164
Journal of Vector Ecology
December, 2000
A taxonomic review of the North American genus Orchopeas Jordan, 1933
(Siphonaptera: Ceratophyllidae: Ceratophyllinae)
Robert E. Lewis
Department of Entomology, Iowa State University, Ames, IA 50011-3222 USA
Received 4 February 2000; Accepted 22 May 2000
ABSTRACT: The 20 named taxa assigned to the North American flea genus Orchopeas Jordan, 1933 are reviewed.
Four of these are treated as junior synonyms and the fifth as a nomen dubium. These are: Orchopeas sexdentatus
firemani Hubbard, 1955 = Orchopeas schisintus (Jordan, 1929); Orchopeas schisintus nevadensis (Jordan,
1929) = Orchopeas schisintus agilis (Rothschild, 1905); Orchopeas howardi bolivari Barrera, 1955 =
Orchopeas howardi (Baker, 1895); Orchopeas howardi texensis Eads, 1950 = Orchopeas fulleri Traub, 1950
and Orchopeas labiatus (nomen dubium). Six taxa originally assigned as subspecies of Orchopeas sexdentatus
are elevated to the rank of full species and distribution and host preferences of all 15 species are discussed.
Illustrations of the diagnostic features of all the species are provided. Journal of Vector Ecology 25(2): 164-189.
2000.
Keyword Index:
Orchopeas, taxonomy, distribution, hosts.
Depending upon whose classification is followed,
historically, the genus Orchopeas has contained 10 or
11 species and 20 named forms. All are exclusively
Nearctic in distribution except for one that was
accidentally introduced into England on the eastern
gray squirrel, Sciurus carolinensis. The southernmost
extension of the genus is said to be the mountains of
Venezuela according to Tipton and Machado-Allison
(1972), although, as pointed out elsewhere in this
study, there is reason to doubt the validity of this
record.
Before becoming involved with the various components of the genus a brief word seems in order about
an enigmatic taxon that has dogged taxonomic students
of the order since it was erected by Baker in 1904.
The status of the monotypic Orchopeas labiatus
(Baker, 1904) has been suspect since Jordan (1929)
suggested that it was closely allied to O. caedens and
O. nepos. Subsequent authors have retained O. labiatus
as a valid species, usually with the caveat that it might
be only an abnormal specimen of O. caedens. I. Fox
(1940) illustrated the seventh sternite of the holotype
female, which is the only known specimen, and bears
the USNM No. 6909. Given the variable nature of this
sternite in Orchopeas, this female could belong to a
number of species assigned to the genus. Perhaps the
strongest reason for retaining O. labiatus as a full
species has been the fact that the spermatheca appears
more globular than in females of other taxa. Adams and
Lewis (1995) noted that this specimen was remounted
in 1927. During the remounting of fleas on microscope
slides it is easy to distort the anatomical structures that
are used for taxonomic discrimination. This includes
the genitalia, which are critical in the identification of
fleas in both sexes. The usual type of spermathecal
distortion involves crushing, which causes the bulga to
appear more globose than is normal. Except for this
character, the specimen of O. labiatus is inseparable
from females of O. caedens (Jordan, 1925) a wellknown ectoparasite of red squirrels throughout their
range in North America. It is my contention that, given
the history, the condition of the specimen (which I
have examined) and its similarity to females of a
number of other taxa assigned to this genus, the true
identity of O. labiatus will never be known with
certainty. However, to synonymize the name with one
of its closest congeners, O. caedens or O. nepos,
would disrupt the stability of one or the other of two
well-established names of North American taxa. For
this reason I am treating the name Orchopeas labiatus
(Baker, 1904) as an unused nomen dubium and it will
not be treated further in this study. The literature
citation pertaining to it is included under Orchopeas
caedens (Jordan, 1925) but this is not to be construed
as a formal synonymy.
It has long been accepted that the diagnostic sex in
fleas is the male and that chaetotaxy and genitalic
configuration form the basis for taxonomic discri-
December, 2000
Journal of Vector Ecology
mination in almost all species. This includes the shape
of the fixed and movable processes of the clasper and
the apex of the ninth sternite. In some species the
shape of the eighth sternite may also be useful, but in
most it is very similar and in species where it is present
it is usually difficult to see due to its highly membranous
nature. Males of all species of Orchopeas possess
spiniform setae along the caudal margin of the movable
process, as well as a single spiniform on the caudal
margin of the ventral lobe of the apical arms of the
ninth sternite. While the number and spacing of these
tend to be relatively constant among individuals of the
same species there are sufficient exceptions to nullify
this character as the sole means of separation,
particularly among closely related taxa.
As is true for most ceratophyllid genera, females
of the species assigned to this genus for the most part
lack distinctive characters that are useful in taxonomic
discrimination. Historically, the most important
characters in this sex have been the form of the
spermatheca and the shape of the caudal margin of the
seventh sternite. In Orchopeas females the bulga and
hilla of the spermatheca are so similar in size and shape
as to render this organ useless for identification
purposes. On the other hand, the marginal contours of
the seventh sternite, while highly variable, are somewhat
more useful when used in combination with the known
range of the taxon in question. However, sufficient
variability exists in closely related species making
absolute determination practically impossible in the
absence of accompanying males, especially in the
sexdentatus and howardi species groups. As Hubbard
(1947) so aptly pointed out “. . .there is little doubt that
as time goes on, range will play a far greater part in
identifications than it has in the past.” As an aid to the
identification of females, I have included illustrations
showing the degree of variability in the seventh sternite.
Admittedly, there is some overlap in a few cases, but
the majority of females fall within the median of the
extremes. Where possible, all specimens illustrated
are from the same population.
Although the species are listed in alphabetical
order in the systematics section, the 15 species
recognized here fall into four reasonably well-defined
species groups based on genitalic characters and host
preferences.
The nepos group also includes O. dieteri and O.
latens. Only O. nepos is well-known. These three
species are restricted to the far western states and all
are evidently specific parasites of squirrels belonging
to the genera Sciurus and Tamiasciurus.
The sexdentatus group also includes O. agilis, O.
cascadensis, O. illinoiensis, O. intermedius, O. penn-
165
sylvanicus, and O. schisintus. All but O. illinoiensis,
which was only recently described, have usually been
treated as subspecies of O. sexdentatus; and all are
specific to woodrats of the genus Neotoma. However,
there are numerous records from other small rodents,
particularly those associated with woodrat lodges,
such as Peromyscus species. Only O. illinoiensis and
O. pennsylvanicus occur east of the Mississippi
River.
The caedens group also includes O. neotomae.
Orchopeas caedens is distributed from eastern Canada
and New England to the western Rocky Mountains
and Alaska and is specific to Tamiasciurus hudsonicus.
Orchopeas neotomae is restricted to New Mexico,
Arizona, and parts of Colorado and Texas, south into
Mexico, and parasitizes Neotoma species. I have seen
one male reputedly from Los Angeles County,
California. The two are grouped together because of
genitalic similarity.
The howardi group also includes O. leucopus and
O. fulleri. As the name suggests, O. leucopus is mainly
a parasite of Peromyscus leucopus and P. maniculatus.
This species is widespread throughout most of subarctic
North America. The remaining species are associated
with Sciurus species. Orchopeas howardi is confined
to North America east of the Rocky Mountains.
Orchopeas fulleri is so poorly known that its range has
not been completely determined. It is treated here as a
parasite of tree squirrels based on records reported in
Ayala-Barajas et al. (1988) and may be only a
southwestern variant of O. howardi.
Genus Orchopeas Jordan, 1933
Bakerella. Wagner, 1930, Parazit. Sb. 1: 101, 119.
Generitype: None designated at publication. See
Lewis and Lewis (1989) for a detailed history of
this name. Synonymized with Orchopeas by
Wagner, 1935, Konowia 14: 220. Smit and
Wright, 1965, Hamb. Zool. Mus. Inst. 62: 6,
designated Pulex wickhami Baker, 1895, as the
type species of Bakerella Wagner, 1930.
Orchopeas. Jordan, 1933, Novit. Zool. 39(1): 71.
Generitype: Pulex wickhami Baker, 1895
(=Pulex Howardii Baker, 1895). This genus
falls into Jordan’s (1933) “Group A” in which
the outer surface of the forefemur bears one or
no lateral bristles.
Until now there have been no keys to all of the
species in this genus. I. Fox (1940) provided one for
the four eastern species known to him, and Hubbard
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Journal of Vector Ecology
(1947) one for the nine western taxa known to him.
However, with the exception of O. s. neotomae, he did
not distinguish between the subspecies of O.
sexdentatus, and although he mentioned O. labiatus,
he did not list diagnostic features for this taxon.
