a taxonomic review of the ceratophyllus curtis, 1832 of north … · 2017-10-11 · december, 2001...

December, 2001 Journal of Vector Ecology 119

A taxonomic review of the Ceratophyllus Curtis, 1832 of North America(Siphonaptera: Ceratophyllidae: Ceratophyllinae)

Robert E. Lewis1 and Terry D. Galloway2

13906 Stone Brooke Circle, Ames, IA 50010-41742Department of Entomology, University of Manitoba, Winnipeg, Manitoba, Canada R3T 2N2

Received 8 January 2001; Accepted 15 March 2001

ABSTRACT: The 22 North American taxa assigned to the flea genus Ceratophyllus Curtis, 1832, are reviewed.All are parasites of birds and include 17 species in the nominate subgenus, 3 in Emmareus and 2 in Celeophilus.Keys to the subgenera are provided, distinguishing characters of the individual species are discussed and illustratedand the various aspects of distribution and host preferences are cited. Journal of Vector Ecology 26 (2): 119-161. 2001.

Keyword Index: Ceratophyllus, taxonomy, distribution, hosts.

The genus Ceratophyllus has historically containedspecies now assigned to many other families. Untilthe 1930’s no serious effort was made to separate thesetaxa into their respective genera for various reasons.Although workers at the British Museum (NaturalHistory) at Tring had long since divided the myriadspecies assigned here into groups of species equivalentto genera, their classification had never been formallypublished. By the early 1930’s it was obvious that arevised interpretation of the genus was necessary toaccommodate the influx of new species being described.In his survey of the American species of Ceratophylluss. lat., Jordan (1933) stated: “If I here inflict on sciencea number of new generic names, I plead the excuse thatthe divisions of Ceratophyllus I here define wouldinevitably be named by somebody else, probably bysomeone who does not know these fleas, as has happenedin other orders of animals.” He then discussed thecomposition of 18 genera of ceratophyllids, 8 of whichbore his new name designations. In this discussion, hedivided the genera into 3 groups based on the chaetotaxyof the forefemur and mid- and hindcoxae. Ceratophyllushe placed in group C, in which the outer surface of theforefemur bears a number of small lateral bristles, andthere are no long, thin bristles on the basal half of theinner surface of the mid- and hindcoxae. Wagner (1934)followed by erecting Citellophilus for a group of speciesassociated with ground squirrels in the PalaearcticRegion. However, the first major effort at perfecting aceratophyllid classification was that of Ioff (1936).Although the entire classification suffered for want ofstrong representation of North American genera andspecies, Ioff did divide the Ceratophyllus species

available to him into 3 infrafamilial groups. Here thematter remained for the next 45 years. During thisperiod G. H. E. Hopkins constructed a key to theceratophyllid genera known to him, but it also was neverpublished. Goncharov (1981) discussed theinfrafamilial classification of both the Leptopsyllidaeand Ceratophyllidae, but only listed Ceratophyllus andone of the 15 genera he placed in the tribe CeratophylliniDampf, 1908, as 1 of 7 tribes assigned to the subfamilyCeratophyllinae Dampf, 1908. He went on to say thatperhaps this tribe could be further divided into severalgroups, of which he listed five. Ceratophyllus s. str.alone made up one of these groups.

Perhaps the greatest contribution to theclassification of the Ceratophyllidae to date is that ofthe late Frans Smit. The family was discussed in detailin the introductory chapter of Traub et al. (1983),comparing it to the closely related Leptopsyllidae andproviding a key to the ceratophyllid genera andsubgenera of the world. Eight new genera and 9 newsubgenera were erected or exhumed in this work andsome existing genera were reduced to subgenera of eitherCeratophyllus or Oropsylla. Ceratophyllus itselfsustained considerable modification. Two newsubgenera, Celeophilus and Emmareus were erected forsome species infesting birds that were originally placedin the nominate subgenus. Amonopsyllus Wagner, 1938,as a subgenus, was resurrected for the 4 subspecies ofMonopsyllus ciliatus (Baker, 1904). The genusMonopsyllus Kolenati, 1857, was placed inCeratophyllus as a subgenus, and Rosickyiana Smit,1972, was retained as a subgenus.

While a case can be made for the retention of the

120 Journal of Vector Ecology December, 2001

subgenera Ceratophyllus, Celeophilus and Emmareus,since they are all bird fleas and equate with distinctspecies groups within the genus, the same cannot besaid of Amonopsyllus, Monopsyllus and Rosickyiana.The single species belonging to Amonopsyllus is aparasite of arborial and scansorial sciurids in westernNorth America. The 1 North American species ofMonopsyllus and the monotypic Rosickyiana are bothprimarily parasites of mustelid carnivores. For theseand other reasons we are retaining these 3 as full generaand they will be discussed in a larger work in progress.

The genus Ceratophyllus shares the distinction witha few other taxa of fleas in that its species are associatedwith birds and many of these are hosts to more than 1member of the genus simultaneously. Thus, theEuropean house martin, Delichon urbica, a broad-ranging Palaearctic swallow, may be infested with C.farreni, hirundinis and rusticus at the same time. InNorth America the cliff swallow Petrochelidonpyrrhonota, hosts C. arcuegens, calderwoodi, celsus,petrochelidoni and scopulorum in different parts of itsrange and the distributions of some of these species areat least partly sympatric. Therefore, as populations offleas increase in the nesting sites there is ampleopportunity for attempted mating of one species withanother. The latest mention of this is that of Benton(1998) but the possibility has been alluded to by earlierauthors such as Rothschild (1955). Further commenton this subject may be found in the species accounts ofC. calderwoodi, celsus and idius.

To our knowledge only C. gallinae has been rearedin the laboratory under controlled conditions as reportedby Vysotskaya & Kir’yakova (1970) and this wouldappear to be fertile ground for further experimentation.

Lewis (1990) listed 48 species in the world faunain the 3 subgenera considered here, and 3 additionalspecies have been described since. Of the 22 speciesreported from North America, 17 belong to the nominatesubgenus, 2 to Celeophilus and 3 to Emmareus.

Before embarking on a taxonomic resumé of theNorth American species of Ceratophyllus, anexplanation of the synonymy section of the individualspecies accounts is in order. Five of the 22 species areconsidered to be subspecies of taxa that are found inother parts of the Northern Hemisphere. There are also19 junior synonyms assigned to these taxa, albeit 15 ofthese apply to C. gallinae and C. garei. There are 7species names and 3 subspecies names that are treatedas junior synonyms in this study. Both junior synonymsand the names of North American subspecies areincluded in the synonymy section of each species.Unless information concerning the actualsynonymization is included in the citation, the name is

not formally considered to be a synonym in this work.That is, it is not formally synonymized.

In a recent review of the 2581 names of species-group taxa in the Siphonaptera of the world that werestill treated as valid in July of 2000, 576, or slightlyover 22% were named subspecies. When compared toother orders of insects it would seem that eitherpulicologists are among the most perspicacious oftaxonomists, or the subspecies category is being usedas a repository for speculation. We are firmly convincedof the latter. So few subspecies complexes have beenanalyzed, and the results so tentative, that there is littleevidence of the validity of the subspecies concept in theorder. As a result, subspecies are acknowledgednomenclaturally in the individual treatments withoutdebating the desirability of such taxonomic application.

Host names in the following accounts conform withusage in the AOL Check-list (1998) for birds and Wilsonand Reeder (1992) for mammals.

Ceratophyllus Curtis, 1832

Ceratophyllus. Curtis, 1832, British Entomology 9(417). Generitype: Pulex hirundinis Curtis, 1826.

Ceratopsyllus. Curtis, 1832, British Entomology 15 (nopage number) errata & addenda. An unwarrantedemendation.

Key to the subgenera of Ceratophyllus(modified from Smit, 1983)

1 Occipital setal row not widely interrupted abovethe ventralmost seta; spiculose area of male t VIII poorlyto well developed .................................................... 2 Occipital setal row widely interrupted above theventralmost seta; spiculose area absent (onwoodpeckers) ...................................... Celeophilus2 Apophysis of st IX and penis rods of male making atleast one convolution; in females the basal portion ofthe spermathecal duct is not strongly dilated or wrinkled;basal portion of the ductus obturatus strongly sclerotized;bulga of spermatheca cylindrical, much longer than hilla;spermathecal duct usually inserted in the direction ofthe long axis of the bulga (mostly on birds buildingrelatively dry nests above ground level, especiallyhirundinids) ...................................... Ceratophyllus Apophysis of st IX and penis rods making at most oneconvolution; in females the basal part of the spermathecalduct is dilated and wrinkled; ductus obturatus usuallyweakly sclerotized through-out; bulga swollen, aboutas long as hilla; spermathecal duct inserted near the endof the dorsal side of the bulga, forming an angle of atleast 45° with the latter’s long axis (birds with relatively


damp nests on or near the ground) ....... Emmareus

Subgenus Celeophilus Smit, 1983

Ceratophyllus Curtis, 1832 - Celeophilus subgen. n.Smit, 1983: 14.

Ceratophyllus (Celeophilus) Smit, 1983. Smit, 1983:30.

Diagnostic CharactersThe subgeneritype for this subgenus is

Ceratophyllus (Celeophilus) adustus Jordan, 1932.Head Frontal tubercle present but extremely small.

Eye well developed and heavily pigmented, oval.Frontal setal row of 0 to 3 setae. Ocular row of 3 setaein both sexes. Postocular seta present, extending wellbeyond the apex of the genal lobe. When not broken,all or most of the setae on the antennal pedicel extendbeyond the apex of the clavus in females. Preoccipitalsetae usually 1 in addition to a row of minute setulaeextending along the dorsal margin of the antennal fossa.Occipital setal row usually 4 per side, widely interruptedabove the ventralmost, separated from the upper 3 by awide space. Labial palpi extending about 3/4 the lengthof the forecoxa.

Thorax Pronotal ctenidium of 28-30 spines.Mesepisternum with a few small setae; mesepimeronwith only 1 or 2 slender bristles. Caudal margin ofmetanotum usually without spinelets, or if present theyare minute.

Legs Forecoxa sparsely setose. Sulcus ofmidcoxa with a short dorsal interruption. Forefemurwith small setae on its lateral surface. Striation of mid-and hindfemur and tibia of medium density. All tibiaewith 6 notches in the caudal margin bearing heavy setae.Chaetotaxy of the hindtibia in members of this subgenusreduced (Figures 1, 2). First and third pairs of lateralplantar setae slightly displaced on to the plantar surface.

Abdomen Abdominal tergites I-IV usually with 1or 2 marginal spinelets per side. Outer and inner of the3 antepygidial setae minute in the male, somewhat longerin the female. Remaining characters are sexual in nature.

Male Wagner’s organ present. Apodeme of st VIIIand penis rods coiled ~1 convolution. Apex of penisplate with a long tendril that is also coiled ~360°.Antepygidial setae 1 per side bordered dorsally andlaterally by 1 minute setula. Tergum VIII lacking aspiculose area along its dorsocephalic margin. SterniteVIII gently bent dorsad, its apex with a membranousvexillum. Movable process of the clasper long andnarrow. The 2 acetabular setae arising about midwaydown the caudal margin of the fixed process.

Female Three antepygidial setae, the median the

longest, 5X the length of the ventral, ~3.3 the length ofthe dorsal. Genital ducts weakly sclerotized, ductusbursae and ductus obturatus rather short. Base ofspermathecal duct wrinkled and somewhat dilated.Bulga rather large, cylindrical, with an apical orifice forthe duct. Hilla much shorter than bulga and set off fromit by a prominent constriction. Anal stylet normally witha dorsal as well as a ventral lateral seta.

