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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.

Descrz@~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.

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