Many of the taxa assigned to this genus were
poorly circumscribed originally, and a number of those
assigned as subspecies of O. sexdentatus were based
on characters that have subsequently been shown to be
so variable as to be worthless for taxonomic
December, 2000
discrimination. The following is an attempt to deal with
the currently recognized taxa. I am fully aware of the
presence of populations in both the east and the west
that do not conform to all of the parameters followed
in this revision. However, until additional material
from relatively unexplored areas of North America
comes to hand, a thorough understanding of the genus
will not be possible. In the interim, following is a
modest attempt to bring some order to a genus that
historically has had a confusing nomenclatural past.
Key to the Species of Orchopeas Jordan, 1933
1 Labial palpi not reaching apex of forecoxae ............................................................................................... 2
Labial palpi extending to or beyond the apex of the forecoxae ................................................................. 4
2 Movable process of male clasper 2.5X as long as wide at widest point (Fig. 3), dorsal apex of male fixed
process rounded, almost extending to apex of movable process, apex of male st IX blunt (Fig. 18), squared
apically and bent caudad, apical 3/4 of male st VIII spiculose and with a vermiform appendage on its dorsal
margin (Fig. 33), caudal margin of female st VII with a lobe subtended by a deep sinus which sometimes
shows a slight submarginal thickening (Fig. 48). (Coastal southern
California on Neotoma species) ............................................................................................... sexdentatus
Movable process of male 2X or less as long as wide, clavate, dorsalapex of male fixed process
oblique apically, caudal margin of female st VII bearing a lobe subtended by a deep sinus ...................... 3
3 Movable process of male with a uniformly convex caudal margin (Fig. 13), apex of fixed process noticeably
flexed cephalad, apex of distal arm of male st IX with dorsal and ventral lobes pointed caudally (Fig. 28),
st VIII with a distinct bulge on dorsal margin but lacking a vermiform appendage (Fig. 43), dorsal lobe of
female st VII straight or slightly convex and subtended by a deep sinus above
the rounded ventral margin (Fig. 58). (North-central to southern California on Sciurus species.) . latens
Movable process of male clavate, its caudal margin angulate in the middle, its ventrocaudal margin slightly
concave (Fig. 10), male fixed process bluntly pointed dorsally, apex of distal arm of male st IX with dorsal
lobe rounded caudally, ventral lobe pointed apically (Fig. 25), male st VIII small, fimbriated basally, its apex
forming a vermiform, unspiculated appendage (Fig. 40), caudal margin of female st VII with a dorsal lobe
subtended by a shallow sinus and a rounded ventral lobe (Fig. 55). (Widespread throughout the United States
and Canada on small rodents, especially Peromyscus
leucopus and P. maniculatus.) ..................................................................................................... leucopus
4 Frontal setal row incomplete, consisting of 0 or 1 long seta, not counting the minute setulae bordering the
cephalic margin of the antennal groove, especially in males, (O. illinoiensis, included here, though
obviously a member of the sexdentatus species group, frequently has a single seta on
one side and 2 on the other in both sexes.) ................................................................................................. 5
Both ocular and frontal rows complete, the latter typically with 2-4 long setae per side ....................... 11
5 Movable process of male shaped like a scalene triangle, its apex extending ~1/3 beyond the apex of the
narrowly rounded, rather pointed apex of the fixed process (Fig. 4), cephalic margin of apical lobe of male
st IX distinctly concave, its caudal apex roundly pointed (Fig.19), distal arm of st VIII with a spiculate apex
and a well-developed hump and vermiform appendage on its dorsal margin (Fig. 34), female st VII variable,
from smoothly rounded (Fig. 49) to bearing a caudal lobe subtended by a shallow sinus. (Presently known
only from southern Illinois, central Missouri, and western
Kentucky where it parasitizes Neotoma floridana.) ................................................................. illinoiensis
December, 2000
Journal of Vector Ecology
167
Movable process of male clavate, more rectangular or square in shape, its caudal margin bearing 4-5
spiniforms, contours of male st VIII and IX variable, caudal margin of female st VII variable but
usually lobed and/or bearing a sinus ........................................................................................................... 6
6 Movable process of male clavate, its ventrocaudal margin strongly concave, with 5 along the dorsocaudal
margin (Fig. 9), dorsal apex of fixed process blunt, with acetabular setae arising on the dorsocaudal 1/3,
dorsal lobe of apical arm of male st IX (Fig. 24) smoothly rounded, acuminate dorsocaudally, ventral lobe
pointed ventrocaudally, apical 3/4 of distal arm of st VIII slightly inflated, spiculate and bearing a small
vermiform appendage on its dorsal margin (Fig. 39), caudal margin of female st VII variable, from bearing
a sharp to blunt lobe subtended by a deep to shallow sinus, to entire (Figs. 54, 69). (Widespread throughout
North America, mainly on Tamiasciurus
hudsonicus and its predators.) ........................................................................................................ caedens
Movable process more rectangular or square, contours of male st VIII and IX variable, caudal margin of
female st VII variable but usually bearing a small to pronounced lobe subtended by at least a small
sinus ............................................................................................................................................................. 7
7 Movable process of male roughly rectangular, ~2.5X as tall as wide, its caudal margin bearing 4
spiniforms (Figs. 14, 15), female st VII with lobe and sinus ..................................................................... 8
Movable process of male more square, much less than 2X as tall as wide at widest point, its caudal
margin bearing 4-5 spiniforms (Figs. 11, 12), female st VII with ill-defined lobes and sinusus .............. 9
8 Male movable process 2.5X as tall as wide at widest point (Fig. 14), dorsal lobe of fixed process pointed,
acetabular setae arising about midway up the caudal margin on a slight hump, dorsal margin of dorsal lobe
of distal arm of male st IX flat to slightly concave (Fig.29), its ventral lobe broadly pointed, ~ basal 1/2 or
apical 3/4 of male st VIII with parallel margins (Fig. 44), without a vermiform appendage on dorsal margin,
caudal margin of female st VII slightly concave above a small, pointed
ventral lobe and sinus (Fig. 59). (Pacific coastal southern California on Sciurus griseus.) .......... dieteri
Movable process slightly less rectangular, 2.75X as tall as wide at widest point (Fig. 15), dorsal extension
of fixed process broad, smoothly rounded apically, almost reaching apex of movable process, acetabular
setae arising ~1/3 down caudal margin, dorsal margin of dorsal lobe of distal arm of st IX smoothly rounded
and convex, its ventral lobe strongly attenuated and sharply pointed (Fig. 30), apical 3/4 of male st VIII
strongly spiculose, its dorsal margin bearing a small vermiform appendage (Fig. 45), caudal margin of
female st VII usually with a conspicuous, well-developed marginal lobe subtended by a deep sinus (Fig 60).
(Southern British Columbia to south-central
California, usually on Tamiasciurus douglasi, sometimes other squirrels and their predators.) .... nepos
9 Male movable process more angular, only slightly taller than wide at its widest point, its caudal margin with
a distinct angle above which are 4 spiniforms (Fig. 11) both upper and lower lobes of distal arm of st IX are
rounded caudally but the dorsal margin of the upper lobe is straight or slightly convex (Fig. 26), apical 3/
4 of male st VIII spiculose, its dorsal margin bearing a small vermiform appendage (Fig. 41), caudal margin
of female st VII usually slightly concave above a slight lobe and sinus (Fig. 56). (Widespread in North
America east of the Rocky Mountains where Sciurus niger
and S. carolinensis occur, or where they have been introduced.) .................................................. howardi
Male movable process obviously taller than wide, its caudal margin smoothly rounded and usually bearing
5 spiniforms (Fig. 12), its ventrocaudal margin slightly concave, both upper and lower lobes of distal arm
of st IX rounded caudally but dorsal margin of upper slightly concave, apical 4/5 of male st VIII spiculose,
its dorsal margin with a small vermiform appendage (Fig. 42), caudal margin of female st VII straight or
slightly convex above the shallow sinus (Fig. 57). (Southwestern United States, into Mexico where it
parasitizes Sciurus aberti and other members of that genus. Poorly
known.) ............................................................................................................................................... fulleri
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Journal of Vector Ecology
December, 2000
10 Male movable process over 3X as tall as wide at widest point, its upper half subtriangular with itscephalic
and caudal margins subparallel, 4 spiniforms on caudal margin, the upper 2 widely spaced, the lower 2 arising
close together near caudal angle (Fig. 8), upper lobe of distal arm of male st IX claw-like, its dorsal margin
almost straight, its apex hooked (Fig. 23), apical 4/5 of st VIII heavily spiculose, its dorsal margin bearing
a vermiform appendage (Fig. 38), caudal margin of female st VII with a broad, paddle-shaped lobe subtended
by a deep sinus (Fig. 53). (Rocky Mountain region of New Mexico and Arizona, extending into Colorado,
Texas, and Mexico, as a
parasite of various species of Neotoma.) ..................................................................................... neotomae
Male movable process 2.5X or less as tall as wide at widest point, upper spiniform on caudal margin remote
from remaining spiniforms, lobes of distal arm of male st IX variable, as is the configuration of st VIII, caudal
margin of female st VII variable but usually bearing some kind of a lobe, with or
without a sinus ........................................................................................................................................... 11
11 Caudal margin of male movable process bearing a distinct projecting angle, the margin below it strongly
concave, dorsal projection of fixed process with parallel margins and a broadly rounded apex (Figs. 6, 7),
caudal projection of upper lobe of distal arm of male st IX drawn out to a relatively sharp to bluntly rounded
point (Figs. 21, 22), male st VIII with a vermiform appendage on dorsal margin, caudal margin of female st
VII usually with a conspicuous lobe subtended by a deep sinus (Note: From this point, females are so
variable that accurate identification in the absence of males
is practically impossible.) ......................................................................................................................... 12
Caudal margin of male movable process rounded dorsally, the margin below it almost straight, slightly
concave (Figs. 1, 2, 5), dorsal projection of fixed process with subparallel margins and tapering to a rounded
apex, upper lobe of distal arm of male st IX either verticalor oblique and rounded apically (Figs. 16, 17) or
clavate and bearing a rounded caudal point (Fig. 20), male st VIII
variable, caudal margin of female st VII variable, with or without a lobe and/or sinus ............................ 13
12 Projection on caudal margin of male movable process pronounced causing the margin below it to be deeply
concave, caudal margin of male fixed process straight, acetabular setae not arising on a bulge in the margin
(Fig. 6), upper lobe of apical arm of male st IX drawn into a slightly deflected, hooked apex (Fig 21), male
st VIII relatively short, its apical 2/3 drawn out into a tapering, spiculate point, its dorsal margin with a small
vermiform appendage (Fig. 36), caudal margin of female st VII usually with a sharp, pointed lobe subtended
by a V-shaped sinus (Fig. 51). (A parasite of Neotoma
species in northern Texas, New Mexico, eastern Colorado, southern Kansas, and Oklahoma.intermedius
Projection on caudal margin of male movable process and the concavity below it less pronounced, caudal
margin of fixed process slightly concave, acetabular setae arising on a slight bulge in the caudal margin (Fig.