RemarksThis is a small subgenus with only 2 known species,

both of which occur in Traub’s (1983) Forested andNorth Canadian Sector of the Nearctic Region and theSiberian/Northeastern Sector of the Palaearctic Region,where they are evidently parasites of woodpeckers. Themale of C. (C.) adustus Jordan, 1932, has just beendescribed by Lewis and Haas (2001).

Key to the species of Ceratophyllus (Celeophilus)

1 Movable process of male clasper ~3.5X as long aswide at its widest point; apex of apical arm of male stVIII with an apical fimbriated cone subtended by a fewlong setae and a fimbriated vexillum; apical arm of stIX of male with parallel margins terminating in arounded apex; caudal margin of female st VII entire,without a sinus (Figure 76) ......................... adustus

Movable process of male clasper >4X as long aswide at its widest point; apex of apical arm of male stVIII with a subapical bladder-like vexillum and 2 longsetae; apex of male st IX with subparallel marginsconverging to a rounded tip; caudal margin of female stVII usually with at least a shallow sinus (Figure 77)

............................................................ zhovtyii

Ceratophyllus (Celeophilus) adustus Jordan, 1932(Figures 2, 8, 27, 69, 71, 76)

Ceratophyllus adustus. Jordan, 1932, Novit. Zool. 38:253, Figures 10-11. Canada,British Columbia,Atlin, 59.31N 133.41W, from Erethizon epixanthum[=dorsatum], 3.IX.1931, H. S. Swarth leg. Holotypefemale, by monotypy, British Museum (NaturalHistory) [BMNH].

Diagnostic CharactersMale One long antepygidial bristle per side

bounded above and below by a single tiny setula. TergumVII lacking an area spiculosa. Movable process ofclasper ~3.5 as long as wide at its widest point, about1/3 of its apex extending beyond apex of the fixedprocess. Apex of st VIII (Figure 69) bearing afimbriated, cone-shaped projection subtended by a few


long setae and a narrow, fimbriated vexillum. Distallobe of st IX broad basally (Figure 27) tapering apicallyto a rounded apex with parallel margins. Crochet(Figure 69) acutely pointed apically.

Female Three antepygidial setae per side, thelaterals much shorter than the medial but much longerthan those of the male. Seventh sternite (Figure 76)usually with a caudal lobe that is not subtended by ashallow sinus. Anal stylet slightly <2X as long as wideat base, sharply tapered apically and bearing 3 subapicalsetae in addition to the apical seta. Spermathecal bulga~1.5X as long as hilla, the latter usually with a welldefined apical papilla.

RemarksThis is the subgeneritype for the subgenus. As is

frequently the case with rare species, much of theliterature about it refers to previous references withoutadding materially to our knowledge. Originallydescribed from a single female from an unusual host fora member of this genus, additional material was notreported until Haas et al. (1980), 1 female, and Haas &Wilson, (1984), 66 males, 102 females. Thesecollections came from various localities in Alaska. It isnow beyond doubt that this species is not a parasite ofporcupines, as originally thought, but rather is awoodpecker flea infesting the nests of the flicker,Colaptes auratus, the three-toed woodpecker, Picoidestridactylus, and other members of this genus in theextreme northwestern part of North America. So farthis flea has only been collected in Alaska but it mightbe expected to occur in the northern parts of some ofthe Canadian provinces as well.

In his treatment of morphological anomalies in fleas,Haas (1983) described the presence of a vestigial secondspermatheca in 5 of the 103 females examined by him.While such anomalies are uncommon in ceratophyllidadults, they are not unknown, and over 25% of femaleNosopsyllus medus Jordan, 1938, from Iraq in the LewisCollection have similar vestigial second spermathecae.

Distribution & Host AssociationsMaterial upon which this account is based was all

taken from a few localities in Alaska. With respect tohost associations, only the flicker and the three-toedwoodpecker have been identified with certainty.However, as Haas & Wilson (1984) pointed out, thedowny (Picoides pubescens), the hairy (Picoidesvillosus) and the black-backed (Picoides arcticus)woodpeckers also occur in the area where these fleaswere collected. Of these, the former 2 species wereconsidered to be the most likely preferred hosts sincethe nests were in dead tree stubs of birch, poplar and

spruce, and the latter species of woodpecker prefers tonest in living trees.Ceratophyllus (Celeophilus) zhovtyii Yemel’yanova

& Goncharov, 1966(Figures 1, 9, 45, 70, 72, 77, 85)

Ceratophyllus zhovtyi. Yemel’yanova & Goncharov,1966, Izv. Irkutsk. Gos. Nauch-Issled. Protiv. Inst.Sibiri Dal’n Vost. 26: 309, Figures. 1-4. USSR,Irkutskaya Oblast, Cheremkhovskiy rayon, environsof Sakhir, from Glaucidium passerinum, 21.X.1962,V. D. Sonin leg. Holotype male, Irkutsk AntiplagueInstitute. [IAI]

Diagnostic CharactersMale One long antepygidial bristle per side,

bordered above and below by a single minute setula.Tergum VIII lacking an area spiculosa. Movable processof clasper (Figure 9) >4X as long as wide at its widestpoint, its apical 1/3 extending beyond the apex of thepointed fixed process. Apex of st VIII (Figure 70)bearing a few subapical setae and a fimbriated, bladder-like vexillum. Distal lobe of st IX (Figure 45) not sobroad basally, tapering to a rounded apex withsubparallel margins. Crochet (Figure 70) triangular, wellsclerotized.

Female Three antepygidial setae per side, thelaterals much shorter than the median but longer thanthose in the male. Anal stylet ~2X as long as wide atbase, tapering apically and with 1 apical and 2-3subapical setae. Seventh sternite (Figure 77) usuallywith a caudal lobe subtended by a shallow sinus. Bulgaof spermatheca cylindrical, ~1.5X as long as thepapillate hilla but not as distinctly set off from it as inthe preceding species.

RemarksLittle is known about this species. That it has finally

been collected in North America, albeit for the most partin the form of fragments, should stimulate an effort toexamine woodpecker nests in the northern part of thecontinent for additional material of better quality.

Distribution & Host AssociationsAs far as can be determined, the reference by Haas

& Wilson (1984) to this species is the only mention ofits occurrence in the Western Hemisphere. Their seriesof 28 specimens came from three woodpecker nests inthree different localities in Alaska. Two collectionscame from flicker nests and the third from the nest of aspecies of Picoides. Most of the specimens were deadwhen collected and they were taken in association withspecimens of C. adustus and C. rauschi.


This species was originally described from a smallseries collected from the Eurasian pygmy owl(Glaucidium passerinum) and a woodpecker, presumedto be Picoides major, in the Irkutsk region of the SovietUnion. As with many other bird fleas, its apparent rarityis probably due to the infrequency with which bird nestsare examined for fleas.

Subgenus Ceratophyllus Curtis, 1832Ceratophyllus (Ceratophyllus) Curtis. Smit, 1972:

16.

Diagnostic CharactersThe subgeneritype for this subgenus is

Ceratophyllus (Ceratophyllus) hirundinis (Curtis,1826), a species extralimital to this study.

Head Frontal tubercle present but inconspicuousin both sexes. Eye well developed and heavilypigmented. Frontal setal row of 5 or 6 setae extendingup the cephalic margin of the antennal fossa in males;usually only 4 in females and not extending so high.Ocular row of 3 setae in both sexes. Postocular setapresent but frequently obscured. In females there are 4or 5 setae arising on the antennal pedicel that extendwell beyond the apex of the clavus. There are 20 to 25small setulae along the dorsal margin of the antennalfossa in males. In females the average is ~20 and therow expands in width as it progresses caudad.Preoccipital setae usually 3 arranged in 2 oblique rowsof 1 and 2. The occipital setal row usually has 5 setaeper side in males. There are usually 7 setae per side infemales, the penultimate ventral seta by far the longestand usually accompanied by a much smaller seta arisingbelow it near its base. Labial palpi extending about 4/5the length of the forecoxa.

Thorax Pronotal ctenidium of 29 or 30 spines inmales, 30 to 32 in females. Usually 5 pseudosetae underthe mesonotal collar per side in both sexes.Mesepisternum usually with 1 long seta per side in males,1 or 2 in females. In the former there are 3 to 5 minutebristles along the cephalic margin, in the latter there maybe from 4 to 10. Usually 5 bristles per side on themesepimeron in both sexes arranged in 2 rows of 2 and3. Males have 1 minute marginal spinelet per side onthe metanotum; females may have 1 or 2 per side and inboth sexes they are smaller than those on the anteriorabdominal segments. There are 1 to 5 setae on the lateralmetanotal area in males, usually 3, females with 3 or 4per side. With only 1 seta per side on the metepisternumin both sexes. The number of setae on the metepimeronvaries. Typically there are 6 to 8 in males arranged in 3irregular rows. Females vary from 6 to 10 but mostfrequently there are 8, also arranged in 3 irregular rows.

Legs There is a small seta near the base of theforefemoral guard seta, 5 notches in the margin of theforetibia bearing heavy bristles, and 5 pairs of lateralplantar bristles of which pairs 1 and 3 are slightly shiftedon to the plantar surface. This condition prevails in bothsexes and all 3 pairs of legs. Midtibia with 5 notches inmales, 6 in females. Hindtibia with 5 notches in males,6 or 7 in females.

Abdomen Number of abdominal tergal spineletsper side variable but approximating: males I, 3; II, 3; III3; IV, 1 to 3; females, I, 2; II, 2 or 3; III, 1 to 3; IV, 0 or1. No lateral setae on st II in either sex. The remainingcharacters are sexual in nature.

Male Wagner’s organ present. Apodeme of st VIIIand penis rods coiled ~1.5 or more convolutions (exceptin C. enefdeae where they only complete about 360°).Apex of penis plate long and also coiled. Antepygidialsetae 3, the ventral the shortest, the dorsal ~2X as longand the middle, >2X the length of the dorsal. TergumVIII with a well developed spiculose area along its dorsalmargin in most species. Sternite VIII gently bent dorsad,its sides subparallel, its apex bearing 2 or 3 long setaeand a membranous vexillum of variable shape. Claspervariable. Anal tergite triangular. The 2 acetabularbristles arise about the level of the dorsal margin of theacetabulum.

Female Bursa copulatrix weakly sclerotized. Theapical papilla of the hilla is usually well developed andthe spermathecal duct of the bulga arises apically. Ofthe 3 antepygidial setae, the dorsal and lateral bristlesare nearly the same length, the upper a bit shorter, themedial ~2X its length. Anal stylet approximately 2X aslong as wide; with a long terminal bristle and shorterdorsal and ventral subapical bristles.

RemarksThis is by far the largest subgenus and of the 22

North American species of Ceratophyllus, 17 belonghere. Nine of these are parasites of hirundinids(swallows and martins), 3 more are probably parasitesof gulls and other sea birds, another is evidently aparasite of crows, and Ceratophyllus rauschi may berestricted to woodpeckers and flickers, but it is so poorlyknown that this is only conjecture. The remainingspecies are known from a broad assortment of hostsincluding galliform and passerine birds. Darskaya(1950), Peus (1967) and Main & Wallis (1972) haveassigned species belonging to this subgenus to a numberof species groups. However, based on host preferencealone, the North American taxa appear to segregate into4 categories. The idius-group includes C. affinis,arcuegens, calderwoodi, celsus, coahuilensis, idius,riparius, petrochelidoni, and scopulorum which are


parasites of hirundinid passerines. The vagabundus-group includes C. lari, pelecani and vagabundus thatare mainly associated with sea birds and a fewfalconiform hosts. The gallinae-group includes C.enefdeae, gallinae, niger and rauschi, all associatedwith a broad range of passeriform families, especiallyemberizids and fringillids. Ceratophyllus rossittensisstands alone as a parasite of corvids, although one mightexpect it on hosts that use abandoned crow nests, suchas owls.

