THREE NEW SPECIES OF FROM CANADA (SIPHONAPTERA) G.P. HOLLAND Biosystematics Research Institute, Agriculture Canada, Ottawa KIA OC6

Abstract Can. Ent. 111: 713-719(1979) Three new species of fleas are described, two of the family Tiraboschi 1904 (Nearctopsylla grahami n. sp. and Rhadinopsylla (Micropsylla) rauschi n. sp.) and one of the family Dampf 1908 ( cald- erwoodi n. sp.). N. grahami is known only from western Ontario where it has been collected from marten, Martes a, americana. R. (M.) rauschi was collected from deer mlce, Peromyscus maniculatus osgoodi, in southwestern Saskatchewan. C. calderwoodi was collected from a nest of cliff swallow, Petrochelidon pyrrhonota, in eastern New Bmnswick.

HYSTRICHOPSYLLIDAE Tiraboschi 1904 Nearctopsylla grahami n. sp. Figs. 1, 2, 3, 5 Resembling N. brooksi (Roths.) and like that species found on mustelids, especially Martes americana . Male. Head much as in N. brooksi but pronotum with about 16-19 longer spines rather than 25-29 shorter ones (Fig. 1). Mesonotum and metanotum with about 3 irregular rows of setae with about 2 stout pseudosetae high up on each side of the mesonotum and none on the metanotum (Fig. 3) (vs. about 5 rows on the mesonotum and 4 on the metanotum in brooksi) and about 4 long slender pesudosetae per side on the mesonotum and 2 shorter ones, high up, on the metanotum (Fig. 4). Mesepisternum with about 6 setae per side; (brooksi with 9- 11). Metepimeron with 15- 17 setae per side. Abdominal tergum I with 3 rows of setae; 11-VI with 2 rows, and VII with 1 row. Two or three setae below each (circular) abdominal spiracle. (In brooksi, tergum I with about 4 rows, I1 and 111 with about 3 rows.) Apical spinelets on abdominal terga I (2-4); I1 (2, 4); 111 (2); IV (0-2). Single antesensilial seta, not on a posterior protuberance of tergum VII; in brooksi the slightly shorter seta is located on such a protuberance. Stigma of tergum VIII somewhat larger than that of brooksi. Clasper shaped much as in brooksi but the movable process broader (Fig. 2) and the patch of mesal setae sparser. Acetabular seta inserted on a level with the acetabulum rather than below it. Sternum IX very much as in brooksi with the posterior half of the ventral arm bent sharply dorsad, though shorter and broader than in that species and without the slight terminal anterior bending. Groups of long and short setae disposed very much as in brooksi, the row of about 20 setae along the dorsal half of the ventral arm being somewhat longer (Fig. 2). Female. Head similar to the male but with the usual sexual differences. Pronotum with 2 irregular rows of setae and 15-17 spines in the ctenidium (vs. 18-23 in brooksi). Mesonotum and metanotum as in the male, there being about 2 stout pesudosetae a side, high up, on the mesonotum; metanotum lacking pseudosetae (cf. with brooksi, which has them). Mese- pisternum with about 6 setae per side; brooksi with 9-1 1. Metepimeron with about 16 setae per side. Hind tibia with 8 pairs or triplets of dorsal setae set in notches and about 19 lateral setae. Metacoxa with about 12 spiniforms arranged mesally, near the anterior apex. Abdominal terga with about 3 irregular rows of setae becoming 2 around terga VI and VII. Two subequal antesensilial setae as in brooksi. Abdominal apical spinelets as follows: I, 3(2-4); 11, 4(3-5); 111, 3(2-5); IV, l(2); V, 0; VI, 0 (but one specimen with a large spinelet on one side). The spinelets of N. brooksi are very much the same. Two or three long setae of the distal row of each tergum extending below the spiracle. Abdominal sterna 111-VI with about 8 setae in 2 rows of which the distal 6 are long. About 16 setae on sternum VII, in 2 rows, of which the distal 9 are long. The posterior margin of sternum VII has a rounded lobe and a broad, deep, sinus (Fig. 5). Tergum VIII with about 6 long setae and about 12 shorter ones on its distal ventral surface. Anal stylet about 6 times as long as broad and with 1 long apical seta and 2 small hairs on either side of the base of the seta. Spermatheca with a sub-globular bulga and with the hilla tapering gradually and curled around 714 THE CANADIAN EM0MOLOC;IST Volume 11 1 THE CANADIAN ENTOMOLOGIST 715

