Seale, A. Hamilton, S. Johnson, 5 females; 21
May 2006, S. J. Taylor, J. Krejca, M. E. Slay, 3 males, juveniles; Lehman Caves, 26 May 2006, S. J. Taylor, J.
K. Krejca, M. E. Slay, G. Baker, 3 males, 3 females; Little Muddy Cave, 23 May 2003. S. J. Taylor, J. K.
Krejca, M. Porter, K. Dittmar de la Cruz, a juvenile male presumed this species; Wheeler’s Deep Cave, 26
May 2003, S. J. Taylor, J. K. Krejca, M. Porter, K. Dittmar de la Cruz, 2 males, 2 females.
Description: Male: Length, 4.5 mm, width 0.45 mm (Fig. 37). Antennae short, clavate; head about 40%
wider than collum. Collum with arcuate anterior margin bearing 12 short, clavate setae, posterior margin
slightly procurved, with 6(5) marginal setae. Second segment slightly wider than collum. Typical midbody
segment (segment 10; Fig. 3) with inconspicuous paranota, two paranotal teeth on each side (possible third
tooth is directly opposite ozopore), posteriolateral corner right-angled to slightly obtuse, not drawn out
beyond ozopore into sharp process. Metazonital setae on modest tubercles, short, clavate; anterior row of six,
with lateralmost setae on each side posteriorly displaced; middle row of four setae, slightly procurved;
marginal posterior row of four (six on some segments) setae; usual triad of setae subtends ozopore. Pygidium
(Fig. 7) short, rounded, with eight setae, pygidial process very short, blunt, bearing usual four spinnerets.
Pregonopodal legs encrassate. Gonopods (Figs. 25–31) with hemispherical coxae tightly appressed in
midline, excavated to receive retracted telopodites. Prefemora densely setose, strongly transverse, articulating
process acute; exomere arising from anteriomesal margin of prefemur, long, thin, sinuous. Endomerite from
distolateral edge of acropodite, basally broad, incurved, ending in three unequal processes, middle one
longest. Subtending process of solenomere is set low and laterally on acropodite, seminal pore with relatively
few cuticular teeth.
Female: Length, 4.8 mm, width 0.5 mm. Nonsexual characters as in male; cyphopods not studied.
Distribution and habitat: As shown by the map (Fig. 36), the caves occupied by Nevadesmus
ophimontis are relatively closely clustered, and all are located in Great Basin National Park. Model Cave is a
large cave (length 599.9 meters [1968.1 feet]) at an elevation of 2080 meters (6824 feet). The fauna of Model
Cave is dominated numerically by Collembola, followed by mites, mayflies, and flies. Globular springtails
(Arrhopalites spp.) including an undescribed species, were particularly abundant. In addition to N.
ophimontis, three troglobitic or troglophilic species are present, including the sclerobunine harvestman
Cyptobunus ungulatus ungulatus Briggs, the conotylid milliped Idagona lehmanensis Shear (see Shear 2006),
and the pseudoscorpion Microcreagris grandis Muchmore. Surrounding vegetation (Fig. 38) for all the caves
listed as supporting populations of N. ophimontis is Great Basin pinyon-juniper woodland (SWReGAP land
cover type; Lowery et al. 2006).
Lehman Caves, at an elevation of 2096 meters (6877 feet), is the largest (length ~3352.8 meters [~11,000
feet]) cave in the Great Basin National Park. The fauna includes Collembola, Diptera, and mites, in addition to
M. grandis and N. ophimontis. Notably absent from the Lehman Caves, in spite of intensive collecting, were
C. ungulatus ungulatus and Idagona lehmanensis. Eight taxa of Collembola have been collected in this cave.
SHEAR ET AL.60 · Zootaxa 2151 © 2009 Magnolia Press
Little Muddy Cave is a large cave (length 309 meters [1010.5 feet]) at an elevation of 2045 meters (6709
feet). Microcreagris grandis occupies this cave; it is a possible predator on N. ophimontis.
Snake Creek Cave is a large (length 51.3 meters [1682 feet]) cave at an elevation of 2030 meters (6660
feet), and located on a fairly barren, south facing slope (Fig. 38). Springtails and the psocopteran Speleketor
sp. were dominant in collections.
FIGURES 25–29. Gonopods of Nevadesmus ophimontis. Fig. 25. Posterior view. Fig. 26. Anterioventral view. Fig. 27.
Lateral view. Fig. 28. Gonopods in gonostome, ventral view. Fig. 29. Solenomere, posterior view.
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Wheeler’s Deep Cave is one of four interconnected caves making up the Baker Creek Cave System,
Nevada’s longest cave (length 1315 meters [4315 feet]). It is located at an elevation of 2147 meters (7044
feet) in a limestone outcrop adjacent to the riparian zone of Baker Creek. Cyptobunus ungulatus ungulatus
and N. ophimontis are fairly common in the deeper, wetter parts of this cave, where a perennial stream is
Etymology: The species name refers to the Snake Mountain Range, and to Snake Valley, major
geographical features associated with the collection localities.