Following is a key to the members of the subgenusCeratophyllus reported from North America. In usingit the reader should bear in mind that females of a numberof species cannot be identified with any degree ofcertainty in the absence of accompanying males. Also,some of the diagnostic characters of the males,specifically the chaetotaxy and configuration of theapical lobe of the distal arm of st IX and the membranousappendages associated with the apex of st VIII may bedifficult or impossible to discern in undissectedspecimens.

Key to the species of the subgenus Ceratophyllus(Ceratophyllus)

1 Metathorax lacking a pleural arch; abdominalsegments with differential pigmentation producingvertical bands in lateral view .................. arcuegens

Metathorax with well developed pleural arch;abdominal segments uniformly pigmented, no distinctvertical bands evident ............................................ 2

2 Males .............................................................. 3Females ........................................................ 18

3 Movable process widest in its proximal half; dorsallobe of fixed process produced apically (Figure 16);apical lobe of distal arm of st IX expanded apically(Figure 39); apex of st VIII with 3 or 4 long setaesubtended by a long, narrow vexillum which may beslightly spiculose basally; crochet triangular ..........

................................................. petrochelidoniMovable process widest in the middle or in its distal

half; apical lobe of distal arm of st IX with parallel sides,not greatly expanded apically; other characters variable

........................................................................ 4

4 Fixed process long and slender, nearly reachingapex of movable process (Figure 7); movable processwidest in the middle, cusp in cephalic margin near apex(Figure 7) ............................................ scopulorum

Fixed process short and/or rounded (Figures 8, 20),or if slender, not reaching beyond three fourths the length

of the movable process (Figures 10, 12) .............. 5

5 Fixed process slender from base to apex, notreaching beyond three fourths the length of movableprocess; apical fovea on caudal margin of fixed processengaging cephalic, marginal, incrassate cusp of movableprocess (Figures 10, 12) ........................................ 6

Fixed process triangular or broad and bluntlyrounded at apex (Figures 6, 14, 18) ...................... 7

6 Fixed process engaging cephalic margin of movableprocess near middle of the latter (Figure 10); apical lobeof distal arm of st IX with subparallel margins, itscephalic margin with a slight median bulge (Figure 31);membranous vexillum of st VIII with a trace of basalspicules on dorsal margin (Figure 54) ........... celsus

Fixed process engaging cephalic margin of movableprocess about one third down the latter (Figure 12);rounded apical lobe of distal arm of st IX with parallelmargins, its cephalic margin smoothly rounded (Figure30); membranous vexillum of st VIII lacking spiculeson dorsal margin .................................. calderwoodi

7 Movable process only ~2X as long as wide; fixedprocess approximately oval, broad and bluntly roundedat apex (Figure 14); upper and lower lobes of distal armof st IX separated by a conspicuous gap in caudal margin(Figure 32); rounded apex of st VIII with 3 short setae;vexillum widest in the middle; crochet only slightlylonger than width of st VIII, sharply pointed and lightlysclerotized on dorsal and ventral margins ...............

.................................................... coahuilensisMovable process >2X as long as wide, apex variable

(Figures 9, 15); no conspicuous gap in caudal margin ofdistal arm of st IX separating upper and lower lobes(Figures 34, 38); apex of st VIII with a variable numberof longer setae; crochet usually much longer than widthof st VIII, if sclerotized, only along dorsal margin(Figures 57, 62) ..................................................... 8

8 Movable process >4X as long as wide (Figure 6);apical lobe of distal arm of st IX curved caudad to arounded, acute tip (Figure 44); apex of st VIII with acone-shaped, spiculated projection, 3 subapical setae anda knife-shaped, membranous vexillum with spiculesbasally and along dorsal margin; crochet tapering to arounded tip apically ............................ vagabundus

Movable process shorter but >2X as long as wide(Figures 13, 17); apical lobe of distal arm of st IX broadlyrounded, without an acute tip (Figures 38, 42); apex ofst VIII without a cone-shaped projection (Figures 55,64), or if present, vexillum curved dorsally and crochetcurved ventrally and bluntly tipped (Figure 63) ... 9


9 Upper lobe of fixed process acuminate (Figures 11,19) ......................................................................10

Upper lobe of fixed process blunt and roundedapically (Figures 17, 19) ...................................... 11

10 Caudal margin of movable process slightly concavein middle (Figure 11); apical lobe of st IX strongly flexedcaudad, caudal margin strongly concave (Figure 33);apex of st VIII with 4 or 5 bristles, membranous vexillumbroad and knife-like; crochet broad and blunt (Figure60) ........................................................... enefdeae

Caudal margin of movable process with apronounced fovea on its caudal margin, subtended by aslight sinus (Figure 19); cephalic margin of st IX slightlyconcave (Figure 40); apex of st VIII with 3 long setaeand a long, thin, straight vexillum which lacks spicules;crochet slightly curved, tapering to a blunt tip (Figure52) ............................................................. rauschi

11 Apex of fixed process triangular, but blunt (Figures22, 23) .................................................................. 12

Apex of fixed process blunt and broadly rounded(Figures 17, 18) ................................................... 13

12 Less than half the length of movable processextending beyond apex of fixed process, lacking apronounced incrassation in cephalic margin (Figure 22);apical lobe of st IX relatively small, its margins parallel(Figure 42); apex of st VIII with 2 long bristles subtendedby a relatively short, curved, aspiculate vexillum; crochetblunt and slightly curved (Figure 53) .... rossittensis

More than half the length of movable processextending beyond apex of fixed process, with apronounced incrassation in cephalic margin (Figure 23);apical lobe of distal arm of st IX with subparallelmargins, widest apically, rounded (Figure 34); apex ofst VIII with long bristles and a membranous vexillumthat lacks spicules; crochet with a concave ventralmargin, slightly sclerotized dorsally (Figure 50) ....

.............................................................gallinae

13 Movable process 3X or more as long as broad(Figure 15) ........................................................... 14

Movable process <3X as long as broad (Figure 21)..................................................................... 16

14 Movable process extending no more than one fourthof its length beyond tip of fixed process (Figure 18);vexillum with a distinct basal spiculate lobe and spiculatealong entire dorsal margin; crochet wide and curvedventrally, with a blunt tip (Figure 63) ............ idius

Movable process with one third or more of its length

extending beyond tip of fixed process (Figure 15);vexillum at most with small cone-shaped process (Figure57), vexillum with spicules limited to basal portion ofdorsal margin; if curved ventrally, crochet narrow(Figures 55, 57) ................................................... 15

15 Apical lobe of distal arm of st IX tapering slightlyapically, broadest basally (Figure 37); apex of st VIIIwith about 3 long setae, membranous vexillum spiculatebasally on dorsal margin; crochet concave ventrally,slightly sclerotized dorsally (Figure 55) ......... niger

Apical lobe of distal arm of st IX slightly wider, itsmargins parallel, its apex bent caudad (Figure 28); apexof st VIII bearing a cone-shaped process subtended by2 setae and a curved vexillum bearing a few spicules onits dorsal margin; crochet triangular, flexed ventrally

............................................................... affinis

16 Movable process with a distinct angle andincrassation at the point where it engages the fovea onthe fixed process, the latter of which is flexed cephaladapically (Figure 21); apex of st VIII with 3 or 4 longsetae, vexillum much longer than width of st VIII (Figure49) ............................................................ pelecani

Movable process lacking a distinct angle in itscephalic margin though an incrassation may be evident,its overall shape clavate (Figure 13); vexillum at mostonly slightly longer than width of st VIII (Figures 58,64) ...................................................................... 17

17 Movable process clavate, widest in its apical third(Figure 13); apex of st VIII usually with 2 long setae,vexillum very short, usually less than width of st VIII,without spicules (Figure 64) ............................. lari

Movable process clavate, widest in the middle(Figure 17); apex of st VIII with a spiculate terminus,subtended by a cluster of long setae, length of vexillumabout the width of st VIII (Figure 58) ........ riparius

18 Sternite VII with a conspicuous lobe subtended bya sinus .................................................................. 19

Sternite VII undulate or lacking a lobe subtendedby a sinus, though a broad, shallow depression may bepresent .................................................................. 26

19 Marginal lobe of st VII squared caudally, subtendedby a shallow emargination, not a distinct sinus (Figure86) ............................................................... affinis

Marginal lobe of st VII pointed or broadly rounded,with a distinct sinus below it (Figures 78, 83) .... 20

20 Upper and lower lobes of st VII approximatelyequal in size (Figures 79, 80) .............................. 21


Upper and lower lobes of st VII subequal (Figures81, 83) .................................................................. 22

21 Upper and lower lobes separated by a broad,shallow sinus (Figure 79), on Petrochelidon fulva

................................................... .coahuilensis Upper and lower lobes separated by a narrow, shallowsinus (Figure 80) ....................................... pelecani

22 Marginal lobe of st VII pointed, its margin belowthe sinus not projecting as far as the apex of the lobe(Figure 83) ....................................................... idius

Marginal lobe of st VII blunt or rounded above thesinus (Figure 82) .................................................. 23

23 Marginal lobe of st VII blunt, subtended by a deepsinus, the margin of which is incrassate (Figure 81)

...................................................... scopulorumMarginal lobe of st VII blunt, margin of sinus not

incrassate (Figure 82) .......................................... 24

24 Abdominal tergites II and III usually with 4marginal spinelets; marginal lobe of st VII blunt, thelower lobe as long as or longer than the upper lobe(Figure 78) ................................................. riparius

Abdominal tergites II and III usually with only 2marginal spinelets; marginal lobe of st VII blunt, thelower lobe shorter than the upper (Figure 82) ...25

25 First and third antepygidial setae very short, each<1/4 as long as second; ~8 short setae above the spiracleon t VIII; on Petrochelidon spp; emarginate forms of .

........ celsus, calderwoodi and petrochelidoni*First and third antepygidial setae at least 1/4 as long

as the second; ~11 short setae above the spiracle on tVIII; not usually associated with Petrochelidon spp.;emarginate form of ......................................... niger

26 Caudal margin of st VII undulate or with a broad,shallow sinus (Figures 84, 89) . ........................... 27

Caudal margin of st VII smoothly rounded, entire,without a shallow sinus (Figures 90, 92) . .......... 28

27 Caudal margin of st VII irregularly rounded withslight undulations, sometimes with a broad, shallowconcavity, particularly caudoventrally; bursa copulatrixpartially sclerotized (Figure 84) ................... celsus

Caudal margin of st VII with a slightly undulatelower lobe, above which is a broad, shallow sinus; bursacopulatrix without sclerotization (Figure 89) .. rauschi

28 Only a small portion of the ductus obturatussclerotized (Figure 95) ......................... ..rossittensis

Proximal 1/2 of ductus obturatus and usually atleast a small part of the bursa copulatrix sclerotized29

29 Sclerotized portion of bursa copulatrix about 1/2as long as sclerotized portion of ductus obturatus(Figures 90, 92) ................................................... 30

Sclerotized portion of bursa copulatrix, if present,much shorter than sclerotized portion of ductus obturatus(Figures 87, 93) ................................................... 31

30 Spiracle on t VI circular; darkly pigmented fleas,mainly on passerine birds; non-emarginate form of .

................................................................. nigerSpiracle on t VI large and appearing doubled, dark

fleas mainly on gulls and raptors ........ vagabundus

31 Mesepisternum with 12 or more thin setae; darklypigmented fleas mainly on gulls and cormorants ....

.................................................................... lariMesepisternum with 10 or fewer thin setae . 32

32 Frontal tubercle minute ...................... gallinaeFrontal tubercle prominent ........................... 33

33 Hilla of spermatheca half as long as bulga (Figure94) ........................................................... enefdeae

Hilla of spermatheca less than half as long as bulga;non-emarginate form of ................ calderwoodi andpetrochelidoni**

*Identity of the emarginate form of females of these 3species can only be inferred from the presence of males.**Identity of non-emarginate form of females of these2 species can only be inferred by the presence of males.

