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PROC. ENTOMOL. SOC. WASH.
116(3), 2014, pp.
NEW SPECIES AND NEW RECORDS OF JUMPING BRISTLETAILS
FROM THE ROCKY MOUNTAINS (MICROCORYPHIA:
MEINERTELLIDAE, MACHILIDAE)
G
RANT
D. D
E
J
ONG
GEI Consultants, Inc., 4601 DTC Boulevard, Suite 900, Denver, Colorado 80237
U.S.A. urn:lsid:zoobank.org:author:E99DB80C-5A7D-4D51-AB42-F22D569D92FC
(e-mail: gdejong@geiconsultants.com)
Abstract.—The Microcoryphia are extensively distributed in the Nearctic Region,
with over 30 species in 12 genera in two families. The genus Machilinus Silvestri
1904 has previously been reported from Arizona, British Columbia, California,
Nevada, Utah, and possibly Colorado. I report and describe three new species of
Machilinus based on specimens from Colorado, New Mexico, Arizona, Idaho,
Montana, and Nevada. Machilinus matadero, new species, was collected from June
through November in metropolitan Denver, Colorado. Machilinus taoseno, new
species, was collected in April through August in northern New Mexico, central
Colorado, and central Arizona. Machilinus zingiberus, new species, was collected
in July through August in central Idaho and northern Nevada. A few biological notes
are presented for each species. The machilid Pedetontus californicus (Silvestri 1911)
is reported from Idaho for the first time.
Key Words: —description, distribution, new record, epigaeic, Nearctic
DOI: 10.4289/0013-8797.116.3.xxx
urn:lsid:zoobank.org:pub:CF51FA6B-68E3-47CC-910D-DBAA4C40CC0E
Microcoryphia (=Archaeognatha), or
jumping bristletails, is a small order of
Hexapoda (approximately 500 species
world-wide) with a Nearctic fauna of
over 30 described species in two families
and a dozen genera (Poole and Gentili
1997, Sturm and Bach 1992, Packauskas
and Shofner 2010). Distribution of the
Microcoryphia appears to be widespread
in the United States, but only reported
sporadically due to limited collecting
and few taxonomic specialists working
on the group (Wygodzinsky and Schmidt
1980, Allen 1995). Machilinus Silvestri
1904, with several species in Europe,
Yemen, South Africa, southern South
America, southern Mexico, and western
USA and Canada (Sturm and Bach 1993,
Notario-Mun
˜oz et al. 1997, Sturm 1997),
is represented in the Nearctic by two
subspecies of M. aurantiacus (Scho
¨tt
1897): the nominal subspecies and M. a.
setosus Sturm and Bach 1992. Machili-
nus nevadensis Sweetman 1937, in-
cluded in Poole and Gentili (1997), is
anomen nudum and was considered by
Sturm and Bach (1992) to be a junior
synonym of M. a. aurantiacus.
Machilinus is easily separated from
the other genera of Meinertellidae by the
shape and placement of the ocelli, which are
subrectangular to elliptic and anterolaterally
placed in relation to the compound eyes
in Machilinus, but birotulate-elongate
(shoe-sole shaped) and generally ante-
rior to the compound eyes in the other
genera. Other key characters separating
Machilinus and its subgenera involve the
mandibular and lacinial teeth, maxillary
palpi, labial palpi, coxal styli on the legs,
and the genitalia (Sturm and Bach 1993).
The genus has been split into four sub-
genera (Sturm and Bach 1993, Sturm
1997); only Nearctolinus Sturm and Bach,
with reduced abdominal coxal vesicles
and reduced inner distance between sty-
lets on abdominal segments II and III,
occurs in North America.
Scho
¨tt (1897) described M. (Nearcto-
linus) aurantiacus from the Sierra Nevada
and Monterey, California; the original
types appear to have been destroyed
(Sturm and Bach 1992). Silvestri (1911)
discussed specimens of M. aurantiacus
he had seen from northern California,
writing on p. 350, “Exempla nonnulla per
terram apricam vagantia ad Shasta
Springs (California) legi, alia ad Boulder
Can
˜on collecta vidi.” Silvestri did not
indicate which U.S. state “Boulder
Can
˜on” was in, but earlier in the work he
had referred to Boulder Canyon, Colo-
rado, in his descriptions of Petrobius
calcaratus Silvestri and Mesomachilus
nearcticus Silvestri (op. cit., pp. 335 and
341). The reference to “Boulder Can
˜on”
was mentioned by Sturm and Bach
(1992), but attributed to the state of
Nevada.
In this paper, I provide a few additional
notes on M.(Nearctolinus) aurantiacus
and descriptions of three new species of
Machilinus (Nearctolinus).Onewascol-
lected from metropolitan Denver, Colo-
rado, the second was from the Sangre de
Cristo Mountains of northern New
Mexico, and the third was collected in
central Idaho. Other specimens are known
from western Montana, central Colorado,
central Arizona, and northern Nevada. A
few morphological comments are made
on the neotype of M. aurantiacus and the
holotype of M. a. setosus to address a few
characters that were found to be useful in
diagnosing the new species but had not
been included in existing descriptions of
M. aurantiacus. These include the penis
in males and the perianal sclerotization
and setae in both sexes.
M
ATERIALS AND
M
ETHODS
The neoholotype of M. (Nearctolinus)
aurantiacus,designatedbySturmand
Bach (1992), as well as other specimens
determined by H. Sturm to be conspecific,
were examined at the California Academy
of Sciences (CAS), San Francisco, Cal-
ifornia. Most specimens of the new spe-
cies described herein were field-collected
over the last decade by directly collect-
ing the organisms or by extractions with
a Berlese funnel.
Specimens were examined using either
a Bausch and Lomb Stereozoom 7 or
a Leica SHE stereozoom microscope
with 10x oculars. An ocular micrometer
was used for all measurements. Using
both the holotype and some non-holotype
specimens of the new species, selected
parts (antennae, maxillary and labial
palpi, head, legs, butterflied abdomen,
median terminal filament and cerci, and
ovipositor) were dissected and mounted
in CMC-10 on glass microscope slides,
ringed with clear fingernail polish, in
February 2013 and examined with an
Olympus CH-2 compound microscope.
Terminology for general morphologi-
cal features generally follows Sturm and
Bach (1992) while ovipositor type was
determined using Sturm and Bach
(1993). Identification of the basiconic
antennal sensillae and the subsequent
sensillogram is based on Notario-Mun
˜oz
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
et al. (1997). Distribution patterns in the
sensillogram table include only the last
two (n and n-1) chains (Fig. 1), except as
noted.
