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v32n2p111
FLEAS
COLLECTED
DURING A PLAGUE
SURVEY IN VENEZUELA1.2
BY ROBERT TRAUB, Major, MSC, and PHYLLIS
From the
c
Y
T. JOHNSON, Parasitologist
Department of Entomology,
Army Medical Service Graduate School,
Washington, D. C.
At the invitation of the Government of Venezuela and of the Pan
Ameritan Sanitary Bureau, a Mission from the Naval Medical School
at Bethesda, Maryland, made a survey in 1950 of sylvatic plague in
Venezuela. The epidemiological team consisted of Commander J. M.
Amberson, MC, USN, Dr. Ernst Schwarz, Zoologist, and Mrs. H. K.
Schwarz, Laboratory Assistant. The present paper presents data on the
fleas collected by members of the Mission and includes the description
and illustrations of a new species, as well as those of related and/or
potentially medically important forms.
It is of undoubted epidemiological significance that none of the fleas
collected by the Mission in these endemic areas were Xenopsylla, the
genus involved in classical plague transmission. Only one specimen of a
cosmopolitan species was found (Ctenoce$&%!es felis felis); al1 the others
were indigenous fleas.
Unless otherwise specified, fleas in the following lists were collected
at Aragua, Campamento Rafael Rangel, by E. Schwarz, H. K. Schwarz
and J. M. Amberson, July and August 1950. The team’s other collecting
site was at Campamento Chacin, River Tuy, Miranda. The first section
of this paper consists of a systematic list of fleas collected by the Mission,
and in each instance cites the host involved; descriptions and notes are
at times included. The second section lists the host mammals and their
respective fleas. In the first list a fraction in parentheses follows the
host name, i. e., under Polygenis b. bah& (Wagner, 1801), will be found
Akodon urichi urichi Allen and Chapman (8/10). Th% signifies that out
of 10 Akodon parasitized with fleas, 8 were hosts to P. b. bohìsi.
1. SYSTEMATIC
LIST OF FLEAS, WITH DESCRIPTIONS
AND NOTES
Family HYSTFXCHOPSYLLIDAE
Subfamily CTENOPHTHALMINAE
TritopsylIa
Ex Didelphis
Ex Sigmodon
intermedia
m. mursupialis
hispidus
hirsutus
oxyura (Jordan, 1926)
Allen, 7 d, 7 Q
Burmeister (1/42), 1 B
1 PubIished under the auspices of the Surgeon General, Department of the
Army, who does not necessarily assume responsibility for ‘cheprofessional opinions
expressed by the authors.
* Received for publication August, 1951.
111
112
BULLETIN
OF THE PAN AMEBICAN WNITARY
BUEEAU
Family RHOPALOPSYLLIDAE
Polygenis ambersoni sp. nov.
Types.-Holotype
male and allotype female ex Sigmodon hisp$dus
hirsutus Burm. Venezuela: Aragua, Campamento Rafael, Rangel, elev.
1260 meters, coll. J. M. Amberson, E. Schwarz and H. K. Schwarz,
7 August 1950; deposited in United States National Museum. One
paratype male with same data, two paratype males ibid., but 21 September; in collection of senior author.
Diagnosis.-Near
Polygenis peronis (J. & R., 1923) and P. acodontis
(J. & R., 1923), but readily separable in that the male ninth sternum
bears very long subapical bristles, longer than the dista1 arm itself
(excluding the heel) (fig. 4), instead of these bristles being merely about
half the length of the dista1 arm (fig. 5, P. peronis). Further separated
from P. peronis as follows: maxillary palpus with second segment more
than 14 times as long as first (fig. 1, MX.P.), not merely slightly longer
(fig. 7, MX.P.); large acetabular bristle inserted well above acetabulum
(fig. 13, AC.B. and AC.) (as in P. acodonlis) instead of approximately
at same leve1 (fig. 15, AC.B. and AC.); dista1 arm of ninth sternum bearing a submarginal mesa1 row of bristles (fig. 4) which is lacking in P.
peronis (fig. 5); crochet armature (fig. 10, CR.A.) well developed, appearing on each side as a pair of subcontiguous rod-like sclerites, instead of being so reduced that it appears as a single thin line (fig. ll,
CR.A.) ; spermatheca (fig. 14) head with somewhat of a dorsal hump,
being about one and one-half times as long as broad instead of twice as
long as broad (fig. 17). Separable from P. acodontis in that the dista1
arm of the ninth sternum is apically quite broad (fig. 4), not narrowed;
this structure with subapical region about as broad as median portion
of digitoid (fig. 13, F.), not definitely narrower than F. Spermatheca
with head narrower than in P. acodontis, in which it is only slightly
longer than broad.
Description
Head, Male (fig. l).-Frontoclypeal
margin evenly rounded, except
for acuminate, dorsad-directed tubercle inserted slightly above leve1 of
eye and arisíng from margin of subovate sclerotized area. Micropunctations scattered above and in front of first row of bristles on both preand post-antennal regions. Preantennal area with three rows of bristles,
first row consisting of about four to six bristles somewhat variable in
size; second row consisting of three bristles, of which ventromarginal
one is longest; third row of two bristles, one behind eye and one submarginal, ventral to eye. Eye conspicuous,.subovate, broader than Iong;
anteroventral portion associated with tentorial arm, which usually appears as a rod-like structure. Genal process broad. Maxillary lobe extending to apex of third segment of maxillary palpus (MX.P.) ; labial palpus
113
VENEZUELAN FLEAS
k
.
r
.
i
íive-segmented, extending approximately to apex of forecoxa. First antermal segment about two and one-half times length of second; most of
bristles antero- or caudomarginal; second antennal segment about twíce
as broad as long, with a dorsomarginal fringe of bristles; asymmetrical
club with apparently eight segments but in reahty true first segment
lying mesa1 to rest of club. Above club a row of short though fairly
broad bristles, bases of bristles virtually contiguous; with a few ama11
bristles above first two segments. Post-antennal region with three rows
of bristles arranged approximately 4-7-6; small hairs intercalated between bases of those in last row; the ventrahnost bristle of each row
longer than others in row.