it (Fig. 5). Bursa copulatrix almost straight and with the long central portion heavily sclerotized. Holotype 6, Spruce River Road, mile 64, Thunder Bay District, Ontario, 20.XII.77, ex juvenile Martes americana americana, collected by Bob Graham. No. 15894 in the Canadian National Collection of , Ottawa. Allotype 9, same data. Paratypes, 266 and 12 9 9 as follows: same data as holotype and allotype, 16, 2 9 9; Mile 74, Spruce River Road, 5.XI.77, ex M. a. americana, 3 9 9; same data, but 12.XI.77, 3 9 9; same data but Mile 88, 26.XII.77, ex female M. a. americana, 19; same data but Mile 68, 21.11.78, ex male M. a. americana, 16, 19; same data but Mile 78, 12.11.78, ex female M. a. americana, 19; same data but mile 90, 12.11.78, ex male M. a. americana, 19. All the specimens were collected by Mr. Graham but were communicated by Prof. M.W. Lankester of Lakehead University, Thunder Bay, to whom I extend my gratitude. Besides the 16 specimens of Nearctopsylla grahami just described, Mr. Graham collected specimens of the following species of fleas from the 13 specimens of Martes a. americana that he got in his traps along the Spruce River Road: Hystri- chopsylla dippiei ssp., 4 specimens; Rhadinopsylla (Actenophthalmus)fraterna (Baker), 1; Monopsyllus vison (Baker), 1; Tarsopsylla octodecimdentata coloradensis (Baker), 1; Orchopeas caedens durus (Jordan), 99; and arctomys (Baker), 6. From the fleas caught one would judge that the principal prey species of the martens were red squirrels, Tamiasciurus hudsonicus, and woodchucks, Marmota monax.

Rhadinopsylla (Micropsylla) sp. Micropsylla is at present set off as a subgenus of Rhadinopsylla principally on the basis of the absence of a transverse ridge dividing the metepisternum from the metanotum. The subgenus is only distributed in western North America (Rhadinop- sylla is holarctic) and two forms have been described, separated as subspecies. Rhadinopsylla (Micropsylla) s. sectilis Jordan and Rothschild 1923 was described from Kelowna, B.C., and has since been reported from various points in Washington, Montana, Nevada, Oregon, California, and Utah. It is characterized by having a genal ctenidium of 4 spines and male genitalia like Fig. 8, drawn from a topotype. R. (M.) s. goodi (Hubbard, 1941), described from Forest Grove, Oregon, usually has 5 genal spines and the male with the movable process somewhat shorter than in typical sectilis. It has since been taken from Vancouver, B.C., western Oregon and California, and apparently is restricted to west of the Cascades, whereas R. (M.)s. sectilis occurs east of those mountains.

LIST OF ABBREVIATIONSUSED IN ILLUSTRATIONS.Ac. S., acetabular setae; Ant. S., antesensilial setae; B. Cop., bursa copulatrix; Bu., bulla (head) of spermatheca; CI., body of clasper; C. Scl., cervical sclerite; D. o., ductus obturatus; D. Sp., ductus spermathecae; Ephy., epipharynx; F., movable process of clasper; Fv., fovea; Fr. Tb., frontal tubercle; G. Ct., genal ctenidium; GI. W., Wagner's gland; HI., hilla (tail) of spermatheca; Lb. Plp., labial palpus; M., manubrium of clasper; Mb., membranous appendage of sternum VIII; Ms. Epm., mesepimeron; Ms. Eps., mesepisternum; Mt. Eps. metepisternum; Mt. Epm., metepimeron; Mt. N., metanotum; Mx., maxilla; Mx. LC., maxillary lacinia; Mx. Plp., maxillary palpus; Oc., eye; P., fixed process of clasper; Pap., papilla; Pr. N. Ct., pronotal ctenidium; Ps., pseudosetae; Sens., sensilium; Sn., sinus in apical margin of sternum VII; St., abdominal sternum; Stig., stigma, the spiracular opening of tergum VIII; Str., striated area of metepi- meron; Strig., strigillae, chitinous thickenings in the spermatheca; Spic., spiculose area of tergum VIII; Spl., spinelets; Spr., spirzle; T., abdominal tergum; I, 11, 111, IV, etc., segmental numbers.

FIGS. 1-5. 1-3, 5, Nearctop~yllagrahami n, sp.: 1, head and pronoturn of male; 2, sternum IX of male and clasper with fixed and movable processes; 3, metanotum and abdominal tergum' I of female; 5, sternum VII and spermatheca of female. 4, Nearctopsylla brooksi (Rothschild): metanotum showing pseudosetae, and tergum I of female, from Atlin, B.C., ex Mustela erminea. 716 THE CANADIAN ENTOMOLOGIST June 1979 The new species described below was discovered in southwest Saskatchewan and has a genal comb like that of R. (M.) s. goodi of the Pacific Coast strip, but has larger, and differently shaped, genitalia.