I differentiate three types of setae, all
of which are exemplified on the labial
palpi (Fig. 2). Hair setae are thin and
hairlike and may be short or long; hair
setae include the general clothing hairs
as well as some specialized hairs, usu-
ally identified by longer length. Spine
setae are long, a little thicker than hair
setae, and acutely pointed; spine setae
are found on the labial palpi, the basal
articles of the maxillary palpi, and in
some species on the genae laterad of the
antennal bases. Club setae are short or
long, noticeably thicker than spine setae,
and any of acutely pointed, rounded, or
truncate; club setae are found on the
front below the compound eyes, the
distal articles of the maxillary setae, and
ventrally on the tibiae and tarsi. The
frontal club setae are considered in
Sturm and Bach (1992) as “spine-like
setae.”
Exact label data are presented for
holotypes and indicated through the use
of quotation marks. A slash (/) is used to
separate the information on separate
lines. Collection data for other speci-
mens are provided with editorial com-
ments in square brackets.
R
ESULTS
Machilinus (Nearctolinus) aurantiacus
aurantiacus Scho
¨tt, 1897
This species is already adequately
described in Scho
¨tt (1897), Silvestri
(1911), and Sturm and Bach (1992). The
neotype (#16977) of M. a. aurantiacus,
designated in Sturm and Bach (1992),
is deposited in the CAS. The neotype
was completely dissected in 1990 by
H. Sturm and the parts (antennae, maxil-
lary palpi, legs, abdominal coxites II – VI,
cerci and median terminal filament, and
the head) were slide-mounted in euparal.
The slide label reads as follows (hand-
written): “Machilinus au- / rantiacus
(Schoett) / m# 6,8 mm CAS 59 / U.S.A.,
Calif., 3 km / NNE Angwin, 396 m /
12.VI.1980, H. B. / LEECH leg /
NEOH OLOT YPE / 16.VIII.90 Euparal”.
The number “59” is circled, and the word
“NEOHOLOTYPE” is outlined with
a red border.
Atotalof29Machilinus specimens in
the CAS, except the neotype, were iden-
tified as “Machilinus cf. aurantiacus”by
H. Sturm in 1990; of these, 22 specimens
closely fit the description of M. aurantiacus
with little variation, and 6 specimens match
the description of M. a. setosus.One
specimen is not M. aurantiacus and is
included as material examined for
a species described below. Micro-
coryphia are easily damaged during
collection and preservation, and most of
the specimens were collected in the
1960s and 1970s, so many of the de-
termined specimens in the CAS are in
rather poor condition. Nevertheless, ex-
amination of these specimens also re-
vealed characters that had not previously
been described or are no longer visible in
the neotype, since it had been dissected
and slide-mounted.
For example, in non-holotype male
specimens in the CAS, the penis is not
sclerotized and does not have any setae
at the tip, as incidentally drawn in figure
42 in Sturm and Bach (1992). In both
males and females, the anus is ringed
Fig. 1. Diagrammatic outline of Machilinus
antenna; n =ultimate distal chain, n-1 =penulti-
mate distal chain, annuli numbered 1 through 8
(after Notario-Mun
˜oz et al. 1997).
VOLUME 116, NUMBER 3
with a more-or-less shield-shaped ring of
lightly sclerotized cuticle, with lateral
rows of weak, light colored setae arising
from dark bases, running in a diagonal
line from near the lateral base of the
cerci anteromedially to near the anterior
end of the anus (cf. Figure 14). The fig-
ures in Sturm and Bach (1992) indicate
broad pigmentation on the front and this
is visible in the slide mounted heads, but
nearly all pigmentation on most of the
specimens in the CAS appears to have
been washed out over time, so variation
in the pigmented areas is unknown.
Scale patterns are not visible, since
nearly all scales have also been abraded
from the specimens.
Material examined in addition to the
neotype: U.S.A.: California: 2 ♂+1♀,
Contra Costa Co., 6 mi. W. Walnut Creek,
21-X-1967, L. Neil Bell; 1 ♂, El Dorado
Co., El Dorado Nat’l Forest, Blodgett, 32 km
E of Georgetown, 1524 m, 6-V-1973,
Herman G. Real; 1 ♂+2♀, Napa Co., N
side of Howell Mt., 3 km NNE of Ang-
win, 396 m, in garden under pile of
weeds, 12-VI-1980, Hugh B. Leech [ne-
oparatypes]; 1 ♀, San Francisco Co., San
Francisco, el. ;301 [units not given], 14-
X-1969, J. Hjelle; 1 ♂+2♀,Santa
Barbara Co., 7.3 mi E of US Hwy 101 on
St. Hwy 166, 27-VII-1964, P.R. & D.L.
Craig; 1 ♂+1♀, Santa Clara Co.,
IX-1992; W. Pulawski [slide-mounted]; 5
♀,SonomaCo.,3mi.N.ofSebastopol,
on moist ground, 7-VII-1964, P. Rubtzoff;
2♀, Sonoma Co., 2.5 mi N of Sebastopol,
fence post, P. Rubtzoff.
Machilinus (Nearctolinus) aurantiacus
setosus Sturm and Bach 1992
Although Sturm and Bach (1992) did
not see sufficient variation within their
material of M. auranticus to warrant
description of additional species, they
described this subspecies in which the
spine setae on the ventrodistal border of
the second segment of the maxillary palp
in the male were significantly longer
than in the nominate form and the sub-
median rows of spine setae on the male
front were stout, as in the nominate
form, but shorter (i.e., club setae).
A male from Utah was designated as
the holotype in Sturm and Bach (1992),
but no specimens in the CAS are la-
beled as the holotype. One male and
one female specimen were collected
from the type locality, with label read-
ing as follows: “1 m#, 1 f# 9 mi E Oak
City, UT / Oak Ck. Camp 1 IX 1963 /
D.C. Rentz. Molasses trap”. Both
specimens were dissected in 1990 by
H. Sturm, and parts (antennae, maxillary
palpi, legs, abdominal coxites II – VI,
cerci and median terminal filament, and
the head) were slide-mounted in eupa-
ral. Accordingly, the slide-mounted
male should be considered the holo-
type.