Ceratophyllus (Ceratophyllus) affinis Nordberg, 1935(Figures 20, 28, 57, 86)

Ceratophyllus affinis Nordberg, 1935. Memor. Soc.Fauna Flora Fenn. 10: 354-369, Figures 1-15.Finland, Turku Japori District, Nystad [=Uusikau-punki light house], 60.48N 21.30E, host and datenot stated but probably Hirundo rustica, Södermanleg. One male Syntype in Museum ZoologicumUniversitatis, Helsinki [MZHF].

Ceratophyllus affinis neglectus. Smit, 1958, Proc. U.S. Nat. Mus. 108 (3394): 51, Figures 1-2, 5-10.USA, Utah, Cache County, Smithfield, 41.50N111.50W, from Hirundo rustica nest, 6.VI.1951,T. Tibbetts leg. Holotype male, Allotype female,United States National Museum [USNM] No.63891.


Diagnostic CharactersMale Penis rods coiled through two 360°

convolutions. With a single antepygidial seta per sidebordered dorsally and ventrally by a tiny vestigial bristle.The 2 acetabular setae arise on a slight caudal projectionof the fixed process. Spiculose area much reduced onthe dorsocephalic third of t VIII which also bears ~10marginal and submarginal bristles. Movable processclavate, 2.5X as long as wide at its widest point, its apical1/3 extending beyond the dorsal apex of the fixedprocess, its cephalic margin with a distinct denticalmidway between the base and the apex. Margins of stVIII subparallel, with 2 subapical long setae, its apexbearing a fimbriated, cone-shaped lobe subtended by along, pointed vexillum that is basally spiculose on itsdorsal margin. Margins of st IX parallel apically, itsapex smoothly rounded. Crochet blade-like, relativelylightly sclerotized and tapering to a blunt tip.

Female Spermathecal bulga set off from hilla by aslight constriction Three antepygidial setae per side,the medial 4X as long as the upper, 6X as long as thelower. Bursa copulatrix only slightly sclerotizeddorsally. Ductus obturatus well sclerotized for most ofits length. Caudal margin of st VII with a distinct lobe.Anal stylet short and conical, 2.75X as long as wide atits base with 1 long apical seta and 2 short subapicalsetae arising on the dorsal and ventral margins. Analsternite quadrate with a number of marginal setae noneof which are particularly spiniform.

RemarksThis is a poorly known species, the validity of which

as a separate species is debatable. Its describer relatedit to C. borealis, but Smit (1956), in his redescription ofNordberg’s taxa, placed it in the C. gallinae group ofspecies. He later (1958) associated it with C. idius andC. niger. In any case, based on the shape of the malegenitalia, it belongs to the group of species in which themovable process is approximately elongate-oval orclavate and the apex of st VIII bears a fimbriated lobesubtended by a finger-like projection on its cephalicmargin as shown in Figure 57.

DistributionThere are few published records of this species,

aside from the type series from Finland, also cited bySmit (1969). It has been reported from Germany bySmit (1958), Norway by Mehl (1967), Sweden byBrinck-Lindroth & Smit, (1971) and various localitiesin the former USSR. Prokop’yev (1969) cited thisspecies as common on an island in the delta of theSelenga River (~52.15N 106.40E) in Lake Baikal,

Kabansky Rayon, Buryatia. Darskaya (1950) reported11 males and 20 females from the Vologda District,about 450 km north of Moscow. Darskaya’s illustrationof the male movable process, while wider than in theholotype male of C. affinis neglectus, does approximatethat of the paratype male of this taxon as illustrated bySmit (1958). Although her drawings are not as detailedas Smit’s, she did have a good series of males on whichto base them. The slight differences between C. affinisneglectus and the nominate subspecies do not appear tous to be sufficient to support separation into subspecies,in spite of the highly disjunct distribution of the fewcollection records. In erecting the subspecies C. a.neglectus, Smit (1958) suggested that the materialreported by Cross & Knowlton (1953) from Smithfield,Utah, probably belonged to this taxon as well.

Host AssociationsFrom the few records it seems that this is a flea of

the barn swallow, Hirundo rustica. Various authors haveargued that this host has no fleas of its own since thenests are too dry to allow the larvae to develop. Thismay be the case for nests in buildings and on other man-made structures, but where nesting conditionsapproximate the natural state, such as cave entrancesand other protected sites on cliffs, etc. it seems the barnswallow is the primary and preferred host.

Ceratophyllus (Ceratophyllus) arcuegens Holland,1952

(Figures 5, 29, 59, 88)

Ceratophyllus arcuegens. Holland, 1952, Can. Entomol.84: 300, Figures 8-15. Canada, Yukon Territory,Rampart House, 67.25N 140.59W, fromPetrochelidon pyrrhonota nest, 10.VIII.1951, J. E.H. Martin leg. Holotype male, Allotype female,Canadian National Insect Collection [CNIC] No.6000.

Diagnostic CharactersMale The penis rods are coiled about twice 360°.

A single long antepygidial seta is present with a tiny,vestigial bristle at its base above and below. A prominentdenticle is situated on the cephalic margin of the movableprocess about 2/3 down from the apex The shape ofboth the fixed and movable processes are unique. Themovable process is widest subapically and is slightlymore than 3X as long as wide. The dorsal projection ofthe fixed process is short and bears a fovea that engageswith the denticle in the cephalic margin of the movableprocess. The 2 acetabular bristles arise low on thecaudally projecting margin of the fixed process. Tergite


VIII bears an extensive area spiculosa but the spiculesare not particularly prominent and there are ~14 marginaland submarginal bristles along the dorsal margin. Theapex of st VIII in the holotype specimen is not quite asillustrated in Holland (1952). As so often happens withvery delicate structures, the fimbriated projection beyondthe base of the vexillum and the apex of st VIII is subjectto folding over so that it projects ventrad rather thandorsad. This seems to be the case in the holotype andthe other 3 males. As a result, the fimbriated apex,though visible, is presented in a different aspect and thevexillum looks crescentric with fimbriations projectingbelow it. With this exception the illustrations in Holland(1952) are essentially accurate except for slightvariations in the distribution of a few setae which are oflittle or no taxonomic importance. Crochet broad basallybut sharply tapering to a blunt, peg-like apex.

Female The spermathecal bulga is enlarged at theduct end and is set off from the papillate hilla by ashallow constriction. The bursa copulatrix isunsclerotized but the ductus obturatus is sclerotizedbasally. Sternite VII has an angulate caudal margin. Ofthe three antepygidial setae the medial is 2.7X longerthan the dorsal bristle, 6.3X longer than the ventralbristle. The spiracular fossa of t VIII is 3.3X as tall aswide and surrounded by three to five fine setae per side.The anal stylet is 2X as long as wide at the base.

RemarksThis “pot bellied” species is distinctive among the

North American fauna in both sexes by the ventralexpansion of the abdominal segments, the alternatinglight and dark bands on these segments and the absenceof the pleural arch. The reduction of the latter isaccompanied by the loss of the ability to jump. Holland(1952) observed that the species is a crawler, rather thana jumper, as are most other members of the genus. Hopla(1965b) observed that there were hundreds of fleas thatcrawled into cracks and crevices in the woodwork duringhis collecting activities in Livengood, Alaska (65.31N148.40W) and he suspected that these were C. arcugens.As a result, he only collected a single male belonging tothis species, while C. scopulorum abounded in the samenests. Holland (1985) expanded on this further, notingthat this species was similar to the Old World species,C. rusticus, C. caliotes and C. orites, which have alsolost the ability to jump. He noted the reduction of thethorax and legs and the difference in pigmentationbetween the anterior and posterior parts of the abdominaltergites and sternites resulting in the alternation of darkand light vertical bands. This character is even visiblein unmounted material.

While many members of this genus are difficult to

distinguish, even in the males, C. arcugens stands outin both sexes. The male clasper is unlike that in anyother species, as is the apex of the male st VIII, and theabdominal banding easily separates the females fromthose of other North American species.

DistributionThis is a species occurring in Traub’s (1983)

Forested and North Canadian Sector, including Alaska,Yukon and the Northwest Territories and northernBritish Columbia in the extreme northwestern part ofthe range of its preferred host. The British Columbiarecord was given in Pilgrim and Galloway (2000). Aswith other bird fleas whose hosts abandon their nestsduring the colder months, these fleas evidently spendthe winter as adults, enclosed in the pupal cocoon,emerging in the spring upon the return of their hosts.

Host AssociationsAs Hopla (1965b) suggested, this appears to be a

specific parasite of Petrochelidon pyrrhonota, in Alaska,Yukon and Northwest Territories, and it often coexistswith C. scopulorum in many of the same nests. It wasnot reported from Alaska by Haas et al. (1985).

Ceratophyllus (Ceratophyllus) calderwoodi Holland,1979

(Figures 12, 30, 51, 82)

Ceratophyllus (Ceratophyllus) calderwoodi. Holland,1979, Can. Entomol. 111:717, Figures 9-12.Canada, New Brunswick, Kouchibouguac NationalPark, ~46.55N 64.58W, from Petrochelidonpyrrhonota, 7.IX.1977, G. Calderwood leg.Holotype male, Allotype female, CNIC No. 15896.

Diagnostic CharactersMale Antepygidial setae one per side bordered

dorsally and laterally by a tiny setula. Spiculose areaof t VIII well developed, extending about 3/4 the lengthof the anterodorsal margin. Apex of st VIII with fourlong apical setae and a long, blade-like, basallyfimbriated vexillum which is spiculate along the dorsalmargin of the basal half. The penis rods are coiledthrough two 360° convolutions and the two acetabularsetae arise on a slightly developed acetabular projectionon the ventrocaudal angle of the fixed process. Themovable process of the clasper is clavate-fusiform,widest near its apex and ~3.5X as long as wide, itscephalic and caudal margins are almost parallel exceptat the proximal and distal ends. With a well developeddenticle on the cephalic margin slightly less than 1/3of the way down from the apex. Fixed process


cylindrical, its dorsal projection bearing a welldeveloped fovea. Crochet moderately sclerotized,tapering evenly to a rounded point.

Female Spermathecal bulga sausage-shaped, setoff from the papillate hilla by a shallow constriction.The spermathecal duct inserts apically into the bulga.Bursa copulatrix partly sclerotized, ductus obturatussclerotized basally. Caudal margin of st VII is shownby Holland (1985) to be dimorphic, entire or bearing alobe subtended by a sinus. Of the three antepygidialsetae the medial is 6X as long as the upper, 7X as longas the lower. There are a few short setae per side aroundthe spiracular fossa on t VIII which is 3X as high aswide. Anal stylet 3X as long as wide at its base.

RemarksLiterature concerning this species is limited. Based

on the configuration of the male genitalia it is related toC. celsus and C. scopulorum. In these taxa the movableprocess is about 3X as long as wide at its widest pointand the acetabular setae arise low on the caudoventralangle of the fixed process.