Thoraa: (figs. 1 and 3).-Pronotum
(PRN.) and mesonotum (MSN.)
with two rows of bristles, those of second row longer and with small
intercalaries; flange of each notum lacking mesal pseudosetae. Mesepisternum (MPX.) with two bristles. Mesepimere (MPM.) with three
bristles arranged 2-l. Metanotum (MTN.) with three rows of bristles,
those of posterior row longest and with intercalaries; metanotal flange
with five or six somewhat pale, smaI1 apical spinelets on each side. Latera1
metanotal area (L.IM.) distinct, with caudal margin lateral to and extending over most of pleural arch; with three or four small bristles near
anteroventral margin and two longer bristles bordering pleural arch.
Metepisternum
(MT&‘.) with a long bristle near posterodorsal angle;
squamulum absent. Pleural ridge of metasternosome fitting into a wellscelerotized socket, the pleural arch (PL.A.). Metepimere (MTM.) with
two rows of bristles arranged 4-6, 4-5.
Legs.-Procoxa with many lateral bristles scattered over entire length
of segment. Meso- and metacoxae with relatively few sueh bristles and
those submarginal or subapical. Profemur with about sixteen small,
scattered, non-marginal lateral bristles and one mesa1 bristle; with two
fairly long, subapical, ventromarginal
bristles. Mesofemur similar but
with ten to fifteen scattered non-marginal lateral bristles. Metafemur
with a lateral, subventral row of five or six thin bristles and three or
four subdorsal ones; metatibia (fig. 2) with large dorsomarginal notches
bearing bristles as follows (from base to apex): 2-2-2-3-2-(excluding
apical 3). Measurements of tibiae and segments of tarsi (petiolate base
deIeted) of holotype shown in microns:
LEQ
TIBIA
Pro-. . . . . . . . . . 247
Meso- . . . . . . . . 370
Meta- . . . . . . . . 515
1
52
103
330
II
72
155
258
Taasny SEGMENTO
III
Iv
52
41
93
41
134
62
V
103
124
144
Pro- and mesotibiae with one or more apical bristles extending distad
of first tarsal segments. Second and third hindtarsal segments with an
apical bristle extending beyond apex of following segment. First tarsal
segment of each leg bearing three pairs of relatively long lateral plantar
114
BULLETIN
OB THE PAN AMERICAN SANITARY BUREAU
bristles, a fourth pair of much shorter lateral plantar bristles and an
apical submedian pair subequal in size.
Abdomen.-First
tergum (fig. 3, IT.) with two rows of bristles, those
of posterior row longer and with intercalaries; with a total of about fourteen to sixteen somewhat pale, apical spinelets. Xucceeding terga with
apical spinelets in holotype arranged 6-10-6-2 (totals); in female 10-7-4-l.
In males, second and third terga with lowest bristle in first and second
rows inserted slightly ventrad to spiracle; or other terga, first row much
more abbreviated; in females first row always extending to spiracle.
Basal sternum in male with one small ventromarginal bristle and two or
three small lateromedian bristles; in female, with two or three ventromarginal bristles on each side and about five small lateromedian bristles.
Unmodified sterna usually with a row of four or five bristles. In female,
third sternum with a patch of three to five much smaller bristles preceding this row. With only one antepygidial bristle in each sex.
ModijZed Abdominal Segments, Male (fig. 6.).-Eighth
tergum (8T.)
reduced, extending caudad only to a leve1 slightly beyond middle of
sensilium and ventrad to near base of manubrium; its spiracle long and
narrow. Eighth sternum (8S.) very large, covering much of the genitalia,
and extending ventrad from leve1 of lowest bristle of seventh tergum
and apicad to near leve1 of acetabulum; with about four or five long
bristles and three or four smaller ones.
Immovable process of clasper (P. and fig. 13) very large, dorsal margin much longer than movable finger; with apex extending somewhat
distad of movable finger; dorsal margin slightly sinuate; caudal margin
becoming convex at insertion of acetabular bristle (AC.B.); caudal
margin with a small tubercle at apical sixth; opposite this projection a
circular thickening which is frequently accompanied by a small, thin
mesa1 bristle; with about six dorsomarginal, fairly long bristles-two
of
these apical and last of these somewhat longer and stouter than others;
with two submarginal, more median long bristles and a few small scattered thin submarginals; a very long acetabular bristle about as long as
F. inserted slightly below midpoint; with two widely separated smaller,
marginal bristles at leve1 of acetabulum; with two or three submedian,
thin, small, mesa1 bristles and a patch of similar bristles along caudal
margin of curve above base of manubrium (MB.). Manubrium long and
narrow, fairly straight, apically somewhat dilated. Tergal apodeme of
ninth segment (T.AP.9) forming dorsal, proximal portion of clasper
lobe; with anterior margin markedly convex. Ninth tergum greatly reduced to a narrow, indefinite area between its apodeme and clasper
lobe. Digitoid or movable finger (F., & fig. 13) about five times as long
as broad; gently curved cephalad; extending slightly distad of the subapical thickening and tubercle on caudal margin of P.; with small, thin
subapical marginal bristles; caudal margin with a group of about five
PLATE
1
.
FIG. l.-Polygenis
ambersoni sp. nov. Head, male.
FIG. 2.-Ibid.
Posterior tibia; male.
FIG. 3.-Ibid. Thorax; male.
115
PLATE
II
J
l
;W&S
Fm. 4.-Polygenis
FIQ. 5.-Pdygenis
sternum; male.
FIG. ô.-Polygenis
5 & 6 POLYGENIS AMBERSONI SPNOV
POLYGENIS PERONIStJ.&R., 1923)
ambersoni sp. nov. Dista1 arm of ninth sternum; male.
peronis (Jordan and Rothschild, 1923); Dista1 arm of ninth
ambersoni sp. nov., Modified segments; male.
116
February 19&?]
VENEZUELAN FLEAS
117
thin bristles above acetabular bristle and one thin bristIe below acetabular bristle; with two or three small thin anteromarginal bristles.
Proximal arm of ninth sternum (P.A.9) roughly shaped like a short
broad boot; its shank slightly narrowed at the middle. Dista1 arm of
ninth sternum (D.A.9 and fig. 4.) proximally produced into a long heel
(HL.9); about eight times as long (as measured from base of heel) as
broad (measured at midpoint); with sides parallel for most of its Iength;
slightly curved dorsally; apex somewhat rounded; apical third with
four or five very long, lateromarginal bristles; with a mesa1 row of marginal thin bristles commencing shortly above base of heel and extending
to apex; with a group of five or six apical or subapical bristles, some of
which are mesal.