Fig. 2. Distal two articles of labial palp of
Machilinus taoseno, sp.n., illustrating three types
of setae.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Material examined in addition to the
holotype: U.S.A., California: 2 ♂+2♀,
Tuolumne Co., Chipmunk Flat nr. Sonora
Pass, 19-VII-1964, D.C. & K.A. Rentz
[paratypes]; Utah: 1 ♀,MillardCo.,same
data as holotype [slide-mounted paratype]
Figs. 3–15. Machilinus matadero sp.n., hair setae not illustrated except where noted. 3. Head, frontal
view, with hair setae below compound eyes; 4. male left maxillary palp; 5. male right maxillary palp; 6.
female left maxillary palp; 7. female right maxillary palp; 8. basal article of male maxillary palp, an-
terolateral oblique view; 9. ovipositor, with hair setae; 10. male labial palp; 11. male fore leg; 12. male
mid leg; 13. male hind leg; 14. male 9
th
and 10
th
abdominal semgnets, ventral view; 15. penis, with hair
setae. ald =distal extension of anterior lobe, alp =proximal extension of anterior lobe, c =cerci, dl =
dorsal lobe, mtf =median terminal filament, P=penis. 1 mm scale bar in center of figure applies to legs,
maxillary palpi, and male terminalia; other scale bars nearest appropriate part.
VOLUME 116, NUMBER 3
Machilinus (Nearctolinus) matadero
De Jong, new species
urn:lsid:zoobank.org:act:0873B45E-
BDEE-4E3D-A372-F120A061CC80
(Figs. 3 - 15)
Description.—
Male: Body length
8.15 mm (variation among paratypes:
8.11 ±0.34 [s.d.], n =7); length of an-
tennae, including flagellum: 5.67 mm;
length of cerci: 2.11 mm; length of me-
dian terminal filament: 7.04 mm (broken).
Color of body dark gray with silver re-
flections when alive, medium reddish
browntodarkgrayinalcohol.Headlarge,
ratio of contact line between contiguous
compound eyes to total length of eyes
0.71 to 0.89, ratio of total length to width
of eye 1.03 to 1.12, thin light band sep-
arating compound eye from darker pig-
mented areas of head broad laterad of the
ocelli and usually obscured poster-
omedially. Ocelli light colored, round to
oval, and anterolaterally placed in re-
lation to compound eyes, separated from
them by one-fifth width of an ocellus.
Median pigmented area on frons with
lighter areas around antennal bases, ex-
tending laterally below eye margins not
as far as ocelli and anteriorly via a nar-
row section nearly to clypeus where di-
vided into two lobes. Narrow section of
pigmented area on frons more than half
as wide as the greatest width of the an-
terior lobes of pigmented area. One pair
long hair setae submedially below com-
pound eyes; a small field of club setae of
random orientations anteriad of the long,
paired hair setae. Small patch of hair
setae on anterolateral corner of genae
laterad of antennal bases.
Antennal scape, pedicel, and first
several annuli round in cross-section, not
flattened. Distinct band of long hair setae
completely circumscribing the scape api-
cally and otherwise moderately clothed
with smaller hair setae. Antennal flagel-
lum comprised of a series of distal chains
of 4-8 individual annuli, with apical
chains much more distinct and having
more annuli/chain (generally 8). Flagellum
densely covered with hair setae and with-
out scales. Sensillae on annuli 1, 3, 5, and
7 on the n chain and on annuli 1, 3, 5, 6,
and 7 on the n-1 chain.
Maxillary palpi with extensive red-
dish brown pigmentation. Basal article
of maxillary palpi with dorsal lobe digi-
tiform and projecting nearly perpen-
dicular to article; anterior lobe bifid with
proximal extension rounded and lightly
sclerotized and distal extension bluntly
squared. Second segment of maxillary
palpi with subapical lobe angled with
a narrow, deep gap between tip and distal
end of article, broadly rounded with weak
sclerotization. Apical 5 articles of maxil-
lary palpi with dense hair setae. Apical
article with 10 to 11 club setae; penulti-
mate article with 5 club setae distributed
equidistantly within apical half, one club
seta proximally, and one club seta halfway
in proximal half (1+1+5); one club seta
subapically on antepenultimate article.
Penultimate article of maxillary palp 2.5
times length of ultimate article. Labial
palpi lightly pigmented, clothed with
moderately dense brown hair setae; third
article rounded and slightly concave with
6 club setae apically; second article
without medial spine setae.
Legs pigmented medium reddish-
brown, clothed moderately with brown
hair setae. Coxae of all legs with cloth-
ing hair setae throughout and with two
major spine setae along venter. Tibia
with 6 to 12 spine setae subapically on
ventral margin; tarsomeres with 10 to 24
spine setae mostly in apical two-thirds
ventrally; spine setae on ultimate tarso-
mere equidistantly distributed in double
rows from base to insertion of tarsal
claw. Tarsal claws light brown.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Dorsum and venter densely clothed
with scales. Stylets of urosternites light
brown, bare dorsally, and with long, dark
brown/piceous hair setae ventrally. One
pair coxal vesicles submedially on coxites
III through V. Ratio of length of stylet to
medial length of coxite on abdominal
segments II through IX: 1.13, 1.12, 0.72,
0.67, 0.76, 0.61, 1.25, 0.83. Terminal spine
tiny. Ratio of length of terminal spine seta
to length of stylet on abdominal segments
II through IX: 0.17, 0.16, 0.15, 0.25, 0.23,
0.36, 0.27, 0.17. Lightly sclerotized
shield-shaped ring surrounding anus,
broadly pointed anteriorly and rounded
posteriorly; diagonal row of strong setae
on the sclerotized ring laterad of the anus,
extending from near base of cerci ante-
romedially to anterior of the anus. Cerci
with scales throughout; no claw present.
Penis cylindrical with deep v-shaped
notch ventrally and blunt apically, apical
end papulate, entirely unsclerotized, and
with setae arising from papules around tip
and along v-shaped notch.
Female: Body length: 8.22 mm. Col-
oration, pilosity, and scales generally as
in male. Compound eyes and ocelli as in
male. Median pigmented area on frons
broader than in male, but with wider light
margins along bases of antennae and
along eyes; without very dark median
area and without submedian rows of se-
tae. Antennal scape, basal annuli, and
flagellum of antennae as in male, except
that subapical, circumferential row of se-
tae on scape not as pronounced. First ar-
ticle of maxillary palpi with similar lobes
as in male; second article without lobe,
but distinctly angular on anterior face;
apical articles with dense hair setae; 10
to 11 club setae on ultimate article, 7
club setae equidistantly distributed on
penultimate article, and one club seta
subapically on antepenultimate article.