7), dorsal projection of fixed process broad and rounded apically, upper lobe of distal arm of male st IX less
pointed and deflexed caudally, apical 3/4 of male st VIII spiculose, its dorsal margin bearing a small
vermiform appendage (Fig. 37), caudal margin of female st VII bearing a pronounced lobe that frequently
rounded and expanded apically (Fig. 52). (A parasite of Neotoma floridana in mountainous southern New
York, Pennsylvania, Maryland, and Virginia,
south to Alabama and Georgia.) .......................................................................................... pennsylvanicus
13 Movable process of male clasper with a smoothly rounded caudal margin, acetabular setae arising on a slight
bulge midway down the otherwise straight caudal margin of the fixed process (Fig. 1), upper lobe of distal
arm of male st IX gradually tapering to a rounded, slightly bent apex (Fig. 16), apical 2/3 of male st VIII
spiculose and with a vermiform appendage on its dorsal margin (Fig. 31), caudal margin of female st VII
variable, from broadly concave to bearing a short, tooth-like lobe (Fig. 46), always with a small, submarginal
incrassation. (On Neotoma species west of the Cascade
mountain chain in southern Washington to northern California.) ........................................... cascadensis
December, 2000
Journal of Vector Ecology
169
Caudal margin of male movable process with a distinct rounded angle but the margin below the spiniforms
is not concave, male st VIII and IX not as above, caudal margin of female st VII with
some type of lobe ...................................................................................................................................... 14
14 Movable process ~2X as long as wide, its cephalic margin slightly convex, caudal margin of fixed process
with a median bulge from which the acetabular setae arise (Fig. 5), upper lobe of distal arm of male st IX
drawn into a rounded, tapering, caudal point (Fig. 20), apical half of male st VIII spiculate basally, its apical
half forming an aspiculate vermiform appendage (Fig. 35), caudal margin of female st VII with a short, blunt
lobe (Fig. 50). (Southern desert areas of Arizona and
northwestern Mexico on Neotoma species.) ............................................................................... schisintus
Movable process ~2.5X as long as wide, its cephalic margin practically straight, caudal margin of fixed
process also straight, acetabular setae not arising on a bulge in the caudal margin (Fig. 2), upper lobe of distal
arm of male st IX with roughly parallel margins, its apex smoothly rounded (Fig. 17), apical 2/3 of male st
VIII spiculate, its dorsal margin bearing a small vermiform appendage (Fig. 32), caudal margin of female st
VII extremely variable but usually bearing a lobe of various proportions, subtended by a variously developed
sinus (Fig. 47). (Widespread in central North America west of roughly southwestern Texas, north to
Alberta and British Columbia east of the coastal ranges.
Primarily a parasite of Neotoma species.) ..........................................................................................agilis
Orchopeas agilis (Rothschild, 1905)
(Figs. 2, 17, 32, 47, 62) [190511]
Ceratophyllus agilis. Rothschild, 1905, Novit. Zool.
12: 167, pl. VII, Figs. 16-18. Canada, Alberta,
Banff, 51.10N 115 34W, from Neotoma
cinerea, 8.VIII.1899, G.F. Dippie leg. Lectotype
male, BMNH, designated by Smit and Wright,
1978b: 4.
Ceratophyllus sexdentatus nevadensis. Jordan, 1929,
Novit. Zool. 35: 30, pl. I, Fig. 4. USA, California,
Mono County, Pine City, from Mustela
arizonensis [=frenata], 21.XII.1922, A. B.
Howell leg. Lectotype male, BMNH, designated
by Smit and Wright, 1978b: 32. New Synonymy.
Remarks. Under the name Orchopeas
sexdentatus agilis, this taxon has been reported in the
literature from 22 species of hosts, including two
owls, but it is mainly a parasite of woodrats. However,
most published records shed little light on specific
host preferences, seasonal occurrence, sex ratios, and
other biological information. As interpreted here, the
taxon ranges from west southwestern Texas, north
through the western two-thirds of New Mexico and
northern Arizona, west to eastern California and north
into western Alberta and British Columbia. Holland
(1985) shows one record from the Yukon Territory.
As a result, the species has the broadest distribution of
any member of the Orchopeas sexdentatus species
group.
The only detailed analysis of this taxon is that of
Morlan (1955) albeit listed under the name of
Orchopeas sexdentatus. His study of the mammal
fleas of Sante Fe County, New Mexico, showed that the
species occurred on eight rodent and one leporid host
species. Only four species of Neotoma yielded
significant numbers of fleas. The 1571 woodrats broke
down as follows: N. micropus (579); N. albigula
(981); N. mexicana (9); and N. cinerea (2). In the same
order the percentages of animals infested was 78, 31,
11, and 100; the mean number of fleas per host was 6.6,
0.8, 0.1, and 10.0; and the mean number of fleas per
infested host was 8.4, 2.6, 1.0, and 10.0. These numbers
suggest that N. cinerea and N. micropus are the preferred
hosts and N. albigula and N. mexicana are less attractive.
This is in accord with Holland (1985) who states that
the dominant host in Canada is N. cinerea (Canada is
extralimital to N. micropus) and, as will be seen under
the account of Orchopeas neotomae, N. mexicana is
the preferred host of this species as well. According to
Morlan (1955) the nests of N. micropus were much
more heavily infested with fleas than those of N.
albigula. Evidently nests of the other two species of
woodrats were not examined for fleas. Morlan goes on
to say that seasonal abundance of adults on N. micropus
and N. albigula showed maxima from May to August
and minima from December to February, and the
majority of females examined each month of the year
were gravid. Male/female sex ratios showed females
comprising 58 to 61 percent of yearly collections, but
records from the two hosts were not consistent, and
males dominated some months from September
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Journal of Vector Ecology
through January. For further comment on this species
and its distribution see the account for Orchopeas
cascadensis.
A detailed examination of the type of O. s.
nevadensis provided no additional information that
would substantiate that it is distinct from this species,
and variation in the material examined assigned to this
taxon was well within the expected range for O. agilis.
Orchopeas caedens (Jordan, 1925)
(Figs. 9, 24, 39, 54, 69) [192521]
Ceratophyllus labiatus. Baker, 1904, Proc. U. S. Nat.
Mus. 27(1361): 402, pl. XIX, Figs. 6-9. USA,
Idaho, Latah County, Moscow, 46.44N 117.00
W from Lynx canadensis, date not given, J. M.
Aldrich leg. Holotype female, USNM No. 6909.
(See comment in the introduction concerning
the treatment of this name.)
Ceratophyllus caedens. Jordan, 1925, Novit. Zool.
32: 104, Figs. 16-17. Canada, Alberta, Banff,
51.10N 115. 34W, from Mustela [=Martes]
americana, 15.IX.1915, Mackay and G. F. Dippie
leg. Lectotype male, BMNH.