Rhadinopsylla (Micropsylla) rauschi n. sp. Figs. 6, 7 Male. Head with frontal tubercle about half way down the frontal margin. Ocular row of 3 long setae and frontal row of about 5 shorter ones. Genal comb of 5 spines (Fig. 6). Eye vestigial as is usual in this . Post antenna1 region with 3 rows of setae of which the first two rows are oblique. Labial palpus of 5 segments and just attaining apex of fore coxa. Pronotum with about 5 long spines on a side and a comb of 16 spines altogether. Mesonotum and metanotum each with 2 rows of setae of which the second row consists of 5 long setae with small intercalaries between them. Metanotum extending downward where it is fused, without a suture, to the metepisternum which has 2 more setae. Mesonotum with Volume 11 1 THE CANADIAN ENTOMOLOGIST 717 about 2 long pesudosetae per side; metanotum with none. Metepimeron with 5 or 6 long setae as shown and a densely striated area on the ventro-lateral surface. Other thoracic setae distributed as shown (Fig. 6). Hind coxa with a dense patch of 12T short setae on the antero- distal mesal surface. A few other setae distally on the outer surface. Six single setae or pairs of stout setae set in notches along the dorsal edge of the hind femur. Distal tarsomere with 4 pairs of plantar setae. Abdominal terga each with one major row of about 5 long setae, with intercalaries, and a minor row of about 2 slight setae. Single apical spinelets on the first 4 or 5 terga; sometimes on only 3. A long seta of the major row just below the spiracles on T.11-VII. Body of clasper somewhat broader than in Rhadinopsylla (Micropsylla) sectilis ssp, and the movable process longer and less pointed (Fig. 7). Sternum IX differing from sectilis in that the ventral arm is apically broad and rounded rather than relatively narrow and pointed (cf. Figs. 7 and 8). Holotype 6, Cypress Lake, south of Cypress Hills, Saskatchewan, 24-26.X.76, ex Peromyscus maniculatus osgoodi, collected by Dr. Robert L. Rausch. No. 15895 in the Canadian National Collection of Insects, Ottawa. Paratypes, 266, as follows: same data as holotype, 1 6; near E. end of Cypress Lake, Saskatchewan, 13.XI.77, ex Peromyscus m. osgoodi, 16, collected by Eric Hoberg. The female is unknown. The species is named after Prof. Robert Rausch who, over the years, has collected many species of fleas for me, from Alaska, Saskatchewan, south America and the U.S.S.R.

CERATOPHYLLIDAE Dampf 1908 Ceratophyllus (Ceratophyllus) calderwoodi n. sp. Figs. 9, 10, 11, 12 A typical swallow-feeding Ceratophyllus with all the characteristic features of this portion of the genus, but differing only in the genitalia, especially that of the male. Male. Head of the usual shape, armed with a sharp frontal tubercle and with 2 rows of setae before the eye. Eye large. Post antennal region with 2 setae disposed along the upper side of the antennal groove and about 22 small hair-like setae along the length of the groove. About 5 setae along the posterior edge of the head. Labial palpus of 5 segments nearly, but not quite, as long as the fore coxa. Pronotal ctenidium of about 14 spines on each side. In front of the ctenidium, a single row of about 6 setae with small intercalary hairs. Mesonotum with a strong row of about 5 large setae and about 20 small setae, some even hair-like, scattered in about 3 or more rows on the dorsal part of the sclerite. About 6 pseudosetae per side. Mesepisternum with about 10 setae most of which are thin and hair-like, only two being long. Mesepimeron with 3 setae. Metanotum with a posterior row of 5 long setae, and an irregular anterior row of 5 or 6. Metanotum with a single short pseudoseta per side. Metepisternum with only 1 strong seta. Metepimeron with about 6. Pleural arch strongly developed. Hind tibia with 6 notches, each bearing a stout seta or a pair of stout setae, on its dorsal margin. About 6 thin setae in a row on its dorso-lateral surface. Abdominal tergum I with 2 rows of about 5 setae each and I or 2 scattered setae anteriorly. The first two spiracles appear doubled-up; the others, 111-VII, are more or less circular. There are 2 apical spinelets on each side on terga I-IV. Terga 11-VI bear a posterior row of 7 long setae, with intercalary setae, and an anterior row of about 3 or 4 shorter ones and anterior to this, another seta. Tergum VII has 6 long setae in the posterior row, the 7th being replaced by the single antesensilial seta, flanked by 2 tiny hairs. Tergum VIII with about 8 long setae in a dorsal marginal row, and about 5 scattered laterally (Fig. 9). Dorsal part of the tergum beset mesally with a dense patch of spicules. Sterna 11-VII armed ventrally with 2-4 setae per side. Sternum VIII with conspicuous Wagner's gland proximally. Distally the sclerite is more or less parallel-sided with about 20 minute setae along its length and terminating in a point apically with about 3 long setae on a side and a long apical membranous flap which is distally fimbriated. THE CANADIAN ENTOMOLOGIST Volume 1 I I TFlE CANADIAN ENTOMOLOGIST 719