Penultimate maxillary palp segment 2.4
to 2.5 times the length of the ultimate
maxillary palp segment. Labial palpi
lightly pigmented brown. Last article of
labial palp rounded apically with 3 to 6
club setae.
Legs of female light brown, but with
ventral spine setae not as strong as in
male on tarsomeres.
Abdomen as in male, including peria-
nal sclerotization with row of strong setae
extending from base of cerci ante-
romedially. Ovipositor quaternary type,
arising from urosternite IX, slightly
curved dorsally after it passes anus, then
slightly curved ventrally again. Ovipositor
appears quadrate with >35 divisions,
more rounded and more obviously di-
vided in apical third. One seta per division
in apical 0.75 of gonapophysis IX with
one seta at tip; one seta per division in
apical 0.33 of gonapophysis VIII with
one seta at tip. Tips of gonapophyses IX
extending beyond gonapophyses VIII by
a length of not more than two widths of
a gonapophysis.
Holotype.—
♂, dissected and mounted
on glass microscope slide in CMC-10,
coverslip ringed with clear fingernail pol-
ish. Label (computer printed with red
border): “Arapahoe County, CO / Littleton,
Slaughterhouse Gulch / N39°37’00”
W105°00’44” / 6 July 2004 / G. De Jong /
Machilinus matadero De Jong / holotype”.
Specimen is deposited in the National
Museum of Natural History (USNM),
Smithsonian Institution, Washington, DC.
Paratypes.—
7♂, same data as holo-
type; 1 ♂+8♀, same data as holotype
but 3 June 2004. One ♀is dissected and
mounted on glass microscope slide in
CMC-10 similar to holotype; the rest are
in alcohol. Three paratypes are deposited
with the holotype in the USNM; three
paratypes are deposited in the C. P. Gillette
Museum of Arthropod Diversity (CSUC)
at Colorado State University, Fort Collins,
CO; the remaining specimens are de-
posited in the personal collection of GDD.
VOLUME 116, NUMBER 3
Etymology.—
The specific epithet is
a masculine noun used in apposition,
matadero being Spanish for slaughter-
house, in reference to the type locality.
Diagnosis.—
The long line of contact
between the compound eyes (>0.7 of the
total length) separates this species from
M. aurantiacus, which has a much shorter
line of contact (;0.55 of the total
length). The ocelli are separated from the
compound eyes by a distance of 0.2 the
width of an ocellus in this species,
a character shared only by the following
species. The submedian pair of long, hair
setae immediately anterior to the com-
pound eyes is unique in this species. The
distribution of the antennal sensillae is
diagnostic among these new species, but
not compared to M. aurantiacus. The
subapical lobe on the second maxillary
segment is broadly rounded at the tip and
weakly sclerotized. The subapical club
seta on the antepenultimate maxillary
palp article is unique to this species in
Machilinus (Nearctolinus). The tips of
gonapophyses IX barely surpass the tips
of gonapophyses VIII.
Biology.—
Efforts to collect speci-
mens at the type locality have been made
in every month from March to November,
and most specimens have been collected
in June and July. Specimens were col-
lected along unpaved walking and game
trails throughout the western end of
Slaughterhouse Gulch, a small natural
area park in Littleton, Colorado. The
organisms were most common on hard-
pan soils, especially near the defaunated
disks around nests of the harvester ant
Pogonomyrmex occidentalis (Cresson),
along small game trails in the park, and
in cottonwood tree (Populus deltoides
Bertram ex. Marsh) leaf litter. They were
occasionally, but more rarely, found on
large areas of soil laid bare by human
activities (i.e., walking trails or along
railroad tracks). Predominant vegetation in
theareaincludedcommonragweed(Am-
brosia artemisiifolia L), clover (Trifolium
sp.), rabbitbrush (Chrysocamnus nause-
osus Pallas ex. Pursh), cottonwood trees,
and cheatgrass (Bromus tectorum L.).
Additional material examined.—
1♂+
8♀, same data as holotype but 14-
VII-2004. One ♀specimen was collected
1July2005atN39°33’58” W105°02’33”
on a cement walking trail (leg. GDD and
S. Blunt), and another ♀specimen was
collected 14-XI-2008 at N39°33’58”
W105°03’05” on the banks of the South
Platte River (leg. S. Kamin); both lo-
calities are in Littleton, Arapahoe
County, Colorado, about 5 km SSW of
thetypelocality.One♀specimen was
also collected 25-X-2011 at N39°48’46”
W104°57’00” (leg. GDD) on a cement
walking trail along Sand Creek in Com-
merce City, Adams County, Colorado, 22
km NNE of the type locality; this specimen
had an unusually large pale area sur-
rounding the compound eyes from the
ocelli laterally and dorsally, but otherwise
agreed in all aspects with the type material.
The later collection dates of some of these
specimens (October and November) ap-
pear to be related to the unusually warm,
dry weather during autumn of those years.
Nearby vegetation at all sites was similar to
that where the type series was collected.
One additional specimen in the CAS,
identified as “Machilinus cf. aurantiacus”
by H. Sturm, is referable to this species:
U.S.A.: Montana: 1 ♂,RavalliCo.,Birch
Creek, 31-X-1965, William L. Jellison.
Machilinus (Nearctolinus) taoseno De
Jong, new species
urn:lsid:zoobank.org:act:B6E0B3F0-
6E13-4ABD-B954-24F72E95F308
(Figs. 2, 16–23)
Description.—
Male: Body length:
6.52 mm (variation of paratypes: 6.54 ±
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
0.11 mm, n =8); length of antennae, in-
cluding flagellum: 4.81 mm; length of
cerci: 2.30 mm; length of median termi-
nal filament: 6.30 mm: Color of body
dark gray with silver reflections when
alive, light to medium brown in alcohol.
Head large, ratio of contact line between
contiguous compound eyes to total length
of eyes 0.56 to 0.86, ratio of total length
to eye width 1.27 to 1.30. Ocelli light
colored, round to oval, and anterolaterally
placed in relation to compound eyes,
separated from them by about one-fifth
width of the ocellus. Median pigmented
area on frons with lighter areas around
antennal bases, extending laterally not as
far as ocelli and anteriorly via a narrower
section nearly to clypeus where divided
into two lobes. Narrow section of pig-
mented area on frons less than one-fourth
as wide as the greatest width of the an-
terior lobes. Paired, submedian rows of
10 to 12 medially declivent club setae on
frons just below compound eyes.