Ceratophyllus caedens durus. Jordan, 1929, Novit.
Zool. 35: 29, pl. I, Figs. 3a-g. Canada, British
Columbia, Okanagan, 50.02N 119.23W, from
Putorius arizonensis [=Mustela frenata],
16.XI.1917, J. A.Munro leg. Holotype female,
BMNH. New Synonymy.
Remarks. Since Wagner (1936) first indicated
that he questioned the distinctness of the two subspecies
of O. caedens, specialists have disagreed about their
validity. Certainly the strongest advocate for their
retention has been Holland (1949 and 1985) who argued
that the configuration of the female seventh sternum
was justification for their separation, even in the absence
of diagnostic characters in the male and the occurrence
of both “subspecies” on the same host from Banff,
Alberta, Atlin and Kinbasket Lake, British Columbia,
and Great Slave Lake in the Northwest Territories.
Indeed, many subsequent authors have simply referred
to Orchopeas caedens in their works without
deliberating the validity of the subspecies. The only
author to refer to intergrades between the two taxa was
Holland (1985) who cited them as occuring in a band
extending from the Northwest Territories and British
Columbia, east to Labrador. Having examined the types
of both taxa, and studied material from across North
America, it is my contention that the evidence is
December, 2000
insufficient to support the retention of both names.
Therefore O. c. durus is here again treated as a junior
synonym of the nominate taxon.
Although the preferred host is the red squirrel,
Tamiasciurus hudsonicus, host records in the literature
list 46 species, including six species of birds. There
are two records from Oklahoma published by Ellis
(1952 and 1955) that are almost certainly
misdeterminations, and the records from California by
Stewart (1940), Wood (1938) and Augustson (1943b)
are also questionable. There are also predator records,
especially from Martes americana and Mustela
species, all of which prey on red squirrels.
Orchopeas cascadensis Jordan, 1939
(Figs. 1, 16, 31, 46, 61) [193926]
Orchopeas sexdentatus cascadensis. Jordan, 1939,
Novit. Zool. 41: 317, Fig. 267. USA, Oregon,
Klamath County, Odell Lake, 43.32N 122.00W,
from Neotoma fuscipes, 31.VII.1936, C. A.
Hubbard leg. Holotype female, BMNH.
Remarks. Although the holotype female contains
a fair amount of blood and bears some surface detritus
on abdominal segments IV-VI, it is well mounted and
shows the caudal margin of st VII quite well, the single
most important diagnostic character for this taxon.
Jordan did evidently not draw this holotype, but it most
closely approximates his Figure 267e. Jordan did not
cite a type specimen for the taxon, but a slide bearing
Brit. Mus. 1923-615 is labeled “Type,” to which
someone has added “Holotype.” It currently resides in
the British Museum (Natural History) collection. Figure
46 is of this specimen. Variation in Figure 61 is based
on females from Lincoln County, Oregon.
On the basis of the male genitalia, it is difficult to
separate this species from Orchopeas agilis.
Comparison of the figures of the two species show the
caudal margin of the movable process of O. agilis to be
somewhat more angular than that of O. cascadensis,
but this character is somewhat variable in long series.
The acetabular setae arise above the middle of the fixed
process in O. cascadensis, frequently on a slight bulge
in the caudal margin, while the caudal margin of this
structure is relatively straight in O. agilis and the
acetabular setae arise below its middle.
In spite of the variation in the configuration of the
caudal margin of st. VII in females, as shown in Figure
61, when a lobe is present it is always pointed and there
is always at least a slight lateral incrassation in this
sternite, whether a lobe is present or not. An incrassation
is not present in females of O. agilis and the caudal lobe
December, 2000
Journal of Vector Ecology
is never so sharply pointed.
Holland did not report this taxon from Canada,
although he recorded a number of collections of O.
agilis from southern British Columbia, a few from
western Alberta, and one from the Yukon Territory,
which he noted as “possibly fallacious.” There are a
number of published and unpublished records of O.
cascadensis from California as far south as the San
Francisco Bay area and western Oregon. Examination
of limited collections from Washington State did not
yield this species, but Hubbard (1947) reported a
collection from Yacalt, Clark County, Washington.
Material examined by me came from central and
northern Washington and was referable to O. agilis.
Orchopeas dieteri (C. Fox, 1929)
(Figs. 14, 29, 44, 59, 75) [192928]
Ceratophyllus nepos dieteri. C. Fox, 1929, Entomol.
News 40(7): 218. USA, California, Los Angeles
County, sine loco, from Lynx ruffus [=rufus],
1926, collector not given. Holotype male,
allotype female, on separate slides, USNM No.
41820.
Remarks. The last detailed treatment of this
species was that of Eads et al. (1985) based on material
then in the collection of the Centers for Disease
Control at Fort Collins, Colorado, now in the
Smithsonian Institution, and the specimens mentioned
by them are the only ones that I have been able to
examine beyond the holotype male and allotype female.
All extant material was taken in the environs of Los
Angeles and the species is evidently normally a parasite
of the western gray squirrel, Sciurus griseus, but little
else is known about it.
Augustson (1943b) reported a collection of this
species from Sciurus griseus collected at Sunland, a
northern suburb of Los Angeles, but did not indicate
the number of specimens taken. As pointed out by Eads
et al. (1985) males of this species are distinct from O.
latens and O. nepos in the following characters. The
apex of the fixed process of the clasper is much more
acutely pointed; the acetabular bristles arise on the
caudal margin about halfway between the base and the
apex; and the four spiniforms on the movable process
arise on the ventral half of the caudal margin of the
movable process. As with many other members of this
genus, intraspecific variation in the females precludes
definitive determination in the absence of accompanying males.
171
Orchopeas fulleri Traub, 1950
(Figs. 12, 27, 42, 57, 75) [195022]
Orchopeas fulleri. Traub, 1950, Fieldiana Zool. Mem.
1: 60, pls. 36-37. México, Nuevo Leon, near
Villa Santiago, from “tree squirrel”, VI. 1940, H.
Hoogstraal and K. Knight leg. Holotype male,
allotype female, on separate slides, FMNH,
Chicago, IL, (published 28-February-1950).
Orchopeas howardi texensis. Eads, 1950, Ann.
Entomol. Soc. Am. 43(1): 47. USA, Texas,
Menard County, Menard, 30.56N 99.48W, from
Sciurus niger, 29.XII.1946, D. C. Thurman leg.
Holotype male, allotype female, USNM No.
104688, (published 24-April-1950). New
Synonymy.
Remarks. I have studied the primary types of O.
fulleri and O. howardi texensis and have concluded
that the latter is most likely a junior synonym of the
former. There is some variation in the size and the
contours of both the claspers and the ninth sternites,
but the limited amount of material available for study
suggests that only a single taxon is involved, that it is
primarily a parasite of diurnal tree squirrels, and that its
range extends from at least central Texas, south into
southern Mexico, and possibly, even further south.
Ayala-Barajas et al. (1988) list O. fulleri from
Chihuahua and Durango, from Sciurus apache (=S.
nayaritensis). They make no mention of O. howardi
texensis.
As may be seen from the figures of this taxon, it is
definitely a member of the howardi species group. In
the male it differs from other members of this group in
being considerably larger, although size itself is seldom
a valid character in insects in general. The movable
process is much more rounded caudally and it bears 5
submarginal spiniforms, although in the material
examined by me (7 males and 9 females) 1 male had 4
per side while 2 had 6 per side. The cephalic margin is
declivant, forming a distinct incrassate cusp that fits
into a fovea on the inner surface of the fixed process.
The gibbose dorsal lobe of the apical arms of the male
ninth sternite bears a distinct concavity in its dorsal
margin and a prominent convexity on its cephalodorsal
angle bearing 2 short, curved setae. The females
examined were remarkably consistent in the contours
of st VII, but my sample tended to show at least a
shallow depression in the caudal margin above the
ventral sinus. Thus they tended to closely resemble
females of O. howardi except for their generally larger
size.
172
Journal of Vector Ecology
Orchopeas howardi (Baker, 1895)
(Figs. 11, 26, 41, 56, 71) [189504]
Pulex wickhami. Baker, 1895: Can. Entomol. 27(4):
109, 111. USA, Iowa, Johnson County, Iowa
City, 41.39N 91.31W, from Sciuropterus
[=Glaucomys] volans, no date given, H. F.
Wickham leg. Lectotype female, BMNH,
designated by Hopkins, 1954: 198.
Pulex gilletti. Baker, 1895, Can. Entomol. 27(4): 109,
111. USA, Michigan, Ionia County, Portland,
42.52N 84.53W, from “red squirrel” [=
Tamiasciurus hudsonicus] no date given, C. P.
Gillette leg. Lectotype female, USNM No.
104632, designated by Adams and Lewis, 1995:
16.