Aedeagus (figs. 6, 9, and 10) essentially of the type of P. adocetus
Traub, 1950 (q.v., p. 65, pl. 38, fig. 5 and pl. 39, fig. 1). Aedeagal apodeme
(AE.A.) about as long as aedeagus proper; somewhat less than half as
broad as long. Lateral plates dorsoapically produced into acuminate
projections (A.L.L.) overlapping portion of endchamber; ventroproximally produced into an acuminate, conspicuous heel (fig. 6 and 9,
HL.) ; with a long narrow rod arising from heel (probably homologue
of aedeagal apodemal rod); hood flaps (fig. 9 and 10, H.F.) with a saggitate proximoventral
lateral extension on each side. Median dorsal
lobe (M. D.L.) convex; extending to apex of hood flaps; distally associated
with a sclerite on each side (A.M.S.) which is probably the homologue
of the apicomedian sclerite of many other genera of íleas; most of apicomedian sclerite weakly sclerotized but with apical portion distinct as a
narrow, subverticd structure. This portion of A.M.S. laterally bearing
a dorsal tuft of small rugae or spicules and a subventral, mesa1 patch
of short, semimembranous filaments (CR.P.), the crochet processes.
Basal segment of sclerotized inner tube (hg. 9, I.T.-B) shorter but much
broader than elongate, apical portion, (I.T.-A). With only one per& rod
(fig. 6, P.R.) extending cephalad of apex of aedeagal apodeme, and that
slightly; second rod much shorter. Well-developed pseudotube (PS.?‘.)
with most of its ribs (fig. 9, RIB.) inclined distodorsad, extending to
leve1 of A.L.L. Crochets (CR.) conspicuous, appearing as large cordate
structures extending slightly apicad of ventral portion of A.M.S. and
overlapping dorsoapical portion of pseudotube. Endchamber with an
elongate, Y-shaped sclerite here designated the furca (FU.), one arm of
which is lightly sclerotized. Ventral arm of furca well sclerotixed, extending to near apex of crochet. Each crochet associated with a somewhat V-shaped sclerite (CR.A.) here designated the crochet armature,
the ventral arm of which is weI1 sclerotized and consists of two virtually
contiguous rod-Iike extensions reaching ahnost to apex of pseudotube.
Lateral lobes (L.L.) very Iong, fairly well sclerotized; extencling from
below crochet processes to an indefinite area near apodemal strut; in
vicinity of coiled per& rods, each lobe bearing a lateral sclerite which
118
BU’f&ÉTEN ÓF THE PAN AMERICAN SANITARY BUREAU
appears on each side as an oblique, very shallow, elongate ellipsoid resembling a slightly concave disc seen on end; this sclerite herein designated as the fender (FNR.). The fender lying ventrad (but laterad) to
balloon of coil of penis tubes. Side pieces (Á3.P.) well developed, shaped
like a sinuate spindle, vertical, laterad to sclerotized inner tube near
base of A.I.T. Crescent sclerite (C.S.) distinct. Lateroventral sclerite
(L.S.) of apodemal strut large, well developed, as is the adjacent vesicle
(V.). With a semimembranous, mantle-like structure enclosing proximal
portion of endchamber but mesa1 to lateral lobe; extending cephalad to
vicinity of aedeagal heel, and distad to near thickening of pseudotube;
ventral margin fluted. This structure is herein designated the fluted
membrane (F.M.).
Tenth abdominal segment with sensilium large, fairly flat, about
sixteen pits per side. Dorsal anal lobe (fig. 6, D.A.L.) well sclerotized,
subconical, extending slightly more dorsad than sensilium; slightly less
than twice as long as broad at base; with apical and subapical bristles.
Ventral anal lobe (V.A.L.) about twice as long as dorsal lobe; much more
acute; with ventral margin more heavily sclerotized than dorsal margin;
with similar apical bristles.
ModZfied Abdominal Xegments, Female (fig. 12).-Seventh
sternum
with dorsal margin oblique, fairly straight; caudal margin very slightly
sinuate, subvertical; with nine bristles, of which about three are submarginal and two marginal. Eighth tergum very long; with anterior
margin extending from above base of antepygidial bristle to near ventral
margin of seventh sternum; this margin markedly curving caudad at
midpoint before plunging ventrally; with a row of bristles commencing
anterior to and above the elongate spiracle and extending to near ventral
margin of segment; those near spiracle short, rest usually longer; with
a group of about four or five small lateromedian bristles anterior to the
row; with about thirteen caudomarginal or submarginal bristles, the
longest two at dorsocaudal angle below ventral anal lobe; with a patch
of small thin mesa1 bristles below ventral anal lobe and another of two
or three near apex of eighth sternum. Dorsal anal lobe (D.A.L.) with a
row of dorsomarginal bristles and five or six lateromedian ones, of which
one or two are inserted below anal stylet. Anal sytlet (AS’. and fig. 16)
about twice as long as broad; anterior margin with a small subapical
sinus bearing two extremely small hairs; with a long apical bristle.
Ventral anal lobe (V.A.L. and fig. 16) angulate, broad; with a marginal
row of about nine bristles, variable in length, some mesal; with about
four smaller subapical bristles. Spermatheca (SI’. and fig. 14) with head
about twice as long as broad; dorsal margin convex, with a short but distinct tubercle; ventral margin slightly sinuate; tail up-turned, extending
dorsad of head. Bursa copulatrix (B.C.) apparently an indistinct subglobular structure; with a short sclerotized duct arising from median
portion of sphere.
.
PILA’333 III
M&S
; 8r 8 POLYGENIS PERONIS (J. & R., 1923)
POLYGENIS AMBERSONI SI? NO\/:
FIG. 7.--Polvgenis pero& (Jordan and Rothschild,
head : male.
F&. 8.-Ed.
Apex of aedeagus.
FIG. 9.-Polygenis
ambersoni sp. nov. Aedeagus.
119
192) ; Anterior portion of
PLATE
IV
.
e.L.
FIGURES 10 8 12 POLYGENIS AMBERSONI Sí? NOV.