Figs. 16–23. Machilinus taoseno, sp.n., hair setae not illustrated except where noted. 16. male right
maxillary palp; 17. male left maxillary palp; 18. female right maxillary palp; 19. female left maxillary
palp; 20. male head, frontal view; 21. male 9
th
and 10
th
abdominal segments, ventral view; 22. ovipositor,
with hair setae; 23. penis, with hair setae. 1 mm scale bar in center of figure applies to maxillary palpi and
male terminalia; other scale bars nearest appropriate part.
VOLUME 116, NUMBER 3
Antennal scape, pedicel, and first
several annuli dorsoventrally flattened
slightly so oval in cross-section. Scape
with distinct band of long hair setae
completely circumscribing segment sub-
apically and otherwise moderately clothed
with smaller hair setae. Antennal flagel-
lum comprised of a series of distal chains
of 4-8 annuli, with apical chains having
more annuli/chain (generally 8). Flagel-
lum densely covered with hairs and
without scales. Sensillae on annuli 1, 3,
5, 6, and 7 on the n chain and on annuli 1
through 7 on the n-1 chain.
Maxillary palpi with extensive reddish-
brown pigmentation. Basal article of
maxillary palpi with dorsal lobe dig-
itiform and nearly perpendicular to arti-
cle, anterior lobe weakly bifid apically
with proximal extension rounded and
distal extension obtuse. Second article of
maxillary palpi with posterior lobe acute
apically, subcultriform with larger gap
below, and strongly sclerotized at tip.
Apical 5 articles of maxillary palpi with
sparse hair setae (may have been
abraded). Apical maxillary palp article
with 10 to 11 club setae. Penultimate
article with distribution of club setae
asymmetrical – on right maxillary palp,
4 to 5 club setae equidistantly spaced in
apical third and 2 to 3 club setae equi-
distantly spaced in proximal two-thirds
(1+1+1+4); on left maxillary palp, 11
club setae equidistantly spaced along
length of article. Antepenultimate article
sometimes with one club seta subapi-
cally. Penultimate maxillary palp article
2.25 times length of ultimate article.
Labial palpi lightly pigmented, clothed
with dense brown hair setae; third article
of labial palpi subtruncate, slightly con-
cave and with 9 to 11 club setae apically,
second article with 7 spine setae on
medial edge.
Legs pigmented medium reddish-
brown, clothed moderately with brown
setae; long hair setae along venter of tro-
chanter, femur, and tibia of each leg from
base to apex of each leg article. Coxae
with hair setae only on apical third. Tarsi
unpigmented and clothed densely with
brownsetae;9to11spinesetaeventrally
on 1
st
tarsomere, 8 to 14 spine setae
ventrallyon2
nd
tarsomere, 14 to 17 spine
setae ventrally on 3
rd
tarsomere, not in
paired rows. Tarsal claws light brown.
Dorsum of abdomen clothed with
scales (sparsely to abundantly, depend-
ing on amount of abrasion experienced),
but scales dense on venter of abdomen.
Stylets of urosternites light brown and
clothed densely with long, dark brown to
piceous hair setae ventrally. One pair
coxal vesicles submedially on coxites III
through V. Ratio length of stylet to me-
dial length of coxite on abdominal seg-
ments II through IX: 1.10, 0.91, 0.69,
0.85, 0.64, 1.00, 1.08, 0.86. Terminal
spine tiny. Ratio length of terminal spine
seta to length of stylet on abdominal
segments II through IX: 0.27, 0.30, 0.33,
0.27, 0.22, 0.25, 0.29, 0.21. Lightly
sclerotized shield-shaped ring surround-
ing anus, broadly pointed anteriorly and
rounded posteriorly; diagonal row of
weak setae on the sclerotized area laterad
of anus extending from base of cerci
anteromedially not as far as anterior end
of anus, with setae stronger near base of
cerci. Cerci with scales on basal seg-
ments, no claw present. Penis cylindrical
with deep, v-shaped notch ventrally,
unsclerotized, slightly papillated, and
with ring of hairs arising from papules
around tip and along v-shaped notch.
Female: Body length: 7.81 ±0.09 mm
(n =5). Coloration, pilosity, and scales
generally as in male. Compound eyes
contiguous and round/oval ocelli placed
anterolaterally as in male. Median pig-
mented area broader than in male, no
submedian club setae. Antennal scape
and basal annuli of antennae slightly
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
flattened dorsoventrally, but subapical
row of long hair setae not as pronounced
as in male. Antennae divided into units of
distal chains, clothed and with sensillae
as in male. Maxillary palpi reddish-
brown; lobes on first article as in male;
second article without apical lobe, and
slightly angular; hair setae dense and club
setae asymmetrically distributed on api-
cal articles as in male. Penultimate max-
illary palp article 2.25 times longer than
ultimate article. Labial palpi lightly pig-
mented brown. Distal article truncate di-
agonally, slightly concave, and with
lighter color and 3 to 6 club setae apically.
Legs of female light brown, but with
ventral spine setae not as strong as in
male on tarsi. Tarsi light brown.
Abdomen, except genitalia, as in
male, including perianal sclerotization
with weak row of setae extending from
base of cerci anteromedially. Ovipositor
quaternary type, arising from urosternite
IX, slightly curved dorsally after it
passes anus, then slightly curved ven-
trally again. Ovipositor quadrate and
divided into >52 divisions with 1 to 2
setae per division on gonapophyses IX, 1
seta per division on terminal 12 segments
of gonapophyses VIII. Tip of gonapoph-
yses IX extending beyond gonapophyses
VIII by a length greater than four times
width of a gonapophysis.
Holotype.—
♂, dissected and moun-
ted on glass microscope slide in CMC-
10, coverslip ringed with clear fingernail
polish. Label (computer printed with red
border): “Taos County, NM / 7 mi. E
Questa / N36°41’46” W105°29’04” / 12
August 2003 / G. De Jong and R. Stitt /
Machilinus taoseno De Jong / holotype”.
Deposited in the USNM.