Pulex howardii. Baker, 1895, Can. Entomol. 27(4):
109, 111. USA, Georgia, Rabun County,
Tallula Falls, 34.45N 83.22W, from “squirrel”
[=Sciurus niger], no date given, L. O. Howard
leg. Lectotype male, BMNH, designated by
Hopkins, 1954: 198.
Orchopeas howardi bolivari. Barrera, 1955, Revista
Soc. Mex. Ent. 1: 92, Figs. 7-10. Mexico,
Chiapas State, Ocosingo, 18.04N 92.15W, from
Glaucomys volans, no date given, M. del Toro
leg. Holotype male, allotype female, UNAM
(MZFC 251, 252). New Synonymy.
Remarks. The lectotype male of this species,
designated in Hopkins (1954) is poorly cleared, but the
diagnostic characters of the genitalia are clearly visible.
When material throughout its range is examined, it is
evident that there is considerable variation in the
contours of the fixed and movable process in the males,
and in the latter case even the number of spiniforms
along the caudal margin may vary from the usual four.
The form of the aedeagal crochet is also variable, but in
all specimens examined its apex was somewhat blunt.
Since Hopkins (1954) did not designate an
allolectotype, I chose a female from Little Rock,
Arkansas, as the source of the female diagnostic
characters. However, it should be emphasized that
spermathecae are not diagnostic in this genus and the
contours of the caudal margin of the seventh sternite
are notoriously variable as in most species in the genus.
Determination of females in the absence of
accompanying males is always questionable.
Having studied material from the Mexican states
December, 2000
of Durango and Coahuila, the holotype male and allotype
female of O. fulleri from Neuvo Leon, paratypes of O.
howardi bolivari from Chiapas, collections from
several localities in Texas, as well as other parts of
North America and the imported population in southern
England, I am convinced that material described as O.
howardi bolivari is well within the range of individual
variation observed in the nominate form. The slight
variations in the shape of the movable process of the
males and the contours of the seventh sternum of the
females show little if any geographic restriction.
Indeed, the population parasitizing Sciurus
carolinensis and Sciurus vulgaris in southern England
is more distinct than any of the material mentioned
above, and no separate name has ever been proposed for
it.
The description of O. h. bolivari, translated from
the original Spanish reads: “The form of the harpogones
(=movable processes) reaches, in the three paratypes,
approximately that of the typical subspecies; but there
is always an escotadura (a tooth-like projection) on the
dorsocephalic margin, though it may be very slight.
Figures 10A, B, C show examples of variation in the
configuration of the caudal margin of the seventh
sternite of the female.
“Orchopeas howardi bolivari nov. subsp. can
easily be recognized from the forma typica (Fig. 6) and
from O. h. texensis (Fig. 5) by the large size of the
movable process measured from the margin where the
small seta is inserted in the dorsocaudal margin (Fig. 9
sd) as far as the dorsalmost projection of the lobe of the
condyle: by the shape of the ventral border, which in O.
h. texensis is straight, in O. h. howardi is concaveconvex, and in O. h. bolivari nov. subsp. is slightly
concave; and furthermore by the presence of the
dorsocephalic projection in the other two known
subspecies.”
The differences in contours of the movable
processes in the two named taxa are insignificant and
there is no noticeable projection of the dorsocephalic
apex in the figured paratype.
There remains the puzzling record from Venezuela
listed in Tipton and Machado-Allison (1972). The
provenance of these two specimens is questionable. I
have been able to examine a single male in the collection
of the Bean Museum, Brigham Young University, Provo,
Utah, said to have been collected in Venezuela. It is
completely without data and is over cleared. This
specimen is closer to O. latens than to O. howardi, so
it may not be one of the two specimens listed by Tipton
and Machado-Allison. Extensive collecting in Panamá
failed to reveal O. howardi, or any other member of the
genus in that country and there are evidently no published
December, 2000
Journal of Vector Ecology
records of any member of the genus south of México.
However, I am informed that there is 1 male and 6
females in the collection of the Carnegie Museum of
Natural History, Pittsburgh, Pennsylvania, bearing the
following data: Guatemala, Jalapa, 4 mi. NE Volcan de
Jumay Aserrodoro, San Lorenzo, 6000” ex Sciurus
aureogaster, 11.II.1952, Luis de la Torre leg, CNHM,
R. Traub det. Ayala-Barajas et al. (1988) list specimens
from the Mexican states of Chiapas, Guerrero, and
México. Most of these were collected on flying
squirrels.
There are probably more published records for
this species than for any other member of the genus.
Although the literature reports it from at least 53 host
species, including eight species of birds, it is evidently
a true parasite of Sciurus carolinensis and S. niger. It
is a common flea east of the Rocky Mountains and
Benton (1980) reported it from almost every state east
of the Mississippi River. It is suggested that the
California records of Wood (1938) and Augustson
(1943b), if not misdeterminations, were from fox
squirrels imported from the eastern states. This may
also be the case with the records of Campos (1971) and
Hudson et al. (1971) from Colorado. The occurrence
of this species in Utah as reported by Stanford (1944)
was contested by Stark (1959) and the record from
Montana in Dunn and Parker (1923) was changed to
Orchopeas caedens by Jellison et al. (1943).
On more than one occasion this species has been
found on humans, though usually singly, and we know of
no published records of its feeding. However, in a few
instances the species has become an important pest in
commercial mink ranches in central Iowa and elsewhere.
Benton (1980) reported a similar infestation at a mink
farm in Virginia. Infestations develop where the mink
cages are unsheltered and in close proximity to trees,
particularly when sanitation practices in the
confinement area are lax.
Orchopeas illinoiensis Lewis, 1998
(Figs. 4, 19, 34, 49, 64) [199801]
Orchopeas illinoiensis Lewis, 1998, J. Med. Entomol.
35(4): 399-403. USA, Illinois, Union County,
Larue-Pine Hills area, ~37.35N 89.25W,
fromNeotoma floridana, IV.1996, Anne-Marie
Monty leg. Holotype male, allotype female.
Deposited in the Field Museum of Natural
History, (FMNH), Chicago, IL.
Remarks. Representatives of this species were
listed by Palmer and Wingo (1972) under O.
sexdentatus. The specimens were collected by me in
173
1953, during service in the United States Army at Fort
Leonard Wood, Missouri (Pulaski County). Most
specimens have since been lost, but a few of them still
reside in the Enns Collection at the University of
Missouri, and I believe, in the Traub Collection, now at
the Carnegie Museum in Pittsburgh, Pennsylvania.
These were not included in the type series because they
were so poorly cleared and mounted.
Mammalogists now recognize two separate eastern
species of Neotoma. There is little doubt that O.
illinoiensis is a parasite of N. floridana, the range of
which extends to eastern Colorado and Texas, west of
the Mississippi River. Where available, this host prefers
bluffs and caves, or at least a rocky habitat in which to
build its rather bulky lodges. Fleas are mainly confined
to the fibrous nesting material and may occur in large
numbers in old, well established lodges.
Orchopeas intermedius Hubbard, 1943
(Figs. 6, 21, 36, 51, 66) [194306]
Orchopeas sexdentatus intermedius. Hubbard, 1943,
Pacific Univ. Bull. 39(10): 2 plus an unnumbered
figure. USA, Kansas, Meade County (sine loco),
from wood rat [=Neotoma micropus], VII.1940,
J. E. Hill and P. E. Crowe leg. Holotype male,
allotype female, on same slide, USNM No.
57087.
Remarks. To date this species has been reported
in the literature only from collections from Kansas,
Oklahoma, and Colorado, where it was collecetd on
woodrats, specifically Neotoma albigula and N.
micropus. I have been able to examine 40 males and 81
females belonging to this taxon. In addition to 1male
and 2 females from Kansas, 16 males and 30 females
from Oklahoma, and 5 males and 13 females from
Colorado, I have also examined three pairs from New
Mexico and 15 males, 33 females from Texas. All but
occasional strays were from species of Neotoma.
Orchopeas latens (Jordan, 1925)
(Figs. 13, 28, 43, 58, 73) [192522]
Ceratophyllus latens. Jordan, 1925, Novit. Zool. 32:
105, Figs. 19-20. USA, California, Santa Cruz
County, sine loco, from gray squirrel [=Sciurus
griseus], VIII.1900, E. M. Erhorn leg. Lectotype
male, BMNH, designated by Smit and Wright,
1978b: 27.
Remarks. This is another poorly known species
that is evidently a parasite of the tree squirrels, Sciurus
174
Journal of Vector Ecology
griseus and Tamiasciurus douglasii, although
Augustson (1943b) also reported it from Citellus
(=Spermophilus) beecheyi from San Diego County,
California. After a thorough review of the literature,
I am convinced that the record from Oregon listed
in Traub et al. (1983) and repeated in Lewis et al.