FIGURE I I
POLYGENI S PERON I S (J. & R.,1923)
FIG. lO.-Polggenis
Fra. ll.-Polygenis
FIG. 12.-Polygenis
ambersoni ep. nov. Apex of aedeague.
peronis (Jordan and Rothschild, 1923) ; Apex of aedeagus.
ambersoni sp. nov. Modified segmenta; female.
120
PLATE
V
4
c
.
,
FIGURES I3,14 & 16 POLYGENIS AMBERSONI Si? NOV
FIGURES 15 8 17 POLYGENIS PERONI S (J. & R, 1923)
E‘IG.
13.-Polygenis ambersoni sp. nov. Immovable process and digitoid of
clasper. FIG. 14.-Ibid. Spermatheca; female. FIG. 15.-PoEygenis pero& (Jordan
and Rothschild, 1923); Immovable process of digitoid of clasper. FIG. IB.-Polygenis ambersoni, Anal stylet and ventral anal lobe of proctiger, female. FIG. 17.
-PoZygcnis
peronis (Jordan and Rothschild, 1923) Spermatheca; female.
121
PLATE
VI
c
POLYGENIS ROBERT] BEEBEI II. FOX, 1947)
FIG. 18.-Polygenis roberti beebei (1. Fox, 1947), Head; male.
FIG. 19.-Ibid. Posterior tibia; male.
FIG. ZO.-Ibid. Thorax; male.
122
February 1952]
VENEZUl3LAN FLEAS
123
COMMENT ON THE AEDEAGUS OF P. ambersoni AND RELATED FLEAS
The pseudotube (figs. 9 and 10, PS.T) is in reality a stout sclerite in
the middle of the floor of the endchamber. It is subapically dorsally
thickened, and it is this dorsal sclerot,ization which gives the structure
of a tube when seen from the lateral aspe&, as in mounted specimens.
From the proximal portion of the psuedotube arise a series of many thin,
subvertical, convex rod-like sclerotizations, designated above as ribs
(RlB.). Most of the ribs are inclined or pitched dorso-apicad, but some
apical ribs usually are inclined dorsoproximad. The more apical ribs arise
from a slightly sclerotized sternum-like structure which lies on the dorsal
thickening of the pseudotube and which is frequently diflicult to see.
The presente of ribs on the pseudotube is characteristic of this group of
fleas, as are the fluted membrane (F.M.), the side-pieces (S.P.), the
fenders (FNR.), and the furca (FU.), as well as structures previously
mentioned (Traub, 1950).
Remarks.-This
species is named for one of the collectors, Commander
J. M. Amberson, M.C., of the Naval Medical School, who has contributed
much to the field of parasitology
Polygenis roberti
and tropical
(Rothschild,
medicine.
1905)
Pulen robe& Rothschild, 1905, Nov. Zool. 12: 479481, figs. 1, 2.
RhopalopsyUusrobe& Jordan and Rothschild, 1908, Parasitol. 1: 77. Jordan
and Rothschild, 1923, Ectoparasites 1: 330, 350. Dalla Torre, 1924, Ber. Nat.
Med. Ver. Inns. 39: 22. Pinto, 1929, Arthr. Parasit. 1: 326. Wagner, 1937,
Zeits. Parasit. 9: 699. Costa Lima, 1937, Rev. Med. Cirur. Bras. 45(2): 117.
Polygenis roberti Jordan, 1939, Nov. 2001. 41: 446, fig. 329. Guimarães,
1940, Arq. 2001.2: 223. Costa Lima and Hathaway, 1946, Monog. Inst. Oswaldo
Cruz No. 4: 143.
Many specimens of Polygenis roberti were collected during the plague
survey. Study showed that these constitute a distinct subspecies and are
the same as the form previously described as Rhopalopsyllus beebei 1.
Fox, 1947. Dr. FOX kindly verified our belief that the present specimens
are the same as those described as beebez’.For purposes of clarification,
Polygenis roberti beebei (1. Fox) (new combination) is re-figured and redescribed below.
Comparison of the paratype male of PoZygenis tripopsis Guimaráes,
1948, with Venezuelan material and with P. roberti robe& led us to
believe that P. tripopsis is also a subspecies of roberti. Our conclusions
regarding the subspecific status of these two forms are based on the
similarity of details of the specialized aedeagal sclerites and of the other
male modifled segments (with the exception of the ninth sternum). The
chaetotaxy of the metatibia, used as a distinguishing character in the
original descriptions of tripoptis and beebei, was found to be somewhat
variable and does not warrant the retention of these forms as full species.
For example, the Venezuelan specimens from one area exhibit 211 the
listed tibial variations.
124
BULLETIN
OF THE PAN AMERICAN SANITARY BUREAU
The three subspecies may be differentiated on the basis of the shape of
the male ninth sternum. In P. roberti Iripopsis (fig. 29) the dista1 arm is
of almost equal width from base to apex, with apex subrounded. In
P. roberti beebei (fig. 30) the dista1 arm is characterized by being noticeably convex on the ventral (posterior) margin; apex subrounded, the
arm slightly upcurved. P. roberti roberti (fig. 31) is distinct in that the
dista1 arm is narrower at base than at rounded apex.
Polygenis roberti beebei (1. Fox, 1947), new combination
18-28 and 30)
(figs.
Species B (?) Hecht, 1942. Revista de Sanz’dud y Aeistencia Social ‘7(6): 815,
fig. 1-B.
Rhopalopsyllus beebei1. Fox, 1947, Zoologica 32(13) : 117.
[email protected]~on.-(Only differences from P. umbersoni are included.).
Nead, male (fig. 18).-Frontal
tubercle arising from triangular or
subtriangluar sclerotixed area. First preantennal row of five to six bristles,
most dorsal longest. Maxillary
lobe extending beyond apex of third
segment of maxillary palpus. Labial palpus about five-sixths length of
forecoxa. Preantennal region with three rows of bristles arranged approximately 4-5-6.
Thorm (fig. 18 and 20).-Metanotum
with three to five apical spinelets on each side. Lateral metanotal area with one to four small bristles
near anteroventral margin.
Legs.-Profemur
with two mesa1 bristles. Metatibia (fig. 19) with
large dorsomarginal notches bearing bristles as follows: 2-2-2-3-l(2, or
missing)-3; apical group of three or four bristles, usually four. Second
and third hindtarsal segments with one or two apical bristles extending
to or beyond apex of following segment.