Paratypes.—
8♂+5♀, same data as
holotype. One ♀dissected and mounted
on glass microscope slide in CMC-10
similar to holotype; the rest are in alco-
hol. Three paratypes are deposited with
the holotype in the USNM; three para-
types are deposited in the CSUC; the rest
are retained in the personal collection of
GDD.
Etymology.—
The specific epithet is
a masculine noun used in apposition and
is derived from taosen
˜o, Spanish for
a resident of Taos, in reference to the
holotype county of origin.
Diagnosis.—
The medial contact line
between the compound eyes is 0.56 in
the holotype male because the very
narrow gap between the anterior om-
matidia extends deeper than in other
specimens of this species, in which the
length of contact is about 0.8 the length
of the eyes (e.g., 0.78 in the allotype) and
is longer than in M. aurantiacus. This
species has the longest eyes of any of the
other species of Machilinus (Nearctoli-
nus). The ocelli are separated from the
compound eyes by a distance about 0.2
the width of an ocellus. The frontal setae
on the male are in paired submedian
rows, similar to M. aurantiacus setosus,
but different from the other species. The
gena have only a single hair seta laterad
of the antennal scapes, whereas the other
described species have small clumps of
setae. The perianal sclerite has two di-
agonal rows of setae – one with a few
strong setae and the other with several
weaker setae – other species have only
a single row. The tips of the dorsal/pos-
terior gonapophyses of the ovipositor
clearly surpass the tips of the ventral/
anterior gonapophyses by at least four
times the width of a gonapophysis.
Biology.—
This species is active at
least from April to August; attempts to
collect specimens in the months of
March, September, and October have
been unsuccessful. Soil fauna from an
1800 cm
3
litter and soil sample collected
12-VIII-2003 at the type locality were
extracted in a Berlese funnel for five
days, yielding two of the paratype
VOLUME 116, NUMBER 3
specimens of M. taoseno. Also present in
the Berlese extraction were collembolans
(Hypogasturidae, Entomobryidae, Smin-
thuridae), mesostigmatid mites (No-
thridae), prostigmatid mites (Bdellidae),
one immature jumping spider (Salt-
icidae), and an immature reduviid bug.
These data indicate that M. taoseno were
present in the litter at a density of just
over one organism per liter of soil and
litter; however, a sample of 1000 cm
3
of soil and litter collected 27-IX-2004
yielded no additional specimens.
All of the New Mexican specimens of
M. taoseno were collected at the type
locality, at the mouth of a small canyon
100 m south of the Red River. The
bristletails wandered in the open sun
(rarely observed when overcast) in lit-
ter beneath a large southwestern pon-
derosa pine tree (Pinus ponderosa
brachyptera Engelmann). They were
most common in areas where pine
needle and pinecone duff had accu-
mulated in a thin layer but were un-
common on completely bare ground.
Multiple attempts in 2004 and 2005 to
collect additional specimens elsewhere
in the Red River canyon in apparently
similar habitats were unsuccessful.
Additional material examined.—
6♂
+2♀, same data as holotype but 11-13-
VI-2003, G. De Jong. One ♀speci-
men was collected 21-IX-2005 in Teller
County, Colorado, near the outfall of the
Carlton Tunnel into Fourmile Creek (leg.
GDD). GPS coordinates for the site are
N38°39’40” W105°13’20”, and the site
was at an elevation of 2,103 m above sea
level. This specimen was found wandering
in open sun on bare ground, with little leaf
litter present. Nearby vegetation consisted
primarily of sunflowers (Helianthus sp.).
One ♀specimen was collected 29-IV-
2008 in Pinal County, Arizona, in an
aquatic biomonitoring sample from Queen
Creek (leg. J. Volosin). Because of the
circumstances of its collection, it was
quite damaged, but the asymmetrical
distribution of club setae on the maxil-
lary palpi exactly matches that of this
species.
Machilinus (Nearctolinus) zingiberus
De Jong, new species
urn:lsid:zoobank.org:act:DEDDD1F2-
4949-4FA3-BBF3-EDED3EBCEEAE
(Figs. 24–27)
Description.—
Male: Body length
6.15–7.13 mm; length of antennae, in-
cluding flagellum, >3 mm (broken);
length of cerci: 1.81 mm (broken); length
of median terminal filament: 4.96 mm.
Color of body gray/brown when alive,
light brown/orange in alcohol. Head
large, compound eyes contiguous, ratio
of contact line to total length of eyes
0.65, ratio of total length to width 1.18.
Ocelli light colored, oval, and ante-
rolaterally placed, contiguous with
compound eyes. Pigmented area on
frons extensive, separated from com-
pound eyes and bases of antennae by
a thin margin, and extending laterally
to touch ocelli and anteriorly via a nar-
rower section nearly to clypeus where
divided into two lobes. Narrow section
of pigmented area on frons more than
one half as wide as the greatest width of
the anterior lobes. Field of 24 randomly-
oriented club setae on front medial-
ly below eyes. Anterolateral corner of
genae with small clump of spine setae
laterad of antennal bases.
Antennal scape, pedicel, and first sev-
eral segments cylindrical. Pedicel with
distinct subapical band of long hair setae
medially and extending ventrally, other-
wise moderately clothed with small, light
brown hair setae. Antennal flagellum with
distal chains of 4 to 8 annuli. Flagellum
densely clothed with fine hair setae,
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
scales absent. Sensillae on annuli 1, 2, 3,
5, and 7 on the n chain and on annuli 1,
3, 5, and 7 on the n-1 chain (based on
more proximal chain since antennae
broken).
Maxillary palpi reddish-brown with ir-
regular lighter areas. Basal article of
maxillary palpi with dorsal lobe digitiform
and nearly perpendicular to article; ante-
rior lobe bifid with proximal extension
strongly rounded and sclerotized and
apical extension broadly rounded to
bluntly squared. Second article of max-
illary palpi with posterior lobe acute and
broadly sclerotized. Apical 5 articles of
maxillary palpi with dense hair setae,
which are darker dorsally than ventrally.
Apical article with 6 to 7 club setae,
penultimate article with 7 to 8 club setae
equidistantly arranged along length of
Figs. 24–27. Machilinus zingiberus, sp.n., hair setae not illustrated except where noted. 24. male
head, frontal view; 25. male left maxillary palp; 26. male right maxillary palp; 27. penis. 1 mm scale bar
applies to maxillary palpi; other scale bars nearest appropriate part.