(1988) is either a misidentification or a case of confused collection data, since all the other records
known to me are from the southern half of California.
I have been able to examine an additional 5 males
and 12 females of this species from Nevada, Riverside,
San Francisco, San Mateo, and Santa Cruz counties in
California.
Orchopeas leucopus (Baker, 1904)
(Figs. 10, 25, 40, 55, 70) [190420]
Ceratophyllus leucopus. Baker, 1904, Proc. U. S.
Nat. Mus. 27(1361): 401. USA, New York,
Madison County, Peterboro, 43.00N 75.40W,
from Peromyscus leucopus, VII.1900, G. S.
Miller leg. Holotype female, USNM No. 6908.
Ceratophyllus aeger. Rothschild, 1905, Novit. Zool.
12: 167, pl. VI, Figs. 5, 7, 9. Canada, Alberta,
Red Deer, 52.15N 113.48W, from Peromyscus
arcticus [=maniculatus], 22.V.1901, G. F.
Dippie leg. Lectotype male, BMNH, designated
by Smit and Wright, 1978b: 3. Synonymized by
Jordan, 1928: 179.
Remarks. This species has the most extensive
range of any member of the genus, having been reported
from almost all of the contiguous 48 states and all of
the Canadian provinces. Holland (1985) reported a
single Alaskan record from Anchorage. Although I
have located 76 host species in the literature, including
seven bird species, this flea is mainly a parasite of
Peromyscus leucopus and P. maniculatus, but it may
frequent other species of Peromyscus and Neotoma,
and has often been taken on Reithrodontomys megalotis
and Onychomys leucogaster. Again, precise
information about the species is lacking for most of its
range. However, the species is discussed by Morlan
(1955) in his study of the mammal fleas of Santa Fe
County, New Mexico. He reported O. leucopus from
12 different rodent species, but only six of these were
significant hosts. These were Reithrodontomys
megalotis (130); Peromyscus maniculatus (445); P.
leucopus (285); P. truei (662); Onychomys
leucogaster (405); and Neotoma micropus (579). In
the same order, the percentage of animals infested was
21, 17, 34, 39, 1, and <1; the mean number of fleas per
December, 2000
animal examined was 0.5, 0.3, 0.8, 0.9, <0.1, and <0.1;
and the mean number of fleas per infested animal was
2.2, 1.8, 2.2, 2.2, 1.2, and 5.0. Adult fleas were most
abundant from April through August, and least abundant
from November through February. Adults were
collected from P. truei and P. maniculatus during
all months of the year but were absent from P.
leucopus during December and January. Females
predominated over males on all three hosts the majority
of the time, although ratios varied from month-tomonth. Several pools of this species were infected
with plague bacilli according to Holdenried and Morlan
(1955). The egg of this species was described in Linley
et al. (1994).
I have been able to examine 180 males and 299
females from 20 states as well as Ontario and British
Columbia, Canada, and Durango state, Mexico.
Orchopeas neotomae Augustson, 1943
(Figs. 8, 23, 38, 53, 68) [194309]
Orchopeas sexdentatus neotomae. Augustson, 1943,
Bull. So. California Acad. Sci. 42(1): 49, pl. 5,
Figs. 1-2. USA, Arizona, Coconino County,
south entrance to Grand Canyon National Park,
~ 36.03N 112.04W,
from Neotoma
lepida, 5.VI.1942, R.L. Rutherford leg.
Holotype male, allotype female, LACM, Los
Angeles.
Remarks. There is little mention of this
southwestern species in the literature and most
published records are from Arizona and New Mexico.
The northernmost collection is a single female from
Valley County, Idaho, reported by Baird and Saunders
(1992). Although females of this taxon are distinctive,
I suspect this is a misdetermination or a mistranscription
of the collection data. Campos et al (1985) reported
nine specimens from Larimer County, Colorado, from
Neotoma mexicana. Tipton and Allred (1952) also
reported a single female from Grafton, Washington
County, Utah, which is slightly north of the type locality
for the species. There are three females and one male
in the Lewis Collection from Durago and Zacatecas,
México, and Ayala-Barajas et al. (1988) report
collections from the states of México, D. F. and
Morelos.
This species has been reported in the literature
from 10 different host species. Five of these belong to
the genus Neotoma and are evidently the preferred
hosts, with Neotoma mexicana being the most
frequently infested. Two more hosts are predators,
another is a tree squirrel, and the remaining two are
December, 2000
Journal of Vector Ecology
species of Peromyscus, frequent associates of woodrat
nests. All but the Neotoma records are viewed as
accidental associations.
This and the other members of the Orchopeas
sexdentatus complex are the only taxa that have
woodrats as their preferred hosts, and unlike the latter
taxa, O. neotomae is seldom collected in large numbers.
Most reports refer to single specimens, i.e., Tipton and
Allred (1952), Traub and Hoff (1951), Eads et al.
(1979), and Baird and Saunders (1992). Kartman
(1960a) reported one O. neotomae, but no O.
sexdentatus. However, Kartman (1960b) reported no
O. neotomae but 137 O. sexdentatus from his studies
in New Mexico. Hubbard (1943) listed 40 specimens
of this species, but included no indication of the numbers
of hosts involved. Holdenried and Morlan (1955)
reported only nine specimens of the 757 discussed in
their study. Haas et al. (1973) reported 67 males and
111 females of this species from central New Mexico,
mostly from N. mexicanus. By far the most informative
study of this species is that of Morlan (1955) in his
report on the mammal fleas of Santa Fe County, New
Mexico. During this study the author examined 1,571
specimens of four species of Neotoma: N. micropus
(579); N. albigula (981); N. mexicana (9); and N.
cinerea (2). In the same order he reported the percentage
of animals infested with O. neotomae as <1, 1, 44, and
50; the mean number of fleas per host examined as
<0.1, <0.1, 2.0, and 0.5; and the mean number of fleas
per infested host as 1.7, 2.1, 5.0, and 1.0. This is a clear
indication that N. mexicana is the preferred host of this
species. Morlan also indicated that small collections
of this flea were made from May through November
and that the sex ratio was 5.2 males to 4.8 females. A
number of pools of this species were positive for the
plague bacillus according to Holdenried and Morlan
(1955).
During examination of the 32 males and 31 females
available to me the following variations in genitalic
anatomy were noted. While the fixed process of the
male clasper is always broadly rounded apically and
extends almost to the apex of the movable process, the
position and proximity of the acetabular setae varies.
The arrangement shown in Figure 8 represents the
norm, but there may be a somewhat more pronounced
acetabular projection and the bases of the setae may
be closer together. With respect to the movable
process, none of the males showed fewer than four
spiniform setae along the caudal margin. There were
cases where one or more had been lost, but alveoli
were there to attest to their original presence. The
spacing of these spiniforms showed slight variations.
Typically the three dorsalmost spines are equidis-
175
tant from each other. However the lowermost third
and fourth arise much closer together. In a few males
the second spiniform is closer to numbers 3 and 4
than to number 1, as is the case in a topotypical male
that was examined. There was also some variation in the
depth of the concavity in the ventral half of the caudal
margin. The form of the ninth sternite was remarkably
constant in the series, and only minor variations in the
chaetotaxy and the depth of the sinus in the caudal
margin of the apical lobe were evident. In females, as
can be seen from Figure 53 of a topotype and Figure 68,
contours of the caudal margin of the seventh sternum
are fairly consistent, with little variation in the size and
shape of both the dorsal lobe and subtending sinus.
Orchopes nepos (Rothschild, 1905)
(Figs. 15, 30, 45, 60, 74) [190512]
Ceratophyllus nepos. Rothschild, 1905, Novit. Zool.
12: 168, pl. VII, Figs. 13-14. Canada, British
Columbia, Chilliwack, 49.06N 124.04W, from
Spilogale latifrons [=putorius], 1.XI.1899. A.
Brooks leg. Lectotype male, BMNH, designated
by Smit and Wright, 1978b: 32.
Remarks. This species is reported in the literature
from 26 mammalian host species. It is a specific
parasite of Tamiasciurus douglasi, although there are
a significant number of records from Sciurus griseus
as well. It is restricted to California, Oregon,
Washington, and southern British Columbia, but there
are at least two records from Nevada. The record by
Brown and Yeager (1945) of this species from Illinois
on Sciurus carolinensis and S. niger certainly refers
to some other species, probably O. howardi.
Orchopeas pennsylvanicus (Jordan, 1928)
(Figs. 7, 22, 37, 52, 67) [192814]
Ceratophyllus sexdentatus pennsylvanicus. Jordan,
1928, Novit. Zool. 34: 184, Figs. 8-9. USA,
Pennsylvania, Westmoreland County, Ligonier,
Rolling Rock Club, 40.14N 79.14W, from
Neotoma pennsylvanica [=magister], 29.VI.
1927, K. Jordan leg Holotype male, BMNH.