Abdomen.-Basal
abdominal sternum with four to five small ventromarginal bristles and four or five small lateromedian bristles.
Modi$ed Abdominal: Segments, Mule (fig. al).-Eighth
sternum (SS.)
with four or five long bristles and one or two smaller ones. Dorsal margin
of immovable process of clasper (P. and fig. 28) with about nine mediumsized marginal and submarginal bristles, apical one longest; with severa1
smaller submarginal bristles. Caudal margin with two bristles, longer
inserted just dorsad to acetabulum. Three or four small, submedian
mesa1 bristles posterior to and slightly dorsal to patch of mesa1 bristles
just posterior to base of manubrium. Tergal apodeme of segment nine
(T.AP.9) with anterior margin subtruncate. Dista1 arm of ninth sternum (D.A.9 and fig. 30) slightly upcurved, especially distally; proximally
produced into an obtusely acuminate heel; arm narrowed apically;
posterior (ventral) margin more convex than anterior; with a narrow
rounded apex; with twenty to twenty-four lateromarginal,
subequal
bristles extending from near heel to apex.
Aedeugus (figs. 21 and 23).-Aedeagal
apodeme (AE.A) three times
PLATE
VII
..“.<
i. .
is+.
POLYGE;IS ROBERT1 BEEBEI (IFOX, 1947)
KL.
FIG. 21.-PolygenZs robe& beebei (1. Fox, 1947) Modified segments; male.
FIG. 22.-Ibid. Dorsal and ventral anal lobes of proctiger; male.
Fra. 23.-Ibid. Aedeagus.
1%
PLATE
VIII
24
\
POLYGENIS ROBERT1 BEEBEI (I.FOX, 1947)
FIG.
female.
FIG.
FIG.
FIG.
FIQ.
24.-Polygenis
25.-Ibid.
26.-Ibid.
27.-Ibid.
28.-Ibid.
roberti
beebei (1. Fox, 1947) Ventral anal lobe of proctiger;
Modified segmenta; female.
Anal stylet; female.
Spermatheca; female.
Immovable process and digitoid of clasper; male.
126
February 1962j
VENEZUELAN FLEAS
127
as long as broad. Hood flaps (H.F.) elongate, extending cephaIad beyond
coiled penis rods (P.R.) ; the margin somewhat sclerotized part of its
length. Median dorsal lobe (M.D.L.) convex, extending ahnost to apex
of hood flaps. Apicomedian sclerite (A.M.S.) dorsally with a patch of
semimembranous filaments (CR.P.), the crochet process. One penis
rod (fig. 21, P.R.) extending well beyond apex of aedeagal apodeme,
second rod ending short of apex. Ribs (R IB.) ending distally in a rounded
spiculated process cephalad to flared apex of pseudotube (PS.T.).
Crochets (CR.) apparent as slightly sinuate broad rods; corrugated
along longitudinal
aspe&, extending apicad of A.M.S. With paired
wing-shaped pale sclerites, crochet arms, (CR.A.) extending from base
of crochet distoventrally
to anterior leve1 of pseudotube, with convex
dorsal margin, rounded at postero-apical angle, which is produced caudad
about one-half length of crochet. Ventral arm of furca (FU.) extending
caudad about one-half length of crochet. Fender (FNR.) more or less
haIf-moon shaped. Side piece (S.P.) large, triangular, with longest side
sinuate, apparently arising near base of vesicle, but lateral to vesicle.
Dorsal anal lobe (fig. 21, D.A.L. and fig. 22) extending as far dorsally
as sensilium.
Modified Abdominal Segments, Femaíe (fig. %).--Seventh
sternum
with dorsal margin oblique; caudal margin rounded, vertical; with six
long bristIes and one or two submarginal smaller bristles, which may
be absent. Eighth tergum with definite angle on posterior margin; with
row of bristles, alternating large and small, extending from above spiracle
to well below midpoint. Anal stylet (A.S. and fig. 26) with Iong apical
bristle and one very small subapical hair on ventral margin. Ventral
anal Iobe (V.A.L. and fig. 24) angulate, twice as long as broad at base,
bristles confined to caudal margin. Spermatheca (SP. and fig. 27) about
three-fourths as broad as long; with tubercle on head indistinct.
Since 20 of 44 Sigmodon hispidus hirsutus were parasitized with
this flea, it seems likely that this rat is the preferred host.
Records:
Ex Didelphis m. marsupialis Linnaeus (2/42), 6 3, 4 o
Ex Sigmodon hispidus hirsutus Eurmeister (20/44), 17 CF, 13 0
Ex Alcodon urichi urichi Allen and Chapman (l/lO), 1 d, 1 o
Ex Mus musculus brevirosíris Waterhouse (l/l) 1 o
Ex Rattus rattus frugivorus Raíiiesque (2/6). 1 8, 1 0
Ex Heteromys a. anomalus Thompson (l/ll),
1 d
Ex Oecomyst. trinitatis Allen and Chapman (7/7), 17 $, 7 P
Ex Oryzomys fulvescens delicatus Allen and Chapman (3/5), 4 d 6 p
Ex Neacomys spinosus ssp. nov. (2/2), 1 3, 3 o
Polygenis b. bohlsi (Wagner, 1901)
Like P. roberti beebei, tbis is one of the most dominant species in the
area, not only in the number of individuals,
but in the variety Gf hosts
128
BULLETIN OF TEE PAN AMERICAN BANITARY BUREAU
that it infests. Although this species has often been referred to in the
literature, it has not been adequately illustrated. P. bohlsi is readily
separated from others in the genus by the following: Digitoid (fig. 33, F.),
shaped roughly like a low isosceles triangle; the dista1 arm of the ninth
sternum (fig. 35) bearing an apical dense patch of spiniform bristles;
fender of the aedeagus (fig. 32, FNR.) crescent-shaped; spermatheca
(fig. 34) with a very characteristic shape, its head four-fift,hs as broad
as long, subtriangular, tail much shorter than head and somewhat upturned; female eighth tergum with median patch of about 8 thin bristles
anterior to row of bristles extending from ventral to dorsal margins.