Benton (1998) discussed the occurrence of both C.celsus and C. scopulorum in the nests of Petrochelidonpyrrhonota in New Brunswick, Canada, and presenteda case for treating C. calderwoodi as possibly the productof interbreeding between these 2 taxa but requiringfurther study. He noted that this could explain thedimorphism in the st VII of the females of both C.calderwoodi and C. celsus where the range of the latteroverlaps that of C. petrochelidoni in British Columbiaand parts of western United States. For the present weare treating this as a separate species since no molecularstudies or breeding data are available. Perhaps moreintensive collecting in the area where these species aresympatric will help clarify the relationship of C.calderwoodi to both C. celsus and C. scopulorum.

DistributionAlthough Holland (1985) indicated that this species

was known only from the type locality, his map 61 alsoshows a collection from the Brokenhead River where itcrosses highway #59, ~50.24N 96.36W, near thesouthern tip of Lake Winnipeg, Manitoba.

Host AssociationsTo date this species is known mainly from the type

series of slightly over 50 pairs. It is thought to be specificto Petrochelidon pyrrhonota which it shares with at least4 other species of Ceratophyllus. Probably the only otherswallow with more species of fleas is the house martin,Delichon urbica, native to the Palaearctic Region.

Ceratophyllus (Ceratophyllus) celsus Jordan, 1926(Figures 10, 31, 54, 84)

Ceratophyllus celsus. Jordan, 1926, Novit. Zool. 33:387, Figure 4. Canada, British Columbia, OkanaganFalls, 49.21N 119.33W, from Riparia riparia,26.IV.1913, C. Garrett leg. Holotype male, bymonotypy, BMNH.

Ceratophyllus celsus apricus. Jordan, 1929, Novit.Zool. 35: 37, pl. II, Figure 24. Cuba, Soledad, fromnest of Petrochelidon fulva, 4.IV.1925, J. Bequaertleg. [Holo]Type male, USNM No. 41439.

Diagnostic CharactersMale Antepygidial setae 1 per side bordered

dorsally and laterally by single tiny setulae. Spiculosearea on dorsal margin of t VIII well developed, confinedto the dorsocephalic half. Apex of st VIII graduallytapering toward the apex which bears 3 long bristlesand a subapical, blade-like vexillum which is notfimbriated but does bear a few basal spicules. Penisrods coiled through slightly more than 360°. Acetabularsetae arising low on the ventrocaudal angle of the fixedprocess of clasper. Movable process of clasper clavate,2.75X as long as wide, with a rounded apex and aprominent denticle on its cephalic margin. Fixed processcolumnar, 5.8X as high as wide, its apex blunt. Crochetsclerotized, broad at base but tapering to a rounded pointapically.

Female Spermatheca slightly expanded at ductend, set off from the papillate hilla by a shallowconstriction. Bursa copulatrix and ductus obturatusboth sclerotized, the former only slightly. Caudalmargin of st. VII variable, from smoothly rounded todeeply emarginate. Three antepygidial setae, the median6.5X as long as the upper, 6X as long as the lowerbristle. Spiracular fossa 2.8X as high as wide,surrounded by ~10 fine setae. Anal stylet conical, 2.5Xas long as wide at base, with 1 long apical seta and 2well developed subapical setae on outer surface.

RemarksCeratophyllus celsus belongs to a species group

which includes C. calderwoodi and C. scopulorum inwhich the acetabular setae arise low on the caudal marginof the fixed process of the clasper. In the males of all 3species, the movable process of the clasper is about 3Xas long as wide at its widest point. Although it couldnever be confused with C. scopulorum, the modifiedsegments of the males are sufficiently similar to thoseof C. calderwoodi to suggest that the 2 may beconspecific. Benton (1998) indicated that there issufficient evidence available to consider C. calderwoodi


a synonym on the basis of intermediate forms of femalesthat he felt were possibly hybrids. See additionalcomments on this subject under the C. calderwoodiaccount. A detailed description of the larva of thisspecies is given in Pilgrim and Galloway (2000).

DistributionCeratophyllus celsus is widely distributed in North

America. It occurs from Alaska (Hopla, 1965b, Holland,1985, Haas & Wilson, 1980, Haas et al., 1980, 1981),south into British Columbia (Holland, 1985),Washington, Montana and Nevada (Lewis et al., 1988).Holland (1985) cited C. celsus from Manitoba, eastthrough Ontario (see also Galloway, 1987), Québec andNew Brunswick. Galloway (1987) reported 116specimens collected at 11 locations in Ontario alone. Itis known through the midwestern United States to SouthDakota (Easton, 1982), Nebraska and Arkansas (Lewis& Galloway unpublished), and Oklahoma (Hopla &Loye, 1983, Loye & Hopla, 1983). It should be notedthat the record for C. petrochelidoni for South Dakota(Easton, 1982) is, in fact in error, and should correctlybe for C. celsus. The original report was based on asingle female, one with a sinus on st VII, and whichkeyed out to C. petrochelidoni. Easton subsequentlycollected additional specimens from the same cliffswallow colony in Jackson County, including manymales as well as emarginate and non-emarginatefemales, and these were all C. celsus. The mostsoutherly report of this species is from Texas by Eads(1956). In the east, C. celsus has been recorded byBenton (1980) from Illinois, Michigan, New York andVermont.

Host AssociationsThroughout most of North America, C. celsus is a

parasite of the cliff swallow, Petrochelidon pyrrhonota.However, in Alaska and in British Columbia, it has moreoften been collected in the nests of the bank swallow,Riparia riparia (Hopla, 1965b, Haas et al., 1980,Holland, 1985), and in Alaska, it appears to be quiterare in cliff swallow nests (Haas & Wilson, 1980). Haaset al., (1981) recorded C. celsus from nests of the violet-green swallow (Tachycineta thalassina) in Alaska.

Ceratophyllus (Ceratophyllus) coahuilensis Eads,1956

(Figures 14, 32, 65, 79)

Ceratophyllus coahuilensis. Eads, 1956, J. Parasitol.42: 73, pl. I, Figures A-F. USA, Texas, Uvalde

County, sine loco, from Petrochelidon fulva nests,30.III.1955, R. B. Eads leg. Holotype male,Allotype female, USNM No. 62924.

Diagnostic CharactersMale One long antepygidial seta per side bordered

dorsally and laterally by 1 tiny setula. Spiculose area oft VIII forming a distinct, narrow band along thecephalodorsal 3/4 of the dorsal margin. About 8marginal and submarginal setae along the dorsal marginof this tergite. Apex of st VIII somewhat clavate, bearing3 apical setae per side and an apical, expanded vexillumthat is fimbriated along its dorsal margin. Penis rodscoiled through two 360° rotations. Two acetabular setaearising on the caudal margin of the oval fixed processslightly above the middle on a slight convexity. Movableprocess of the clasper clavate, almost exactly 2X as longas wide at its widest point, its cephalic margin almoststraight above and below the denticle, its apex rounded.Fixed process of clasper oval, rounded dorsally, ~1.25as high as wide. Crochet strongly sclerotized andtapering to a sharp point.

Female Spermatheca sausage-shaped, the bulgawidest near the apical insertion of the spermathecal duct.Hilla slightly narrower than bulga, conspicuously setoff from it and bearing an apical papilla. Bursacopulatrix slightly sclerotized distally, ductus obturatussclerotized over most of its length. Caudal margin of stVII with a shallow sinus below a rounded dorsal lobe.Three antepygidial setae, the medial 3.6X longer thanupper, 4.4X longer than lower bristle. Spiracular fossa3X as high as wide, surrounded by ~10 to 12 fine setae.Anal stylet conical, 2.7X as long as wide at base, with 1long apical seta and 2 shorter subapical bristles.

RemarksBeyond the type series, the only published record

available for this species is that of Reddell (1966) fromEdwards County, Texas. However, there is a long seriesof both sexes in the Lewis Collection taken fromPetrochelidon fulva at Dunbar Cave, also in EdwardsCounty, Texas.

DistributionThis is a distinctive species that seems to have a

range restricted to northeastern Mexico, southern Texasand southeastern New Mexico. There are literaturerecords from Kerr and Uvalde counties in Texas, andHaddow et al. (1983) provided records from southeasternNew Mexico and the Mexican state of Nuevo Leon.


Host AssociationsAll known specimens have been taken from the

nests of Petrochelidon fulva, although the record fromNew Mexico may be close to extralimital to the rangeof this host species.Ceratophyllus (Ceratophyllus) enefdeae Ioff, 1950

(Figures 11, 33, 60, 94)

Ceratophyllus enefdeae. Ioff, 1950, In: Ioff et al., Med.Parazitol., Moskva 19: 269. USSR, northernCaucasus, Teberda, in garret of house, from nest ofunknown bird, no additional data. No typedesignated.

Ceratophyllus balati. Rosicky, 1955, Zool. Entomol.Listy 4: 371, 382-383, Figures 6, 7.Czechoslovakia, High Tatra Mts, 2000 m, Skalnatépleso, from Anthus spinoleta, 17.VI.1955, F. Balatleg. Holotype male, Biological Institute,Czechoslovak Academy of Sciences, Praha, CzechRepublic [BICAS] No. 1358/1/1. Synonymized bySmit, 1966: 72. Included here since Hopla (1965b)reported this species from Alaska.

Diagnostic CharactersMale One antepygidial seta per side bordered

dorsally and laterally by single tiny setulae. In thisspecies the setulae arise somewhat farther from themedial than in most other species in the subgenus.Spiculose area of t VIII well developed on the cephalic2/3 of the dorsal margin which also bears 5 to 7 marginaland submarginal setae. Sternum VIII with a dorsalmargin that is almost straight, its ventral margin curved,causing the structure to be wider in the middle thanproximally or distally. Its apex bears 3 to 5 long bristlesand a long, blade-like, unfimbriated vexillum withoutspicules. Although Holland (1985) stated that the penisrods in this species do not complete a 360° convolution,all but 1 specimen in our sample have penis rods thatdo. Acetabular setae arising on a slight hump about 2/3down the caudal margin of the fixed process. Movableprocess 4X as long as wide, widest apically, its cephalicmargin almost straight above the prominent denticle.Crochet well sclerotized, its margins almost parallel, itsapex blunt.

Female Spermatheca sausage-shaped, its hillaslightly papillate and set off from the bulga by a distinctconstriction. The hilla is slightly narrower than thebulga. Holland (1985) showed an apical papilla on thehilla, Hopla (1965b) did not. Of the 10 females in theLewis Collection, all show a slight papilla, but in 2 it isbarely perceptible. Bursa copulatrix is unsclerotizedexcept near its junction with the well sclerotized ductus

obturatus. Caudal margin of st VII entire and smoothlyrounded, without a sinus. Three antepygidial setae, themedial 7.5X as long as the upper, 9.0X as the lower.Anal stylet 2.4X as long as wide at base, with 1 longapical seta and 2 shorter subapical bristles.

RemarksWhile taxonomic revision is not the particular goal

of this study, there are similarities between C. gallinaeand C. enefdeae that are difficult to ignore. Perhaps themost telling similarities involve the modified abdominalsegments of the male. In both species the movableprocess is slightly more than 3X as long as wide and inboth it extends slightly less than 1/2 of its length beyondthe apex of the fixed process. The 2 acetabular setaearise about 1/3 up the caudal margin of the fixed processand the membranous vexillum on the apex of st VIII islong, broad and lacking fimbriation. However, theaedeagal crochets are somewhat different in profile.Females of both species have similar seventh sternites,a well-developed ductus obturatus and the sausage-shaped bulga of the spermatheca is well set off from thehilla which bears a well developed apical papilla.