Remarks. This is the eastern congener of the O.
sexdentatus complex and it is a specific parasite of the
eastern woodrat, Neotoma magister. I have been unable
to examine much of the material reported in the
literature from Alabama, Illinois, Indiana, Kansas, and
Kentucky, but I suspect that these collections will turn
out to be O. illinoiensis rather than O. pennsylvanicus.
176
Journal of Vector Ecology
December, 2000
Orchopeas schisintus (Jordan 1929)
(Figs. 5, 20, 35, 50, 65) [192904]
Orchopeas sexdentatus (Baker, 1904)
(Figs. 3. 18, 33, 48, 63) [190422]
Ceratophyllus sexdentatus schisintus. Jordan, 1929,
Novit. Zool. 35: 31, pl. I, Fig. 5. USA, Arizona,
Cochise County, Paradise, 31.51N 109.13W,
from Neotoma sp., 1913, O. C. Dufner leg.
Holotype female, BMNH.
Ceratophyllus sexdentatus. Baker, 1904, Proc. U. S.
Nat. Mus. 27: 403, pl. XXVI, Figs. 8-14. USA,
California, Santa Cruz County, Boulder Creek,
37.07N 122.09W, from Neotoma sp., no date,
E. Ehrhorn leg. Lectotype male, allolectotype
female, on separate slides, USNM No. 6910.
Orchopeas sexdentatus firemani. Hubbard, 1955,
Entomol. News 66: 138. México, Sinaloa State,
56 km N Los Mochis, on border of Sinaloa and
Sonora, ~26.26N 109.02W, from Teanopus
[=Neotoma] phenax, 23.III.1954, C. A. Hubbard
leg. Holotype male, allotype female, on one
slide, USNM No. 104622. New Synonymy.
Orchopeas reevesi. Vargas, 1960, Rev. Inst. Salub.
Enferm. Trop. Med. 20: 27-35, Figs. 1-6.
México, Sonora, 22 km S. Hermosilla, 29.15N
110.59W, from Neotoma albigula, 31.III.1960,
K. Murray and O. Soave leg. Holotype male,
allotype female, I. H. D. T., México D. F.
Synonymized by Smit and Wright, 1978a: 38.
Remarks. Since the holotype of this species is a
female, its true identity remains conjectural at this
point. Assuming that the males and females of the 30+
specimens available to me were properly associated,
the apex of the distal arm of the male st IX and the
configuration of the male st VIII appear to be the two
most diagnostic characters. Except for 4 males and 2
females in my collection from Sonora, Mexico, all
material available to me is from Yuma and Maricopa
counties in Arizona. Females illustrated in Figure 65
are from McDowell Mountain near Phoenix (~33.40N
111.45W) from Neotoma albigula. The Yuma County
collections are from the collection of the Centers for
Disease Control, now in the Smithsonian Institution.
Most of these have no precise locality data, having been
assigned collection numbers that presumably relate to
field logs, the presence of which is unknown to me.
Unfortunately the tip of the distal arm of the male
sternite IX of the holotype of O. s. firemani is folded
over and its exact contours cannot be distinguished
with certainty. Accordingly, the pair of paratypes in the
British Museum (Natural History) was borrowed, but
both sides of this structure also show apical folding,
although one side is somewhat less distorted than the
other. From examining these two males I am confident
that the dashed line in Figure 20 closely approximates
the true form of the apex of sternum IX.
Remarks. Sorting out records for this species is
difficult since so many published records include
material only identified to species. Indeed, the literature
suggests a distribution extending as far east as Virginia.
However, O. sexdentatus, as treated here is confined
to a small area in California west of the mountains,
extending north from San Diego to the San Francisco
Bay area. It has been reported from a number of small
mammals, but its preferred hosts are species of
Neotoma. The males are easily separated from the
other members of this species group by the squared
apex of the apical arms of the ninth sternite as shown in
Figure 18. However, as one progresses south from the
San Francisco area, the dorsocaudal apex of the male st
IX becomes more pointed and the caudal margin below
it more concave.
Acknowledgments
A number of colleagues and institutions have been
of assistance during this study. I am particularly
beholden to Ms. Nancy Adams of the Smithsonian
Institution, Washington, D.C.; Ms. Theresa Howard of
the Natural History Museum, London, England; Dr.
Alfred F. Newton, Jr., of the Field Museum of Natural
History; and Dr. Brian Brown of the Natural History
Museum of Los Angeles County, Los Angeles,
California, for the loan of primary types. Other
colleagues who assisted with loans of material, etc.,
include the late A. M. Barnes, Communicable Diseases
Center, Fort Collins, Colorado; S. Bennett, Garden
Grove, California; L. Durden, Statesboro, Georgia; R.
P. Eckerlin, Annandale, Virginia; H. J. Egoscue,
Grantsville, Utah; G. E. Haas, Boulder City, Nevada; M.
W. Hastriter, Provo, Utah; J. D. Lang, San Diego,
California; C. Senger, Bellingham, Washington; and N.
Wilson, Cedar Falls, Iowa. I also wish to thank the two
anonymous referees who reviewed this manuscript.
Journal paper No. J-18751 of the Iowa Agricultural and
Home Economics Experiment Station, Ames, IA,
Project No 3100, and supported by Hatch Act and State
of Iowa funds.
Journal of Vector Ecology
Figures 1-8. Movable and fixed processes of the male clasper. 1. Nontype from Oregon. 2. Lectotype. 3. Lectotype. 4. Holotype. 5. Nontype from Arizona.
6. Holotype. 7. Holotype. 8. Nontype from New Mexico.
December, 2000
177
Journal of Vector Ecology
Figures 9-15. Movable and fixed processes of the male clasper. 9. Lectotype. 10. Nontype from New York. 11. Lectotype. 12. Holotype. 13. Lectotype.
14. Holotype. 15. Nontype from British Columbia.
178
December, 2000
Journal of Vector Ecology
Figures 16-23. Apex of distal arm of male sternite IX. 16. Nontype from Oregon. 17. Lectotype. 18. Lectotype. 19. Holotype. 20. Nontype from Arizona.
21. Holotype. 22. Holotype. 23. Nontype from Mexico.
December, 2000
179
Journal of Vector Ecology
Figures 24-30. Apex of distal arm of male sternite IX. 24. Lectotype. 25. Nontype from New York. 26. Lectotype. 27. Holotype. 28. Lectotype. 29.
Holotype. 30. Nontype from British Columbia.
180
December, 2000
Journal of Vector Ecology
Figures 31-38. Sternite VIII of males. 31. Nontype from Oregon. 32. Lectotype. 33. Lectotype. 34. Holotype. 35. Nontype from Arizona. 36. Holotype.
37. Holotype. 38. Nontype from Mexico.
December, 2000
181
Journal of Vector Ecology
Figures 39-45. Sternite VIII of males. 39. Lectotype. 40. Nontype from New York. 41. Lectotype. 42. Holotype. 43. Lectotype. 44. Holotype. 45.
Nontype from British Columbia.
182
December, 2000
Journal of Vector Ecology
Figures 46-53. Sternite VII and spermatheca of females. 46. Holotype. 47. Nontype from Utah. 48. Allolectotype. 49. Allotytpe. 50. Holotype. 51. Nontype
from Texas. 52. Nontype from Virginia. 53. Topotype from Arizona.
December, 2000
183
Journal of Vector Ecology
Figures 54-60. Sternite VII and spermatheca of females. 54. Nontype from Pennsylvania. 55. Nontype from New York. 56. Nontype from Arkansas. 57.
Allotype. 58. Nontype from California. 59. Nontype from California. 60. Nontype from British Columbia.
184
December, 2000
December, 2000
Journal of Vector Ecology
Figures 61-65. Variation in the caudal margin of female sternite VII. Not all drawn to the same scale.
185
186
Journal of Vector Ecology
December, 2000
Figures 66-70. Variation in the caudal margin of female sternite VII. Not all drawn to the same scale.
December, 2000
Journal of Vector Ecology
Figures 71-75. Variation in the caudal margin of female sternite VII. Not all drawn to the same scale.
187
188
Journal of Vector Ecology
REFERENCES CITED
Adams, N. E and R. E. Lewis. 1995. An annotated
catalogue of primary types of Siphonaptera in the
National Museum of Natural History, Smithsonian
Institution. Smithson. Contrib. Zool. 660: 1-86.
Augustson, G. F. 1943a. A new subspecies of
Orchopeas sexdentatus (Baker) (Siphonaptera:
Dolichopsyllidae). Bull. S. Calif. Acad. Sci. 42:
49-51.
Augustson, G. F. 1943b. Preliminary records and
discussion of some species of Siphonaptera from
the Pacific Southwest. Bull. S. Calif. Acad. Sci.
42: 69-89.
Ayala-Barajas, R., J. C. Morales-Muciño, N. Wilson, J.