VOLUME 116, NUMBER 3
article. Penultimate article 2.2 times
length of ultimate article. Labial palpi
light brown, moderately clothed with
dark hair setae; third article subtruncate
and slightly concave with 8 to 11 spine
setae apically, 5 to 6 spine setae along
medial margin of second article.
Legs light brown, moderately clothed
with fine, brown hair setae. Coxal hair
setae fine and abundant, with one long
seta dorsolaterally. Tarsi light brown
with dense, brown hair setae; 1
st
tarso-
mere with 3 ventral spine setae subapi-
cally on foreleg and 10 to 11 spine setae
on other legs; 2
nd
tarsomere with 14 to
17 spine setae; 3
rd
tarsomere with 9 to 14
pairs of spine setae from base to in-
sertion of tarsal claw. Tarsal claws di-
vergent, lightly pigmented brown.
Dorsum of abdomen sparsely clothed
with scales (which may have been
abraded), and scales dense on venter.
Stylets of urosternites light brown, with
few weak hair setae dorsally and strong,
long, black hair setae ventrally (stylets
broken from abdominal segment II). One
pair coxal vesicles submedially on seg-
ments III through V. Ratio of length of
stylet to medial length of coxite on ab-
dominal segments III through IX: 1.23,
0.88, 0.82, 1.18, 0.89, 0.91, 0.49. Ter-
minal spine tiny. Ratio of length of ter-
minal spine seta to length of stylet on
abdominal segments III through IX:
0.19, 0.20, 0.22, 0.23, 0.38, 0.40, 0.21.
Lightly sclerotized shield-shaped ring
surrounding anus, broadly pointed ante-
riorly and rounded posteriorly; row of
thin anal setae long and evenly placed.
Cerci with scales on basal segments and
with strong, black setae medially on
several basal segments; no cercal claw.
Penis cylindrical with deep v-shaped
notch ventrally, lightly sclerotized and
papulate at tip, with setae arising from
papules along tip and along v-shaped
notch.
Female: unknown.
Holotype.—
♂, dissected and mounted
on glass microscope slide in CMC-10,
coverslip ringed with clear fingernail
polish. Label (computer printed with red
border): “Custer County, ID / Thompson
Cr.1mi.Nconfl.SalmonR./N44°15’50
W114°30’50” / 21 August 2008 /
D. Conklin, G. De Jong, J. Mullen, &
S. Kamin / Machilinus zingiberus De Jong /
holotype”. Specimen is deposited in the
USNM.
Paratype.—
1♂, Elko Co., Nevada,
Well s , N 41°6’11” W114°58’25”, 14-VII-
2012, S. Kamin. Paratype, in alcohol, is in
the personal collection of GDD.
Etymology.—
The specific epithet is
a masculine adjective, from the Latin
zingiber, meaning ginger, in reference to
the marking on the front shaped like
a gingerbread man with extremely long
arms.
Diagnosis.—
The medial line of con-
tact between the compound eyes is 0.65
the length of the compound eyes, a little
longer than in M. aurantiacus and shorter
than in M. matadero and most M. taoseno
The ocelli are contiguous with the com-
pound eyes, a character shared with
M. aurantiacus, but different from the
other species. This species shares the me-
dian field of club setae below the eyes with
M. matadero, and is different from the
paired submedian rows in M. aurantiacus
and M. taoseno. The setae on the perianal
sclerotization are long, fine, and light
colored, separating M. zingiberus from
M. matadero and M. taoseno, which
have at least some strong, dark setae.
The penis is lightly sclerotized at the tip,
a feature not seen in the other species in
Machilinus (Nearctolinus).
A sensillogram is provided for this
species even though it cannot be based
on the n and n-1 distal chains, since the
antennae were broken. Notario-Mun
˜oz
et al. (1997) found that the distal chains
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
in Machilinus (Machilinus) are repeating
units (n =n-2 =n-4, etc. and n-1 =n-3 =
n-5, etc.), so the use of chains that that
are more proximal than n and n-1 may be
acceptable. The sensillogram differenti-
ates this species from M. matadero and
M. taoseno.
Biology.—
This species has been col-
lected in July and August. Vegetation in
the immediate vicinity of the type locality
was primarily riparian willows (Salix
sp.). The single topotypical specimen was
collected at the toe of a slate talus in open
sun. The collection locality for the Ne-
vada specimen of M. zingiberus is only
45 km northeast of Soldier Canyon, the
type locality for M. nevadensis (a nomen
nudum). It is possible that Sweetman
(1937) may have been referring to this
species; however, M. nevadensis was in-
adequately described to be definitive.
D
ISCUSSION
The addition of these three species
increases the known number of Micro-
coryphia in the Nearctic Region to 33
species. Sturm and Bach (1992) de-
scribed M. a. setosus at the subspecific
level because the differences from the
nominate form were primarily in the
distribution and number of club setae.
These new species differ based on a suite
of characters outlined in Table 1, in-
cluding the distribution of spine and club
setae, basiconic antennal sensillae, and
the shapes of the eyes. The new species
are described from specimens generally
along the eastern limit of the known geo-
graphical distribution of M. aurantiacus,
which may explain why they had not
been previously recognized.
The penis of M. aurantiacus was in-
cidentally drawn in Sturm and Bach
(1992) and is not diagnostic as figured,
but examination of non-holotype speci-
mens indicated that the tip does not
have hairs as in the presently described
species. The hairs are visible at 80x
magnification, but the papules are not.
Nevertheless, the penii in the newly
described species are nearly indis-
tinguishable from each other (differ-
ing only in the slight sclerotization in
M. zingiberus), suggesting that this is
not a good character.
Conversely, the sclerotized, shield-
shaped ring of cuticle surrounding the
anus and the pattern of hair setae on that
sclerotized ring have not been described
for species of Machilinus in other sub-
genera but were found in the present
study to be good characters for separat-
ing the species described. There are two
lobes on the basal maxillary palp article:
a dorsal lobe and a divided anterior lobe.
Surprisingly, the anterior lobe has not
been reported earlier, but it was hinted at
in figures (e.g., Sturm and Bach 1992
figure 31 and cf. figure 58 for Prae-
machilellus rentzii Sturm and Bach).
This character is visible when focusing
past the dorsal lobe with the microscope,
since the dorsal lobe is the one that is
most obvious when the palp is mounted
dorsal side up.
The sensillogram, which has been
described for several European Machi-
linus (Machilinus) and has proved to be
a good character for species diagnosis (e.
g., Notario-Mun
˜oz et al. 1997, 2000), is
likewise useful in Machilinus (Nearcto-
linus) despite not examining it in
M. aurantiacus. Notario-Mun
˜oz et al.