Records:
Ex DideZphis m. marsupialis Linnaeus (2/2), 2 o
Ex Sigmodon hispidus hirsutus Burmeister (31/44), ll d, 38 9
Ex Akodon urichi urichi Allen and Chapman (8/10), 4 8, 13 o
Ex Rattus rattus frugivorus Raíinesque (3/6), 4 Q
Ex Heteromys a. anomalus Thompson (5/11), 2 8, 7 o
Ex Oecomyst. trinitatis Allen and Chapman (1/7), 1 #, 1 Q
Ex Oryzomys pulvescens delicatus Allen and Chapman (2/5), 2 3, 4 o
Ex Neacomys spinosus ssp. nov. (1/2), 1 Q
Ex Proechimys cayennensis trinitatis Allen and Chapman (2/4), 1 d
(Also one female from Campamento
Chacin, River
Tuy).
Ex Xciurus granatensis chapmani Allen (3/3), 3 Q
From the above list it will be seen that Sigmodon hispidus hirsutus and
Akodon urichi urichi are the preferred hosts of this species, although its
tastes are quite catholic.
Polygenis klagesi samuelis (Jordan and Rothschild,
Ex
Ex
Ex
Ex
1923)
Didelphis m. marsupialis Liiaeus (1/2), 1 Q
Akodon urichi urichi Allen and Chapman (l/lO) ; 1 CF
Rattus rattus frugivorus Rafinesque (1/6), 1 c?, 1 Q
Proechimys cayennensis trinitatis Allen and Chapman (3/4), 3 Q
(and one female from Campamento Chacin)
Polygenis steganus (Jordan and Rothschild,
Ex Didelphis m. marsupialis
1923)
Linnaeus (1/2), 2 8
Polygenis peronis (Jordan and Rothschild,
ll, 15 and 17)
1923) (figs. 5, 7,8,
This species has not been reported since the original description. It
has been above compared with P. ambersoni sp. nov. and pertinent drawings are included. The following records indicate that it is a fairly charactetitic parasite of Heteromys a. anomulus:
Ex Sigmodon hispidus hirsutus Burmeister (2/43), 1 d, 1 P
Ex Akodon urichi urichi Allen and Chapman (2/10), 2 0
Ex Heteromys a. anamalus Thompson (6/11), 6 d, 10 ?
PLATE
IX
.
FIG. 29 POLYGENIS ROBERT1 TRIPOPSIS GUIMARiíESJ948
FIG. 30 POLYGENIS ROBERT1 BEEBEI (1. FOX, 1947)
FIG. 3 I POLYGENIS ROBERT1 ROBERT1 (ROTHS., 1905)
FIG. 29.-Polygenie
robe& tripopsis
Guimarães, 1948; Distal arm of ninth
sterum; male. Fra. 30.-PoEygenis
roberti beebei (1. Fox, 1947) ; Dista1 arm of
ninth sternum; male. FIG. 31.-E’olygenis robe& robe& (Ftothschild, 1905); Dktal
arm of ninth sternum; male.
í29
PLATE
X
POLYGENIS B. BOHLSI (WAGNER, 1901)
FIG. 32.-Polygenis
b. bohlsi (Wagner, 1901) Aedeagus; male.
FIG. 33.-Ibid. Immovable procesa of digitoid of clasper; male.
FIO. 34.-Ibid.
FIO. 35.-Ibid.
Spermatheca; female.
Dista1 arm of ninth sternum; male.
130
u
Febrmry ILVZ?]
131
vENEzlJxLAxFLEAs
Polygenis sp.
Females of this genus are often extremely difficult to identify unless
accompanied by males. It is unfortunate that only females were represented in the material collected in Campamento Chacin, at River Tuy.
Some of these females are very close to Polygenis ambersmi sp. nov.
Ex Raftus rattus alexandrinus Geoffrey (2/2), 2 Q
Ex Proechimys cayennensis trinitatis Allen and Chapman (l/l),
Ex Heteromys a. anomahs Thompson (l/l), 3 Q
Ex Xigmodon hispidus hirsutu-s Burmeister (l/l), 1 Q
Rhopalopsyllus
austral% australis (Rothschild,
Ex Akodon u. wichi Allen and Chapman (l/lO),
2 Q
1904)
1 d
Ex Didelphis m. marsupialis Linnaeus (1/2), 1 Q
Rhopalopsyllus
Family PULICIDAE
cacicus saevus (Jordan and Rothschild,
Ctenocephalides
1923)
felis felis (Bouché, 1835)
Ex Rattus rattus frugivorus Rafinesque (1/16), 1 Q
II. LISTS OF HOSTS AND THEIR
ECTOPARASITES
LOCALITY A. CAMPAMENTO RAFAEL RANGEL
Didelphis marsupialis marsupialis Linnaeus
Polygenis roberti beebei(1. Fox, 1947) 6 d, 4 Q
Polygenis sp. 1 Q
Tritopsylla intermedia oxyura (Jordan, 1926) 7 3, 7 Q
Polygenis bohlsi bohlsi (Wagner, 1901) 2 Q
Polygenis steganus (Jordan and Rothschild,
1923) 2 3
Rhopalopsyllus cacicus saevus (Jordan and Rothschild, 1923) 1 Q
Polygenis klagesi samuelis (Jordan and Rothschild, 1923) 1 Q
Rhopalopsyllus australis australis (Rothschild, 1904) 1 d
Polygenis roberti beebei(1. Fox, 1947) 1 c?, 1 0
Mus musculus brevirostris Waterhouse
Polygenis roberti beebei(1. Fox, 1947) 1 Q
Rattus rattus frugivorus Rafinesque
Polygenis klugesi samuelis (Jordan & Rothschild, 1923) 1 3, 1 Q
Ctenocephalidesfelis felis (Bouché, 1835) 1 Q
Polygenis b. bohlsi (Wagner, 1901) 4 Q
Polygenis roberti beebei (1. Fox, 1947) 1 #, 1 Q
Heteromys a. anomalus Thompson
Polygenis roberti beebei(1. Fox, 1947) 1 d
Polygenis b. bohìsi (Wamer, 1901) 2 d, 7 Q
Polygenis peronis (Jordan & Rothschild, 1923) 6 c?, 10 Q
132
BULLETIN OF TIIE PAN AMERICAN SUlITARY
BUREAU
Oeeomyst. tridztis Allen & Chapman
Polygenis roberti beebei(1. Fox, 1947) 17 3, 7 ?