E. Llorente-Bousquets and H. E. Ponce-Ulloa.
1988. Catalogo de las pulgas (Insecta:
Siphonaptera) en el Museo de Zoologia, Facultad
de Ciencas, Universidad Nacional Autonoma
México. Coleccion Alfredo Barrera. Ser. Catal.
Mus. Zool. “Alfonso L. Herrera” 1: 1-102.
Baird, C. R. and R. C. Saunders. 1992. Annotated
checklist of the fleas of Idaho (Siphonaptera).
Idaho Agric. Exp. Sta. Res. Bull. 148. 34 pp.
Baker, C. F. 1895. Preliminary studies in Siphonaptera.I-VII. Can. Entomol. 27: 19-22; 63-66; 108-111;
130-132; 162-165; 186-191; 221-222.
Benton, A. H. 1980. An atlas of the fleas of the eastern
United States. Marginal Media. Fredonia. 157 pp.
Brown, L. G. and L. E. Yeager. 1945. Fox squirrels and
gray squirrels in Illinois. Bull. Illinois Nat. Hist.
Surv. 23: 449-536.
Campos, E. G. 1971. The Siphonaptera of Colorado.
M.S. Thesis. Colorado State University. Fort
Collins. 274 pp.
Campos, E. G., G. O. Maupin, A. M. Barnes, and R. B.
Eads. 1985. Seasonal occurrence of fleas (Siphonaptera) on rodents in a foothills habitat in Larimer
County, Colorado, USA. J. Med. Entomol. 22:
266-270.
Dunn, L. H. and R. R. Parker. 1923. Fleas found on wild
animals in the Bitterroot Valley, Montana. Publ.
Hlth. Rep. Wash. 38: 2763-2775.
Eads, R. B., E. G. Campos, and A. M. Barnes. 1979.
New records for several flea (Siphonaptera) species
in the United States, with observations on species
parasitizing carnivores in the Rocky Mountain
region. Proc. Entomol. Soc. Wash. 81: 38-42.
Eads, R. B., B. C. Nelson, G. O. Maupin, and A. M.
Barnes. 1985. Orchopeas fleas (Siphonaptera:
Ceratophyllidae) of the Western Gray squirrel,
Sciurus griseus. J. Med. Entomol. 22: 630-636.
Ellis, L. L., Jr. 1952. A tentative key to the mammalian
December, 2000
ectoparasites of the Wichita Mountains Wildlife
Refuge. Proc. Oklahoma Acad. Sci. 33: 111-115.
Ellis, L. L., Jr. 1955. A survey of the ectoparasites of
certain mammals in Oklahoma. Ecology 36: 1218.
Fox, I. 1940. Fleas of eastern United States. Iowa State
College Press. 191 pp.
Haas, G. E., R. P. Martin, M. Swickard, and B. E. Miller.
1973. Siphonaptera-mammal relationships in
northcentral New Mexico. J. Med. Entomol. 10:
281-289.
Holdenried, R. and H. B. Morlan. 1955. Plagueinfected fleas from northern New Mexico wild
rodents. J. Infect. Dis. 96: 133-137.
Holland, G. P. 1949. The Siphonaptera of Canada. Can.
Dept. Agric. Tech. Bull. 70: 1-306.
Holland, G. P. 1985. The fleas of Canada, Alaska and
Greenland (Siphonaptera). Mem. Entomol. Soc.
Canada 130: 1-631.
Hopkins, G. H. E. 1954. Nomenclatorial notes on the
flea of the grey squirrel. Entomologist 87: 197198.
Hubbard, C. A. 1943. The fleas of California (with
check lists of the fleas of Oregon, Washington,
British Columbia, Alaska, Idaho, Nevada, Arizona).
Pacif. Univ. Bull. 39(8): 1-12.
Hubbard, C. A. 1947. Fleas of western North America.
Iowa State Coll. Press, Ames, 533 pp.
Hudson, B. W., M. I. Goldenberg, J. D. McCluskie, H.
E. Larson, C. D. McGuire, A. M. Barnes, and J. D.
Poland. 1971. Serological and bacteriological
investigations of an outbreak of plague in an urban
tree squirrel population. Am. J. Trop. Med. Hyg.
20: 255-263.
Jellison, W. L., G. M. Kohls, and H. B. Mills. 1943.
Siphonaptera: Species and host list of Montana
fleas. Misc. Publs. Montana St. Bd. Entomol. 2: 122.
Jordan, K. 1925. New Siphonaptera. Novit. Zool. 32:
96-112.
Jordan, K. 1928. Siphonaptera collected during a visit
to the eastern United States of North America in
1927. Novit. Zool. 34: 178-188.
Jordan, K. 1929. Notes on North American fleas.
Novit. Zool. 35: 28-39.
Jordan, K. 1933. A survey of the classification of the
American species of Ceratophyllus s. lat. Novit.
Zool. 39: 70-79.
Kartman, L. 1960a. The role of rabbits in sylvatic
plague epidemiology, with special attention to
human cases in New Mexico and use of the
fluorescent antibody technique for detection of
Pasteurella pestis in field specimens. Zoonoses
December, 2000
Journal of Vector Ecology
Res. 1: 1-27.
Kartman, L. 1960b. The role of rabbits in sylvatic
plague epidemiology, with special attention to
human cases in New Mexico and use of the
fluorescent antibody technique for detection of
Pasteurella pestis in field specimens. Zoonoses
Res. 1: 169-195.
Lewis, R. E. and J. H. Lewis. 1989. A catalogue of
invalid or questionable genus-group and speciesgroup names in the Siphonaptera (Insecta). Theses
Zoologicae. 11. Koeltz Scientific Books. Koenigstein, Germany. 263 pp.
Lewis, R. E., J. H. Lewis, and C. Maser. 1988. The fleas
of the Pacific Northwest. Oregon State Univ.
Press, Corvallis. 296 pp.
Linley, J. R., A. H. Benton, and J. F. Day. 1994.
Ultrastructure of the eggs of seven flea species
(Siphonaptera). J. Med. Entomol. 31: 813-827.
Morlan, H. B. 1955. Mammal fleas of Santa Fe County,
New Mexico. Texas Rep. Biol. Med. 13: 93-125.
Palmer, D. B., Jr. and C. W. Wingo. 1972. Siphonaptera
occurring on Missouri mammals. Trans. Missouri
Acad. Sci. 6: 43-55.
Smit, F. G. A. M. and A. M. Wright. 1978a. A list of
code numbers of speciesand subspecies of Siphonaptera. Department of Entomology, British
Museum (Natural History). London, 49 pp.
(mimeographed)
Smit, F. G. A. M. and A. M. Wright. 1978b. A catalogue
of primary type-specimens of Sipho-naptera in the
British Museum (Nat. Hist.) Department of
Entomology, British Museum (Natural History).
London. 71 pp. (mimeographed)
Stanford, J. S. 1944. More Utah Siphonaptera. Proc.
Utah Acad. Sci., Art and Letters 20: 173-178.
189
Stark, H. E. 1959. The Siphonaptera of Utah, their
taxonomy, distribution, host relations and medical
importance. U. S. Publ. Hlth. Serv., Commun. Dis.
Center, Atlanta, Georgia. 239 pp.
Stewart, M. A. 1940. Estado que guarda el Estudio
Hecho sobre la transmission de la Peste Silvestre
en American del North. Sob. Rev. Med. 20: 410415.
Tipton, V. J. and D. M. Allred. 1952. New distribution
records of Utah Siphonaptera with the description
of a new species of Megarthroglossus Jordan and
Rothschild 1915. Great Basin Nat. 11:105-114.
Tipton, V. J. and C. Machado-Allison. 1972. Fleas of
Venezuela. Brigham Young Univ. Sci. Bull., Biol.
Ser. 17(6): 1-115.
Traub, R. and C. C. Hoff. 1951. Records and descriptions
of fleas from New Mexico (Siphonaptera). Am.
Mus. Novit. 1530: 1-23.
Traub, R., M. Rothschild, and J. F. Haddow. 1983. The
Rothschild collection of fleas. The family
Ceratophyllidae: key to the genera and host
relationships, with notes on their evolution,
zoogeography and medical importance. Cambridge
University Press, U.K. Distributed by Academic
Press, Inc. (London). 228 pp.
Wagner, J. 1930. Analytical tables for the identification
of the species of Aphaniptera living on Muridae.
Parasit. Sb., Leningr. 1: 97-192. [inRussian]
Wagner, J. 1935. 23. Ordernung: Flöhe, Aphaniptera
(Siphonaptera, Suctoria) Die Tierwelt Mitteleuropas 6: lfg. 2 abt. 17: 1-24.
Wood, F. D. 1938. Trypanosoma neotoma sp. n. in the
dusky-footed wood rat and the wood rat flea. Univ.
California Publ. Zool. 41: 133-144.