Clasper and processes much as in C. c. celsus Jordan to which it is most closely allied; but the fixed process is far longer than in celsus and the pit opposite the fovea of the movable process is situated at its distal apex, thus changing the contour of the movable process (Fig. 10). Fixed process exceeding the upper of two long setae on the movable process (in celsus the process only exceeds the lower of the corresponding two). Other setae distributed approximately as shown. In Ceratophyllus scopulorum Holland the fixed process is longer again, greatly exceeding both long setae of the movable process. Sternum IX very much as in C. celsus but differing from C.scopulorum in that the proximo-ventral lobe is conspicuously narrower. Acetabular setae inserted slightly below the level of the point of articulation of the movable process. Female. The head is similar to that of the male except for the usual sexual differences. Terminal setae on antenna1 segment I1 exceeding club. Pronotal ctenidium with about 14 spines per side. Setae and pseudosetae as in the male except for the anterior rows on the mesonotum and the metanotum which are somewhat fewer. Abdominal tergum I and spiracles I-VII as in the male. Apical spinelets, 2 on terga 1-111, and usually I on tergum IV, Terga 11-VI bear a posterior row of about 7 long setae with intercalary setae and an anterior row of up to 6 shorter ones and anterior to this, another seta. Tergum VII has a posterior row of 6, an anterior row of about 3 shorter setae, and 3 antesensilial setae of which the middle one is long flanked by a short seta dorsally and a much shorter one ventrally. Tergum VIII with about 7 short setae guarding the spiracle dorsally, about 12 longer setae from near the sensilium, marginally, to the ventral edge of the sclerite, and about 13 grouped ventro-laterally. A patch of about 4 stout setae mesally, just below the stylet. Anal stylet about 3 times as long as broad, and armed with a strong seta apically and 2 smaller setae subapically. Abdominal sternum I1 with 1 seta submarginally and ventrally. Sterna 111-VI with about 4 setae in a row, submarginally and ventrally. Sternum VII usually without a sinus (Fig. 11) but a few specimens with (Fig. 12). About 7 submarginal long setae and just anterior to these about 7 shorter ones. Spermatheca of the usual type found in most species of Ceratophyllus, with a long sausage-shaped (4:l) bulga with con- spicuous strigillae, and a shorter hilla and some strigillae, and with a sclerotized apical papilla. Ductus spermathecae extremely long. A portion of the wall of the bursa copulatrix sclerotized and also the basal portion of the ductus obturatus. Holotype d, Kouchibouguac National Park, New Brunswick, 7.1X.77, ex nest of Petrochelidon pyrrhonota, collected by Gordon Calderwood after whom the species is named. No. 15896 in the Canadian National Collection of Insects, Ottawa. Allotype, 9, same data. Paratypes, 51 d d and 55 9 9, same data. Petrochelidon pyrrhonota, the cliff swallow, is remarkable, even for a swallow, in that it has at least five species of fleas associated with it, C. c. celsus Jordan, C . petrochelidoni Wagner, C . scopulorum Holland, C . arcuegens Holland, and now C. calderwoodi Holland. No other swallow has so many although Riparia riparia, the bank swallow, shares C. c. celsus along with C.r. riparius Jordan and Rothschild, which is specific to it. The tree swallow, Iridoprocne bicolor, has C. idius Jordan and Rothschild which it shares with the purple martin, Progne subis. So far no characteristic has been discovered in the nests of the barn swallow, Hirundo erythrogaster, the violet-green swallow, Tachycineta thalassina, or the rough-winged swallow, Stelgidopteryx ruficollis. Swallow-fleas, when present, are often in large numbers and generally of one species. Exceptions are C. r. riparius and C. c. celsus which sometimes occur together in nests of bank swallows and C. scopulorum and c. arcuegens which occur together in nests of cliff swallows in the north. Though C. scopulorum has been recorded from Kent Island, New Brunswick, C. arcuegens has not. (Received 14 September 1978)

FIGS. 9-12. Ceratophyllus calderwoodi n. sp. 9, genitalia of male. 10, detail of fixed and movable processes. 11, sternum VII and spermatheca, ducts, and bursa copulatrix. 12, sternum VII with distal margin incised.