Polygenis b. bohlsi (Wagner, 1901) 1 9
Oryzomys fulvescens delicatus Allen & Chapman
Polygenis roberti beebei(1. Fox, 1947) 4 8, 6 o
Polygenis b. bohlsi (Wagner, 1901) 2 $,4 P
Neacomys spinosus ssp. nov.
Polygenis robe& beebei(1. Fox, 1947) 1 d, 3 o
Polygenis b. bohlsi (Wagner, 1901) 1 o
Proechimys cayennensis trinitatis Allen & Chapman
Polygenis 6. bohlsi (Wagner, 1901) 1 d, 1 0
Polygenis klagesi samuelis (Jordan & Rothschild,
Xciurus granatensis chapmani Allen
Polygenis b. bohlsi (Wagner, 1901) 3 o
1923) 3 o
LOCALITY B. CAMPAMENTO CHACIN, TUY RIVER
Rattus rattus alexandrinus Goeffrey
Polygenis sp. 2 0
Proechimys cayennensis trinitatis Allen & Chapman
Polygenis b. bohlsi (Wagner, 1901) 1 d, 1 P
Polygenis klagesi samuelis (Jordan & Rothschild,
Polygenis sp. 2 0
Heteromys a. anomalus Thompson
Polygenis sp. 3 9
Sigmodon hispidus hirsutus Burmeister
Polygenis sp. 1 0
1923) 1 o
Rema&.-In
the preceding drawings, mesa1 structures
are indicated
in the conventional
manner, i.e., by broken lines. Bristles which were
missing on the side of the flea which was drawn but which were present
on the other side or on other specimens, are indicated by representing
one of the margins of the bristles with a broken line.
ABBREVIATIONS
AC.
AC.B.
AE.A.
A.I.T.
A.L.L.
A.M.S.
A.S.
B.C.
CR.
CR.A.
CR?.
T.S.
Acetabulum
Acetabular bristle
Aedeagal apodeme
Armature of inner tube
Aocessory lateral lobe of
aedeagus
Apico-median
sclerite of
aedeagus
Anal stylet
Bursa copulatrix
Crochet
Crochet armature
Crochet processes
Crescent sclerite of aedea-
w*
D.A.L.
D.A.9
SS.
IT.
8T.
F.
F.M.
FNR.
FU.
H.F.
HL.
HL.9
I.T.-A.
Dorsal anal lobe of proctiger
Dista1 arm of ninth sternum
Eighth sternum
First tergum
Eighth tergum
Digitoid or movable finger
Fluted membrane
Fender
Furca
Hoodflaps of aedeagus
Heel at base of aedeagal
pouch
Heel of ninth sternum
Apical portion of sclerotiaed inner tube
Febrero íN..]
I.T.23.
L.L.
L.M.
L.S.
MB.
M.D.L.
‘!
4
=
MPM.
MPS.
MSN.
MTM.
MTN.
MTS.
MX.P.
PULGAS VENEZOLANAS
Basal segment of sclerotized
inner tube
Lateral lobes of aedeagus
Lateral metanotal area
Lateroventral sclerite
Manubrium
Median dorsal lobe of aedeaggus
Mesepimere
Mesepisternum
Mesonotum
Metepimere
Metanotum
Metepisternum
Maxillary palpus
P.
P.A.9
PL.A.
P.R.
PRN.
PS.T.
RIB.
SP.
S.P.
T.AP.9
V.
V.A.L.
133
Immovable prooess of ciasper
Proximal arm of ninth sternum
Pleural arch
Penis rod
Pronotum
Pseudotube
Ribs
Spermatheca
Side pieces
Apodeme of ninth tergum
Vesicle
Ventral anal lobe
REFERENCES
Fox, 1.: Notes on Ectoparasites from Venezuela, Zoologica, New York Zoological
Society, %?(13):117-119, 1947.
Hecht, Otto: Las pulgas de las ratas en Venezuela, Rev. San. y Asid. Soc., 7(6) :811820, 1942.
Jordan, Karl: Flöhe sus Venezuela, Zeita. jur Parasitenkunde, 5(2):264-266, 1
fig., 1931.
Traub, R.: Siphonaptera from Central Ameriea and Mexico: A Morphological
Study of the Aedeagus, with Descriptions of New Genera and Species,
Zoological Memo&
oj the Chicago Natural History Museum, l(1) :l-127
(Plates l-54), 1950.
PULGAS
5
COLECCIONADAS
EN UN ESTUDIO
VENEZUELA
(Sumario)
DE LA PESTE EN
A invitación del Gobierno de Venezuela y de la Oficina Sanitaria Panamericana, una Misión de la Escuela Médica Naval, Bethesda, Maryland, llevó a cabo
en 1950 un estudio de la peste selvática en Venezuela. El equipo epidemiológico
estuvo integrado por el Comandante J. M. Amberson, el Dr. Ernst Schwarz y
la Sra. H. K. Schwarz. En este trabajo se presentan los datos sobre las pulgas
coleccionadas por los miembros de la Misión, asf como la descripción e ilustraciones de una especie nueva, y de las formas afines o que poseen importancia
médica potencial.
Reviste sin duda importancia epidemiol6gica el hecho de que ninguna de las
pulgas coleccionadas por la Misi6n en esas zonas endémicas pertenecfan al
género Xenopsylla, que es el que interviene en la trasmisión clásica de la peste.
5610 se encontró un ejemplar de una especie cosmopolita: C¿enocephalkZesfelis
felis (Bouché, 1835); todos los demás ejemplares fueron de pulgas autóctonas.
La mayorfa de las pulgas que aparecen en la lista fueron coleccionadas en
Aragua, Campamento Rafael Rangel; algunas fueron coleccionadas en el
Campamento Chacfn, Río Tuy, Miranda.
134
BOLETfN DE LA OFICINA SANITARIA PANAMERICANA
Se coleccionaron las siguientes especies de pulgas, indicAndose en cada caso
el huésped característico:
Familia HystrichopsyEZidae
Subfamilia Ctenopthalminae
Tritopsylla
intermedia oxyura (Jordan, 1926)
Ex Didelphis m. marsupialis
Familia Rhopalopsyllidae
Polygenis ambersoni sp. nov.