FIGURES 30–35. Drawings of gonopods. Figs. 30, 31. Nevadesmus ophimontis. Fig. 30. Lateral view. Fig. 31. Mesal
view. Fig. 32. Pratherodesmus despaini, posteriolateral view. Fig. 33. P. voylesi, mesal view. Figs. 34,35. P. ecclesia. Fig.
34. Posterior view. Fig. 35. P. despaini, mesal view.
Nevadesmus hubbsi (Chamberlin), new combination
Tidesmus hubbsi Chamberlin, 1943:36, fig. 4. Chamberlin & Hoffman, 1958:74 (list). Shear & Shelley, 2007:63, figs.
SHEAR ET AL.62 · Zootaxa 2151 © 2009 Magnolia Press
Remarks: Shear & Shelley explained why hubbsi could not be congeneric with Tidesmus episcopus
Chamberlin, 1943, the type species of Tidesmus. Repeated collecting efforts have failed to turn up male
specimens, but the close resemblance between females of N. hubbsi and N. ophimontis makes it highly likely
that they are congeneric. The type locality of this species is Cave Valley Cave, in northern Lincoln Co.,
Nevada (see Fig. 25 in Shear & Shelley 2007).
FIGURE 36. Map of parts of the states of Arizona, California, Nevada and Utah, showing localities for milliped
collections (Map by S. Taylor).
Comments on the management status of these species
These four millipede species are sensitive to disturbance for multiple reasons. Local threats include
inappropriate cave management strategies that could directly harm the species by trampling or habitat
alteration (e.g. eutrophication, contamination). The species have extremely limited ranges in a handful of
caves, and these caves may receive visitor impacts that are high relative to the total area of the cave. In the
case of P. voylesi, some 300 caves in the vicinity were surveyed for arthropod populations, and only three of
these, in a tightly circumscribed area, supported milliped populations. In addition, groundwater extraction
could alter microclimate in some of the caves. Regional threats include global climate change, a factor that
recent authors consider likely to become the greatest threat to biodiversity in most regions of the world
(Thomas et al. 2004). For cave species, we typically lack complete distribution information, and do not have
an understanding of their responses to rising surface temperatures (Wynne et al., 2008a). Because their habitat
is discontinuous, their ability to disperse may be dramatically affected by climate change. Sala et al. (2000)
suggest climate change will have the greatest effect on biodiversity in biomes characteristic of extreme
climates. Since caves are characterized as nutrient poor and aphotic, and often exhibit low climatic variability
at depth, caves are indeed an extreme biome. Wynne et al. (2007, 2008b) have established that cave thermal
behavior is influenced by surface temperatures. However, it is uncertain to what extent cave deep zones may
be buffered from global climate change (Wynne et al. 2008a).
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FIGURES 37, 38. Fig. 37. Living example of Nevadesmus ophimontis, photographed in Snake Creek Cave; length of
animal is about 5 mm. Fig. 38. Great Basin pinyon-juniper woodland of the Snake Mountains seen from just inside the
entrance to Snake Creek Cave (photos by J. Krejca).
Given the possible effects of mismanagement and potential groundwater withdrawls, and the uncertainties
regarding the effects climate change on these species, we recommend establishing population monitoring
protocols and regular habitat parameter measurements. Given the sensitive nature of the habitats in which
SHEAR ET AL.64 · Zootaxa 2151 © 2009 Magnolia Press
these four new millipede species occur, we recommend a conservative approach to protecting populations of
species of Nevadesmus and Pratherodesmus species until more is known about their ecological requirements
and distributions. Steps have already been taken in this direction, as BLM-Arizona Strip has closed Millipede
and October Gyp Caves to visitation. Kaweah Cave and the Nevada caves with populations of N. ophimontis
are on National Park Service lands and so receive some protection; further measures are under study.
FIGURE 39. Living example of Pratherodesmus despaini, photographed in Kaweah Cave (photo by J. Krejca).
We thank all of the collectors named above, because considerable effort and some danger is involved in the
biological surveying of desert caves. We also acknowledge the cooperation and help of the staffs of Great
Basin National Park, BLM-Arizona Strip Field Office, Sequoia Kings Canyon National Park (in particular
Joel Despain, Danny Boiano and Shane Fryer), and Cathedral Cave Preserve (in particular Doug Billings and
Tom Gilleland). WS thanks his colleague and frequent collaborator Rowland Shelley (North Carolina State
Museum of Natural History) for sending the specimens described here as Pratherodesmus despaini, and for
enlightening discussions on North American polydesmoidean families. Asa Kreevich provided valuable
comments on the manuscript.
This paper is published with the support of a National Science Foundation PEET Grant (DEB05-2917) to
W. A. Shear, P. Sierwald and J. Bond, and a grant from the Professional Development Committee of
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