(1997) described seven types of basi-
conic sensillae, in addition to the “ro-
setenfo
¨rmige” sensillae, in Machilinus
(Machilinus). Only the “rosetenfo
¨rmige”
and B, C, and D types of basiconic sen-
sillae were found in the Machilinus
(Nearctolinus) species in the present
study.
The biological notes presented ap-
pear to indicate a preference for sunny,
xeric habitats, as has been reported or
VOLUME 116, NUMBER 3
supposed for most other species of
Microcoryphia (Sweetman 1937, Sturm
and Bach 1993, Sturm 1997), although
a couple species have been discovered
elsewhere that prefer more moist en-
vironments (Wygodzinski 1967, Allen
1995). Dominant vegetation in the hab-
itat is inconsistent among the species.
Sturm and Bach (1992) used the
reduction of the abdominal exsertile
vesicles, the terminal spines of the ab-
dominal stylets, and the inner distance
between the stylets on the second and
third abdominal coxites, to indicate that
Nearctolinus was the most derived of the
three subgenera of Machilinus that they
described. Sturm (1997) reported that
Protomachilinus was the least derived
subgenus when it was described. The
subgeneric characters of Nearctolinus
are present in M. matadero,M. taoseno,
and M. zingiberus, except that neither
the coxal vesicles nor underlying muscle
were found in the second abdominal
segment in these three species (they are
present in M. aurantiacus, according to
Sturm and Bach 1992). The specimen of
M. matadero collected on the banks of
the South Platte River (Additional Ma-
terial Examined) had one coxal vesicle
exserted on urosternite V, indicating that
exsertion is possible in this subgenus,
despite doubts by Sturm and Bach (1992)
that they retained that ability; however,
Table 1. Table of diagnostic characters for North American species and subspecies of the genus Machilinus.
Character/Species
M. auranticus
aurantiacus
M. aurantiacus
setosus M. matadero M. taoseno M. zingiberus
Ratio of contact line
to total length in
compound eyes
0.5 – 0.6 0.5 – 0.6 0.71 – 0.89 0.56–0.86 0.65
Ratio of total length
to width in
compound eyes
0.9–1.1 0.9–1.1 1.03–1.12 1.27–1.30 1.18
Ocelli relative to
compound eyes
Contiguous Contiguous Separated by 0.2
width of ocellus
Separated by
0.2 width of
ocellus
Contiguous
Sensillogram
(sensilla types based
on Notario-Mun
˜oz
et al. 1997)
Not examined Not examined annulus n n-1 annulus n n-1 annulus n n-1
1 2C 1B3C 1 5C 5C 1 1B 3C
2 2 1C 2 1C
3 1B1C 1B3C 3 4C 3C 3 1B2C
4 4 2B2C 4 2B3C
5 1B3C 2C 5 2C2D 2C 5 1B1C
6 1B 6 1B1C 1B 6 2B2C
7 1B1C 7 2C 1B2C 7 1B1C
1B3C2D
Medial frontal setae,
in addition to normal
clothing hair setae
Male: Small
field of club
setae; Female:
none
Male: 8–10
club setae
in paired
submedian rows;
Female: none
Male: Paired long
hair setae immediately
below eyes, then large
field of randomly-oriented
club setae; Female: paired
long hair setae present, but
field of club setae absent
Male: 10–12
club setae in
paired
submedian
rows; Female:
none
Male: Large
field of 24
club setae
on front
below eyes;
Female:
unknown
Setae on
anterolateral
corner of genae
Not examined Not examined Small patch of hair setae One long hair
seta
Small clump
of spine setae
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
of the 85 specimens of Machilinus
(Nearctolinus) that I examined, it is
the only specimen in which I have ob-
served this phenomenon. Sturm (1997)
reported that species in Machilinus
(Protomachilinus), from Mexico, have
functional coxal vesicles.
Several other specimens of Machili-
nus were examined in the CAS using
a dissecting microscope; however, be-
cause they were extensively damaged, it
was not possible to refer them to any of
these species. Dissection and slide-
mounting may provide useful in deter-
mining the identity of these specimens.
Mexico: Baja California Norte: 1 ♀, Isla
de Guadelupe, “Twin Cinder Cones”, 14-
VII-1976, Vincent F. Lee; U.S.A.: Cal-
ifornia: 2 ♂+7♀, Mono Co., 4 miles
N Benton, el. 1700 m, 31-VIII-1980 – 25-
IV-1981, Derham Giuliani.
Pedetontus californicus (Silvestri 1911)
Finally, attempts were made 20 August
2009 and 14 August 2013 to collect ad-
ditional specimens of M. zingiberus at
the type locality, but, instead, two female
individuals (one immature and the other
teneral) of the machilid Pedetontus cal-
ifornicus (Silvestri) were collected in
2009 and four mature individuals (three
males and one female) were collected in
2013. This species was identified on the
basis of the lack of scales on the antennal
flagellum; 2+2 eversible vesicles on ur-
osternites II through VI; sternites obtuse
caudally, in length just over half the
length of its urosternite; sole-shaped an-
terolateral ocelli, not greatly constricted;
medial line of contact of compound eyes
less than half the length of the eyes; and
conical terminal segment of the labial
palpi. This collection represents a new
state record for P. californicus for Idaho,
as this species was previously known
only from California (Silvestri 1911,
Ferguson 1990, Mendes 1990). Numer-
ous, weathered machilid exuviae, prob-
ably from P. californicus, were also
found attached to the undersides of pieces
of slate, a common archaeognathan habit
reported in Ferguson (1990).
A
CKNOWLEDGMENTS
I thank James Chadwick, Steve Canton,
Don Conklin, Jr., and Lee Bergstedt (GEI
Consultants, Inc.) for allowing me some
limited time and resources to pursue this
project. Norm Penny and Jere Schweikert
kindly facilitated my visit to the CAS.
Three anonymous reviewers provided
helpful comments on the manuscript.
Literature Cited
Allen, R. T. 1995. Pedetontus gershneri,anew
species of Machilidae from the Interior High-
lands of North America (Insecta: Micro-
coryphia). Entomological News 106: 195–198.
Ferguson, L. M. 1990. Insecta: Microcoryphia
and Thysanura, pp. 930–949. In D. L. Dindal,
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