Ex Sigmodon hispidus hirsutus
Polygenis
roberti beebei (1. Fox, 1947)
Especie muy comtin que se encuentra en varios huéspedes, en particular
m. marsupial&
Sigmoclon hispidus hirsutus, Oecomys t. trinitatis y
Oryzomys fulvescens debicatus.
Didelphis
Polygenis b. bohlsi (Wagner, 1901)
Esta es una de las especies m6s dominantes en la zona y, lo mismo que la P.
roberti beebei, se encuentra en una variedad de huéspedes, por ejemplo, Sigmodon
hispidus hirsutus, Akodon urichi urichi y Heteromys a. anomalus, entre otros.
Polygenis klagesi samuelis (Jordan y Rothschild, 1923)
S610 se tomaron muy pocos ejemplares, de cuatro huéspedes distintos.
Polygenis steganus (Jordan y Rothschild, 1923)
En la colección esta especie está representada por dos machos tomados de
Didelphis
m. marsupialis.
Polygenis peronis (Jordan y Rothschild, 1923)
Se comunica por primera vez desde la descripción original, y es al parecer un
parásito bastante característico del Heteromys a. anomalus.
Rhopalopsyllus
australis australis (Rothschild, 1904)
Se tomó un macho del Akodon u. urichi.
Rhopalopsyllus
cacicus saevus Jordan y Rothschild,
Se tomó un macho del Didelphis m. marsupialis.
1923.
Polygenis ambersoni sp. nov.
Parecida a la P. pero& (Jordan y Rothschild, 1923) y a la P. acodontis
(Jordan y Rothschild, 1923). Puede diferenciarse fácilmente de éstas por llevar
el noveno esterno del macho (fig. 4) cerdas subapicales muy largas, más largas
que el brazo dista1 mismo, en lugar de que esas cerdas ~610tengan como la mitad
de la longitud del brazo dista1 (fig. 5, P. peronis). Se diferencia ademas del P.
peronis en la siguiente forma: los palpos maxilares con el segundo segmento son
más de una y media veces más largos que los del primero (fig. 1, MX.P.),
no
sc510ligeramente más largos (fig. 7, MX.P.) ; cerda acetabular grande que penetra
bien arriba del acetábulo (fig. 13, ACB. y AC.) (como en la P. acodontis) en
vez de aproximadamente al mismo nivel (fig. 15, AC.B. y AC.); el brazo dista1
.
t-
Febrero 1962]
I
,
4
PULGAS T’ENEZOLANAS
135
del noveno esterno lleva en la línea media una fila submarginal de cerdas (fig.
4) que falta en la P. peronis (fig. 5) ; armadura de croché (fig. 10, CB.A.) bien
desarrollada, apareciendo en cada lado como un par de escleritas subcontiguas
parecidas a bastoncillos, en vez de estar tan reducidas que tengan el aspecto de
sola fila delgada (fig. ll, CKA.) ; cabeza del espermaducto (fig. 14) con una ligera
protuberancia dorsal, y de una longitud aproximadamente de una y media veces
la anchura, en vez de ser dos veces más larga que ancha (fig. 17). Puede diferenciarse de la P. acodo& por ser bastante ancho el ápice del brazo dista1 del
noveno esterno (fig. 4), y no angosto; esta estructura con la región subapical es
casi tan ancha como la porción media del digitoide (fig. 13, P.), no marcadamente
más angosta que F. La cabeza del espermaducto es más angosta que en la P.
acodontis, en la que es ligeramente mas larga que ancha.
El estudio del material brasileño y venezolano indica que la Polygenis tipopsis
Gkimurães, 1948, es una subespecie de P. roberli (Rothschild, 1905). Se considera
que la forma descrita previamente como Rhopalopsyllus beebei (I. Fox, 1947)
es también una subespecie de P. TobeTti. Estas conclusiones se basan en la
semejanza de detalles de las escleritas especializadas de los edeagos y de los otros
segmentos modificados del macho (con excepción del noveno esterno). La
quetotaxia de la metatibia, empleada como rasgo distintivo en las descripciones
primitivas de tripopsis y beebei,resultb más variable y no justifica la retenci6n
de estas formas como especies completas. Por ejemplo, los ejemplares venezolanos
de una región exhiben todas las variaciones de la tibia mencionadas.
Las tres subespecies pueden diferenciarse tomando por base la forma del
noveno esterno del macho. En la P. TobeTtikipopsis (fig. 29) el brazo dista1 tiene
casi la misma anchura desde la base hasta el ápice, con ápice casi redondo. En
la P. roberti beebei(fig. 30) el brazo dista1 se caracteriza por ser bastante convexo
en el margen ventral (posterior) ; ápice casi redondo, y el brazo ligeramente encorvado hacia arriba. La P. Toberti roberti (fig. 31) se diferencia en que el brazo
dista1 es mas angosto en la base que en el &pice redondo.
Se vuelve a describir e ilustrar (figs. 18-28 y 30) la especie Polygenis Tobertz
beebei(1. Fox, 1947) por ser muy comti y tener importancia potencial en la
epidemiología de la peste.
La Polygenis b. bohlsi (Wagner, 1901) merece también más estudio como vector
potencial de la peste, por lo menos entre los roedores. Aunque esta especie ha sido
mencionada con frecuencia en la literatura, no se ha ilustrado en forma adecuada.
La P. b. bohZsise diferencia con facilidad de las otras del género por lo siguiente:
el digitoide, o dedo movible (fig. 33, F.), tiene poco mas o menos la forma de un
triángulo isósceles; el brazo dista1 del noveno esterno (fig. 35) lleva en el ápice una
placa espesa de cerdas espiniformes; borde del edeago (fig. 32, FNR.) en fornia
de semiluna; espermaducto (fig. 34) de forma muy caracterfstica, pues la anchura
de la cabeza es de cuatro quintas partes de la longitud y es subtriangular; la
cola es mucho más corta que la cabeza y algo encorvada hacia arriba; el octavo
térgum de la hembra tiene en el medio una placa como de ocho cerdas delgadas
en la porción anterior a la fila de cerdas que se extiende desde el margen ventral
hasta el dorsal.