DistributionThis is another poorly known but widely distributed

species that is clearly related to C. gallinae, if notconspecific with it. Actually, in view of the widespreaddistribution of C. gallinae, a case could be made that C.enefdeae is but a geographical variant of that speciesand that the subspecies C. e. tjanshani is a part of a clineextending from the Swiss Alps to the mountains ofAlaska. That is certainly consistent with the type ofdistribution illustrated in map 18 in Traub et al. (1983).Specimens from Poland, Nepal and Xinjiang Province,China, in the Lewis Collection show the movableprocess of the male clasper growing increasinglynarrower from western to eastern collection localities.Comparative material from Alaska was not available tous but the illustrations of C. balati in Hopla (1965b)suggest that this trend continues into western NorthAmerica.

Host AssociationsHost preferences in this species are less well known

than for many members of the genus but from what isknown it prefers passerine birds which nest on or nearthe ground, a trait shared with C. (C.) gallinae. Thereare 20 specimens belonging to this species in the LewisCollection. The 2 pairs from Poland and 2 males and 1female from Nepal are from birds. However, of theremaining specimens, 2 males were taken on a wooleyhare (Lepus oiostolus) in northern Nepal at ±12,500 ft,


another 2 females were taken from the floor of a hut at11,000 ft, and another female was collected whileflagging for ticks in the Langtang Valley. All 8 of thespecimens from Manas (44.16N 86.02E), Xinjiang,China, were collected in a pasture, not from a host. Likeso many other fleas that parasitize ground-nesting birds,adults evidently disperse from the site where theydevelop when actively seeking new hosts.

Ceratophyllus (Ceratophyllus) gallinae (Schrank,1803)

(Figures 23, 34, 50, 93)

Pulex Gallinae. Schrank, 1803, Fauna Boica 3(1): 195.Germany, Bavaria, Gross Kotzenreuth, nearEschenbach, Oberfalz, 49.46N 11.48E, from nestsof Gallus gallus, 25.III.1966, F. Peus leg. Neotypemale, BMNH. Designated by Peus, 1967: 88.

Ceratophyllus gibsoni. C. Fox, 1914, Bull. Hyg. Lab.Wash. 97: 15, pl. V, Figures 4-5. Canada, PointLepreux, Ottawa, 45.25N 75.43W, from hen house,13.VII.1909, A. Gibson leg. Lectotype male,USNM No. 18457. Synonymized by Ewing & Fox,1943: 69.

Diagnostic CharactersMale One antepygidial seta per side bordered

dorsally and laterally by single small setulae that arelonger than seems typical for most other members ofthis subgenus. Spiculose area of t VIII well developedand conspicuous, covering the anterior 1/2 of thesegment and extending well down the sides to form anarea larger than the sensilium. Dorsal margin of t VIIIwith 6 or 7 marginal and submarginal setae. SterniteVIII tapering gradually to apex which bears 2 apicalbristles per side and a long, slender vexillum that isunfimbriated but bears a few spicules on its dorsalmargin at the base. Penis rods completing not quite two360° convolutions. Acetabular setae arising slightlybelow the middle of the caudal margin of the fixedprocess. Movable process about 3.25X as long as wideat its widest point, its apex acute, the denticle in itscephalic margin well developed, arising approximately3/5 down from the apex. Fixed process oval, round-pointed dorsally, about 2X as high as wide. Crochetsclerotized, its rounded apex deflected slightlydownward.

Female Bulga of spermatheca wider than thepapillate hilla and set off from it by a conspicuousconstriction. Spermathecal duct entering the bulgaterminally. Bursa copulatrix only weakly and partiallysclerotized. Ductus obturatus strongly sclerotized formost of its length. Caudal margin of st VII with a

smoothly rounded lobe and lacking a sinus. Threeantepygidial setae, the median 4X the length of the upper,3.3X the length of the lower. Spiracular fossa 5X ashigh as wide, surrounded by 6 to 8 fine setae. Analstylet conical, 2.6X as long as wide at base, with onelong apical seta and dorsal and ventral subapical setaethat are 3/4 the length of the apical bristle.

RemarksThis is one of the two members of the genus to

warrant a standardized common name from theEntomological Society of America, the other being C.niger. It is called the European chicken flea and reflectsits frequent association with domestic chickens.However, the name is somewhat misleading since thespecies is most commonly encountered in the nests ofpasserine birds. Although the nomenclatural history ofthe species has been confused by the assignment of somany different names, it seems to be the consensus ofmost modern specialists that C. gallinae is a singlespecies that lacks subspecies. This is especially so sinceC. tribulus Jordan, 1926, is now considered a separatespecies rather than a subspecies of C. gallinae.Therefore, Smit and Wright (1965) in their study of theWagner Collection in Hamburg, Germany, synonymizedC. g. kievensis Wagner, 1930a, with C. tribulus Jordan,1926. Smit and Wright (1978a) also synonymized C. g.dilatus Dudolkina, 1946 (in Ioff et al. 1946), with C.tribulus. Neither Traub et al. (1983) nor Lewis (1990)mentioned these subspecies, although Holland (1985)still treated them as valid. Although Holland was aspecialist in bird fleas, we view his inclusion of thesesubspecies as an oversight on his part and not an effortto resurrect the names. The larva has been described bymany authors, most recently by Pilgrim and Galloway(2000).

DistributionThis is truly a cosmopolitan Holarctic species.

Haddow et al. (1983) showed numerous records fromthe temperate portion of the Northern Hemisphere. It isa common flea on a broad range of avian hosts in Eurasiaand was introduced into eastern United States andCanada, probably on domestic poultry. It has a distinctpreference for birds with dry nests. The species is notmentioned by Segerman (1995) in her treatment of theSouth African fauna. However, it was reported fromAustralia by Roberts (1952), Dunnet and Mardon (1974)and Calder (1996). It has also been reported from NewZealand by Thompson (1922), Marples (1942), Smit(1965), King (1976) and Pilgrim (1980). It is found ina variety of habitats from sea level to 3,000 m, but ismost common at lower elevations.


Host AssociationsThe species is known from a broad range of hosts

including a few mammals. Except for records onClethrionomys gapperi, Rattus norvegicus and Mustelanivalis, all mammal records have come from variousspecies of squirrels, hosts most likely to come intocontact with arborial bird nests. The species shows littlespecific preference among avian hosts and there are afew reports from humans, but usually when associatedwith domestic poultry, specifically chickens.

Ceratophyllus (Ceratophyllus) idius Jordan &Rothschild, 1920

(Figures 18, 35, 63, 83)

Ceratophyllus idius. Jordan & Rothschild, 1920,Ectoparasites I: 73, Figures 70-72. Canada, BritishColumbia, Okanagan Landing, 50.12N 119.22W,from Iridoprocne [=Tachycineta] bicolor, VII.1913,J. A. Monro leg. Lectotype male, BMNH,designated by Smit & Wright, 1978b: 24.

Ceratophyllus sternacuminatus. Brown, 1968, Can.Entomol.100: 492, Figures 2-3. Canada, NewBrunswick, Fredericton, 45.57N 66.40W, fromIridoprocne [=Tachycineta] bicolor nest, 7.I.1959,N. R. Brown leg. Holotype male, in collection ofauthor. Synonymized by Smit & Wright, 1978a:42.

Diagnostic CharactersMale One antepygidial seta per side, arising

on a slight projection of the margin of t VII and bordereddorsally and laterally by small setulae. Spiculose areanarrow but well developed on anterior half of thedorsocephalic margin of t VIII which also bears ~6marginal and submarginal setae. Apex of st VIII with 2or 3 long bristles and a strongly fimbriated, U-shapedvexillum. Penis rods usually completing two 360°convolutions. Acetabular setae arising about the middleof the caudal margin of the columnar fixed process.Fixed process elongate-oval, its dorsal apex rounded.Movable process clavate, 2.5X as long as wide at itswidest point, which is only slightly below the apex.Prominent denticle on cephalic margin of the movableprocess very near the dorsal apex. Crochet wellsclerotized, its apex slightly deflected ventrad.

Female Bulga of spermatheca sausage-shaped, its duct end slightly wider than the remainderof the bulga and the papillate hilla is set off by a distinctconstriction. At least the basal half of the ductusobturatus is sclerotized, but the bursa copulatrix is not.Caudal margin of st VII bearing a sharply pointed lobe

subtended by a deep sinus. Three antepygidial setae,the medial 4.3X longer than the upper, 4.6X longer thanthe lower. Spiracular fossa 3.8X as high as wide andsurrounded by ~10 fine setae. Anal stylet narrower andmore sharply pointed than in most species, 3.6X as longas wide at base and bearing one long apical seta and 2short subapical bristles.

RemarksThis parasite is usually associated with tree hole-

nesting swallows and their relatives, such as purplemartins and tree swallows, that nest in bird houses,abandoned woodpecker nests, etc. In Alaska it is aparasite of the violet-green swallow at 40° north (Hopla,1965b). Benton (1998) made a case for interbreedingbetween this species and C. niger based on an analysisof variation in the shape of st VII in females fromWashington and Oregon. The caudal margin of st VIIis deeply incised in this species but smoothly roundedand entire in C. niger. Females in Benton’s sample wereintermediate between these two species, and althoughhe noted considerable variation in his sample hesuggested that this condition could be explained byhybridization. Unfortunately, the males in his samplewere not similarly examined to ascertain if they too wereintermediate in the configuration of their genitalia. Thelarvae of this species are described by Pilgrim andGalloway (2000).

DistributionHolland (1985) showed the northern distribution

for this species as including a number of records fromAlaska, British Columbia, Alberta and Manitoba.Eastern records are from Newfoundland, NewBrunswick, Quebec and Ontario. Records from theeastern United States extend as far south as Virginia,West Virginia and Tennessee (Benton, 1980). Publishedwestern records are confined to Minnesota, NorthDakota, Oregon and Washington and there is a longseries in the Lewis Collection from central Iowa.

Host AssociationsThis is mainly a parasite of hole-nesting birds,

especially certain swallows and martins. Records frombluebirds may be the result of previous occupancy bytree swallows. There are also records from other avianand small mammal hosts.

Ceratophyllus (Ceratophyllus) lari Holland, 1951(Figures 13, 36, 64, 87)

Ceratophyllus lari. Holland, 1951, Can. Ent. 83: 282,Figures 5, 7, 9. Canada, Northwest Territories,


Whaleback Island, Great Slave Lake,~62.00N112.00W, from nests of either Laruscalifornicus or L. argentatus, 23.VI.1947, J. R.Vockeroth leg. Holotype male, Allotype female,CNIC No. 5952.

Diagnostic CharactersMale With 1 long antepygidial bristle on a slight

tergal projection, bordered by 1 tiny seta above andbelow. Spiculose area well developed on dorsocephalichalf of t VIII. Sternite VIII with parallel margins, with2 or 3 long apical setae and a short, triangular vexillumwhich is not spiculose or fimbriated. Acetabular bristlesarising slightly above the middle of the caudal marginof the conical, oblong fixed process on a slightconvexity. Dorsal arm of fixed process rounded dorsally.Movable process clavate, about 2.5X as long as wide atits widest point and with a distinct denticle in the cephalicmargin slightly above the middle. Penis rods convolutedtwice 360°. Crochet bluntly rounded apically, lightlysclerotized.

Female Spermathecal bulga cylindrical, set offfrom the papillate hilla by a slight constriction, bothparts of about equal width. Bursa copulatrix notsclerotized but at least the basal half of the ductusobturatus is. Sternite VII with a slightly undulate marginforming a broad caudal lobe with no ventral sinus. Threeantepygidial setae, the medial 3X as long as the upper,4X the length of the lower bristle. Spiracular fossa2.75X as high as wide, surrounded by ~8 fine setae.Anal stylet 2.2X as long as wide at base, with one longapical seta and two shorter subapical setae on outersurface.

RemarksThis is one of the darkly pigmented members of

the genus. Holland (1959) discussed in some detail acase of teratology involving the modified abdominalsegments of a male of this species from the original typeseries. As indicated below, at least one specimenreported in Clifford et al. (1970) evidently belongs tothis species, and possibly all of those reported by theseauthors.

DistributionClifford et al. (1970) were unable to decide whether

their material from coastal Oregon collections were thisor a related species. We have seen a male from Hunter’sIsland, and although it is extremely poorly prepared, itappears to belong to C. lari. A female from Goat Islandis too poorly mounted to identify. Main & Wallis (1972)reported this species from Ciboux Island, off CapeBreton, Nova Scotia. Holland (1985) added a collection

from Seal Island, Îles de la Madeleine, Quebec, andspecimens of this species are known from Thunder Bay,Ontario, Prince Edward Island and collections fromMichigan. Thus the species seems to be restricted tothe northern regions of North America and will probablyappear in future collections from sea birds.

Host AssociationsHolland (1985) summarized our knowledge of the

host preferences of this species as a parasite of gullsand double-crested cormorants (Phalacrocorax auritus).

Ceratophyllus (Ceratophyllus) niger C. Fox, 1908(Figures 15, 37, 55, 90)

Ceratophyllus niger. C. Fox, 1908, Entomol. News 19:434. USA, California, San Francisco County, SanFrancisco, 37.45N 122.27W, from man, collectorand date not given. Lectotype male, BMNH,designated by Smit & Wright, 1978b: 33.

Ceratophyllus niger inflexus. Jordan, 1929. Novit.Zool. 35: 37, pl. II, Figure 25. USA, Colorado,Custer County, sine loco, no host or date given, T.D. A. Cockerell leg. Holotype female, bydesignation, USNM No. 41425. Synonymized byWagner, 1936b: 201.

Diagnostic CharactersMale With 1 long antepygidial bristle per side

arising on a slight tergal projection, bordered above andbelow by 1 minute seta. Spiculose area well definedand extending along the cephalodorsal half of the marginof t VIII which also bears ~10 marginal and submarginallong setae. Sternite VIII slightly wider in the middle,gently tapering toward the apex which bears 1 long setaand a long, blade-like vexillum that is slightly spiculatebasally on the dorsal surface. Acetabular setae arisingslightly below the middle of the caudal margin of theelongate-oval fixed process which is bluntly roundeddorsally and is 2.5X as high as wide at its narrowestpoint. Movable process clavate, 2X as long as wide atits widest point, its cephalic margin bearing a distinctdenticle about midway down from the apex. Penis rodsconvoluted twice 360°. Crochet slightly curved,moderately sclerotized.

Female Spermathecal bulga cylindrical, sausage-shaped, set off from the slightly papillate hilla by ashallow constriction. Bursa copulatrix sclerotizeddorsally, ductus obturatus with at least the basal halfsclerotized. Caudal margin of st VII a broadly roundedlobe without a sinus. Three antepygidial setae, themedian 4.1X longer than the upper, 5X longer than thelower. Spiracular fossa 3.3X as high as wide and


surrounded by ~8 fine setae. Anal stylet 3X as long aswide at the base, with 1 long apical and 2 shortersubapical setae on the outer surface.Remarks

This is another of the few flea species that havebeen assigned a standardized common name by theEntomological Society of America. It is known as theWestern chicken flea from its association with domesticpoultry. Again, the name is somewhat misleading sincethe species parasitizes a broad range of avian hosts, issometimes taken on small mammals and is reported tofeed readily on humans. As noted in the remarks sectionof the C. idius account, Benton (1998) has suggestedthat these 2 species may interbreed in the northwestwhere the taxa are sympatric. The larva of this specieswas described by Pilgrim and Galloway (2000).

DistributionThis western species is mainly known from the

Great Basin and northwestern North America fromcentral California, north to Alaska. We have seen asingle male from Larimer County, Colorado, east of theRocky Mountains, which is tentatively assigned to thisspecies. There are records from Utah and Montana butmost collections come from the coastal states, BritishColumbia and Alberta and Holland (1985) and Pilgrimand Galloway (2000) provided records from Manitoba.The record originally reported by Stewart (1928) andrepeated by I. Fox (1940b) and Benton (1980) fromNew York state is probably in error, or perhaps it wasbrought in on poultry from the west.

Host AssociationsThis species has been associated with many species

of avian hosts and is also known to feed on a number ofmammals including man. However, there is no evidencethat C. niger can complete its life cycle feeding onmammalian blood alone and to our knowledge thespecies has never been colonized in the laboratory.

Ceratophyllus (Ceratophyllus) pelecani Augustson,1942

(Figures 21, 38, 49, 80)

Ceratophyllus pelecani. Augustson, 1942, Trans. S.Diego Soc. Nat. Hist. 9: 437, Figures 1-2. México,Baja California, Los Coronado Islands, ~32.27N117.17W, from Pelecanus occidentalis, 28.IX.1937,L. W. Walker leg. Holotype female, Allotype male,Los Angeles County Museum, #11 S.D.Zoo, 1937-83. [LACM].

Diagnostic Characters

Male With 1 long antepygidial seta per sidearising on a slight tergal projection and bordered aboveand below by a tiny setula. Spiculose area well definedand extending along the anterior half of the dorsalmargin of st VIII which also bears ~8 marginal andsubmarginal bristles. Sternite VIII curved, with parallelmargins, its apex bearing 2 or 3 long setae and asubapical, awl-shaped vexillum which lacks fimbriationand spicules. The acetabular setae arise on a slight bulgeabout half way up the caudal margin of the oblong-ovalfixed process which is rounded dorsally. Movableprocess of the clasper about 2.5X as long as wide at itswidest point, clavate, its cephalic margin bearing adistinct denticle slightly above the middle. Penis rodsconvoluted slightly more than 360°. Crochet triangularand moderately sclerotized.

Female Spermathecal bulga slightly wider at ductend, sausage-shaped, set off from the hilla, which curvesmore or less at a right angle and bears a large,conspicuous apical papilla. Caudal margin of st VIIwith a median sinus bordered dorsally and ventrally byrounded lobes. Bursa copulatrix moderately sclerotized.Ductus obturatus short but well sclerotized for most ofits length. Three antepygidial setae, the median 2X aslong as the upper, 3X as long as the lower. Spiracularfossa almost 8X as high as wide and surrounded by ~9fine bristles. Anal stylet conical, 2.5X as long as wideat base, with 1 long apical seta and 2 shorter subapicalsetae on the outer surface.

RemarksThe type series of this taxon consisted of 7 males

and 4 females. One pair of paratypes was retained inthe collection of the describer, which now is in the LosAngeles County Museum [LACM], and the remainingmaterial was said to have been deposited in the SanDiego Natural History Museum [SDMC]. The onlyspecimens that we were originally able to examine werethe holotype and allotype. These are poorly mountedand over-cleared, and, as so often with specimensprepared by Hubbard, Augustson and a few other earlyworkers, they are mounted in the reverse of the generallyaccepted position employed by most workers sinceJordan & Rothschild. The drawings presented here aremade from the holotype and allotype but the diagnosticcharacters are based on specimens subsequentlycollected by Dr. Thomas Schwan.

In passing, another irregularity practiced by theauthor of this species is the designation of the femalesex as the holotype. In this group of insects females arenot the diagnostic sex in most genera. Indeed, femalesin many ceratophyllid genera are practically inseparableamong the species unless accompanied by males. As


the late G. H. E. Hopkins once opined, “... Since otherAmerican authors have also followed this practice, itmust be in accordance with some theory with which weare not acquainted, but it is particularly unfortunate in agenus in which females are determinable with difficulty,if at all. The holotype should always belong to the moredistinctive sex.” Although this statement was made inreference to the ctenophthalmid genus Catallagia, itcertainly applies to Ceratophyllus as well.

Very little is known about this species. There areonly 2 reports in the literature that shed much light onhost preferences and distribution. Marshall & Nelson(1967) report it from a number of sea birds on SouthFarallon Island, ~37.40N 123.00W, as well as from theEuropean rabbit and humans, but these may bemisidentifications of Ceratophyllus lari. Schwan (1984)reported on his 1980 observations of the species onSoutheast Farallon Island where it parasitizes Larusoccidentalis. While the paper deals with another speciesof flea, he mentions the unpublished observation thatC. pelecani readily feeds on white mice in the laboratory.

DistributionBeyond the type locality off the western coast of

Mexico, there are few records of this species and all but1 of these are off the coast of California and Oregon.There is a single record from British Columbia in Tatum(1971).

In passing it should be noted that the exact typelocality has not been determined with certainty. In afootnote under Actenopsylla suavis Jordan & Rothschild,1923, Hopkins & Rothschild (1953) noted that Jellison& Good (1942) suggested that the type locality for thisspecies is the Los Coronados Islands, off the extremenorthwest tip of Baja California. They appear to haveoverlooked a Coronados Island in the Gulf of Californiaat about 26.00N. Even so, they are probably correctsince P. aleuticus, the host species of A. suavis, and oneof the host species of C. pelecani, is known to breed onthe Los Coranados Islands, but not off the east coast ofthe peninsula For this reason we have changed the mapcoordinates to agree with those given for Actenopsyllasuavis.

Host AssociationsThe few collections of this species suggest that it is

primarily a parasite of sea birds, and collections fromother hosts are probably accidental.

Ceratophyllus (Ceratophyllus) petrochelidoniWagner, 1936

(Figures 16, 39, 56, 91)

Ceratophyllus petrochelidoni. Wagner, 1936a, Z.Parasitenk. 8: 655, Figure 2. Canada, BritishColumbia, Kamloops, 50.39N 120.24W, fromPetrochelidon lunifrons [=pyrrhonota], VII.1934,G. J. Spencer leg. Holotype male, allotype female,CNIC 4551.

Diagnostic CharactersMale With 1 antepygidial bristle per side borne

on a slight projection and bordered dorsally andventrally by single very small setulae. Spiculose arearelatively narrow and restricted to the anterior 1/2 ofthe dorsocephalic margin of t VIII, which also bears ~7marginal and submarginal setae along its dorsal margin.Sternite VIII with parallel margins, its apex bearing 2or 3 long setae and a long, awl-shaped, curved, subapicalvexillum bearing 2 to 4 small spicules near its base onthe dorsal margin. Acetabular setae arising low on thecaudal margin of the columnar-conical fixed process ona slight caudal process. Apex of fixed process flexedcaudally. Movable process of the clasper rectangular,about 3.3X as long as wide and with pronounced angleson the dorsocephalic and ventrocephalic margins. Witha distinct denticle on the upper quarter of its cephalicmargin that fits into a fovea on the apex of the elongatedfixed process. Penis rods convoluted over twice 360°.Crochet triangular, sclerotized, especially proximally.

Female Spermathecal bulga sausage-shaped, setoff from the papillate hilla by a slight constriction. Bursacopulatrix slightly sclerotized dorsally, ductus obturatussclerotized throughout most of its length. Caudal marginof st VII variable, from bearing a distinct, smoothlyrounded lobe that is not subtended by a sinus to havinga vestigial sinus. Three antepygidial setae, the medial6.3X as long as the upper, 5.7X as long as the lowerbristle. Spiracular fossa 5X as high as wide. Anal styletabout 1.6X as long as wide at base, with 1 long apicalseta and 2 shorter subapical bristles, the lowermost abouta third longer than the upper.

RemarksThis is 1 of the North American species that stands

out with unique genital characters in the male. Althoughthe vexillum is similar to that in C. pelecani, rossittensis,gallinae, rauschi, enefdeae, and calderwoodi, none showthe small spicules on the dorsal margin, at least inmaterial examined by us. However, both celsus andniger do have a few spicules on the dorsal surface of thebase of the vexillum. The apex of the upper lobe of stIX is also much more expanded in this species. Certainlythe 2 most conspicuous characters involve the clasper.The apex of the upper lobe of the fixed process is bentcaudally so as to engage with the denticle in the cephalic


margin of the movable process. The latter is roughlyrectangular, with the widest portion slightly belowmidway down the cephalic margin, whereas the denticlearises about a quarter of the way down from the apex,where the process is noticeably narrower. As isfrequently the case, identification of the females in theabsence of accompanying males can only be tentativeat best since many of the swallow hosts of members ofthis genus may be simultaneously parasitized by morethan 1 species of flea. The larva of this species wasdescribed by Pilgrim and Galloway (2000).

In checking the published information on the typesin the original description it was discovered that datafor the holotype male and allotype female had beenreversed in Wagner (1936a). The holotype male isactually from Kamloops and bears the data given above.The allotype female is from Chilcotin, British Columbia,and bears the data given in Holland (1949), Smit &Wright (1965), Holland (1985) and Lewis (1990).

Hopla (1965a), in his account of C. riparius,mentioned a record in the United States NationalMuseum of 5 females identified as C. petrochelidoniand he indicated that he was confident that they were C.riparius. The computerized inventory of the USNMlists these specimens but provides no collection data.In view of the fact that Holland (1985) did not list thisspecies from Alaska, it seems likely that these specimensare misidentified and that they are in fact C. riparius.

DistributionThis is another western species and Traub et al.

(1983) and Holland (1985) combined have records fromas far north as the British Columbia / Yukon border,south to Alberta, Montana, Wyoming, Utah andCalifornia. Haas et al. (1972) and Wheeler et al. (1970)provided records from New Mexico; Baird & Saunders(1992) from Idaho; Nelson (1972) from Nevada; Jellison& Senger (1976) from Oregon and Wyoming. Therecord of this species from South Dakota (Easton, 1982)was incorrect and actually refers to C. celsus.

Host AssociationsAlthough the species has been collected on a few

other birds, i.e. Passer domesticus, Hirundo rustica andSialia sialis, the preferred host is Petrochelidonpyrrhonota .

Ceratophyllus (Ceratophyllus) rauschi Holland, 1960(Figures 19, 40, 52, 89)

Ceratophyllus rauschi. Holland, 1960, Can. Entomol.92: 792, Figures 6-8, 10. Canada, Yukon Territory,48.3 km N. Stewart River crossing on Klondike

Highway, ~63.25N 136.45W, from nest of Colaptesauratus, 16.VIII.1959, G.P. Holland & J.E.H.Martin leg. Holotype male, Allotype female, CNICNo. 7154.

Diagnostic CharactersMales One antesensilial seta per side, bordered

above and below by tiny vestigial bristles. Spiculosearea present along most of the dorsal margin of t VIIIwhich also bears a few marginal and submarginal setaealong dorsal margin. Sternite VIII with roughly parallelmargins, 3 apical setae per side and a long, narrow,blade-like vexillum that lacks spicules and fimbriation.Acetabular setae arising on a slight acetabular processon the caudal margin of the conical fixed process whichis pointed dorsally and 3.2X as high as wide. Movableprocess clavate, widest in the middle, 2.8X as long aswide at its widest point and with a denticle in the middleof its cephalic margin. Penis rods are coiled throughtwo 360° convolutions. Crochet pointed, slightlydecurved apically, lightly sclerotized.

Female Spermathecal bulga slightly wider at theduct end. The slightly papillate hilla is set off from thebulga by a shallow constriction. Bursa copulatrixslightly sclerotized dorsally. Ductus obturatus wellsclerotized throughout most of its length. Sternite VIIwith an undulate caudal margin but no lobe or sinus.Three antepygidial setae, the median 5.7X as long asthe upper, ~4.6X as long as the lower bristle. Spiracularfossa ~4X as high as wide, surrounded by ~15 fine setae.Anal stylet 2X as long as wide at base.

RemarksHolland (1960) compared this species to

Ceratophyllus niger, another western flea, pointing outthat the movable process was narrower than in C. niger,that there were fewer marginal setae on t VIII, and thespiculose area was less well developed. He alsocommented that the females of C. rauschi generallytended to be smaller and less setose but that they werenot reliably distinguishable from those of C. niger.

Although the original description placed the typelocality on the Alaskan Highway and this was repeatedin Holland (1985), Galloway et al. (1989) corrected thisto read Klondike Highway. While it may be mainly aparasite of flickers and other hole-nesting birds, thereare collections from domestic chickens, as well asturkeys. Further collections should clarify the nature ofits host associations.

DistributionHolland (1985) reported additional collections from

Alberta, Alaska, British Columbia and the Yukon


Territory. All Canadian records are from east of theRocky Mountains.

Host AssociationsThe type series of this species consisted of 3 males

and 11 females, all from the nest of a flicker (Colaptessp.). Though Holland (1985) reported 1 male from aCanada jay (Perisoreus canadensis) and 3 males and17 females from a turkey (Meleagris galloparvo), healso listed an additional 12 males and 29 females fromYukon Territory and Alaska, also from flickers. Hestated that the species apparently infests poultry onoccasion.

Ceratophyllus (Ceratophyllus) riparius Jordan &Rothschild, 1920

(Figures 17, 41, 58, 78)

Ceratophyllus styx. Rothschild, 1900, Novit. Zool. 7:543, pl. IX, Figures 5, 7, 8, 16. England, Hampshire,Lyndhurst, 50.52N 01.34W, from Sand martin[=Riparia riparia] nest, 11.VII.1900, G. Tate leg.Lectotype male, BMNH, designated by Smit, 1955:352.

Ceratophyllus riparius. Jordan & Rothschild, 1920,Ectoparasites I: 71, Figures 67-69. USA, Wisconsin,Milwaukee County, Bayview, Milwaukee, 43.03N87.56W, from nest of Riparia riparia, 9.V.1911,R.A. Muttkowski leg. Holotype male, BMNH.

Ceratophyllus riparius concinnus. Scalon, 1977, Zool.Zh. 56: 1820-1825. USSR, Sakhalin, sine loco, fromRiparia riparia, no date or collector. A nomennudum.

Diagnostic CharactersMale One long antepygidial seta per side arising

on a slight projection, bounded dorsally and ventrallyby 1 (occasionally 2) small setulae. Spiculose areareduced, confined to the dorsocephalic half of t VIIIwhich also bears ~60 setae marginally andsubmarginally. Sternite VIII slightly wider distally thanproximally, its caudal margin with a row of setae on itsdistal half, its apex bearing a cluster of 3 or 4 long setaeper side, and an apical, cone-shaped, fimbriatedappendage subtended by a short, triangular, unfimbriatedvexillum. Acetabular setae arising about half way upthe caudal margin of the columnar fixed process whichis 4.1X as high as wide at its narrowest point. Movableprocess of clasper clavate, about 2.3X as long as wideat its widest point, its apex rounded, its cephalic marginwith an inconspicuous denticle. Penis rods convolutedno more than 360°, frequently less. Crochet sclerotized,tapering to a rounded tip apically.

Female Spermathecal bulga crescentric, sausage-shaped, set off from the papillate hilla by a slightconstriction. Bursa copulatrix well sclerotized, as is theproximal half of the ductus obturatus. Caudal marginof st VII with a broad, rounded lobe subtended by adeep sinus, the margin of which may be slightlyincrassate in some specimens, though never as much soas in C. scopulorum. Spiracular fossa of t VIII 3.3X ashigh as wide and surrounded by ~8 fine setae per side.Anal stylet about 3X as long as wide at base, with asingle long apical seta and 2, occasionally 3 shortersubapical bristles.

RemarksAfter a thorough comparison of North American

material with collections from various localities inEurope, including England, we are convinced that C.riparius is sufficiently different from the European formsof C. styx to justify its treatment as a full species

Certainly the most detailed account dealing withthis species is that of Hopla (1965b) in his treatment ofAlaskan fleas. An abridged review of this discussionfollows.

Although many bank swallow colonies are infestedwith only C. riparius, others may yield small numbersof C. celsus and Hopla speculated that future workersmight report C. idius from colonies of this host as well.He pointed out that some colony sites are shared with afew violet-green swallows and even kingfishers. Whilethe latter have no flea that is specific to them, they maybecome infested by fleas of other ground-hole nestingbirds.

With respect to behavior, Hopla observed that,unlike most fleas, individuals of this species appearedto be positively phototaxic. In the field he noted thatlarge numbers of adults were encountered near theburrow entrances in late summer after their hosts haddeparted. When nests were placed in Berlese funnels inthe laboratory, it was necessary to tape the reflectors tothe funnels to prevent the adults from escaping. This heattributed to positive phototaxis, although an equalpossibility is that they may have simply been movingtoward a source of warmth.

The life history of C. riparius in Alaska is not unlikethat reported for C. styx styx Rothschild, 1900, by Batesand Rothschild (1962) in England. Ceratophyllusriparius has but a single generation in the far north andthis is almost certainly the case for colonies farther south.With young on the wing in 18 to 22 days after hatchingand an incubation period of 14 to 16 days, hosts areonly present for a little over a month. Fleas producedduring this short breeding season evidently remain inthe cocoon until the following spring unless disturbed,


and can endure below freezing temperatures during thewinter months, emerging in the spring prior to or uponthe return of their hosts.

Hopla also noted that Alaskan bank swallows, unlikethose reported by Bates and Rothschild (1962), useestablished burrows over a period of several years,although these may be somewhat “renovated” annuallyby further excavation. Thus, while newly excavatedburrows may be only 2.5 feet deep, older ones may becloser to 4.5 feet in depth.

Approximately 2,000 adults of this species wereplaced in a large glass container containing 1/2 inch ofsand, covered with a tightly woven cloth and placed ina cold room at 45° F and a relative humidity of 75%.There they remained from September to the followingMarch. When removed to a warmer temperature theywere still very active and readily infested 2 two-weekold chickens. Several fed, some for as long as 1.5 hours.Some of these deposited fertile eggs from which larvaeemerged, but none of them completed the life cycle tothe adult stage. In other experiments, Hopla was able tomaintain adults at 60° F for as long as 18 months withlittle or no mortality as long as relative humidities of 70to 80% were maintained. The larva of this species wasdescribed by Pilgrim and Galloway (2000).

DistributionThe map in Haddow et al. (1983) show this taxon,

as C. styx riparius, extending from northeasternKamchatka, western Mongolia and Tuva ASSR, southto Sakhalin Island and Hokkaido, Japan. NorthAmerican records extend from across northern UnitedStates and southern Canada, west to Alaska. The specieshas not been reported from much of the range of itspreferred host, including much of northern Russia andCanada, as well as much of central and southern UnitedStates.

Host AssociationsThe bank swallow (Riparia riparia) is the primary

host of this flea but it has been taken on a few otherswallows, as well as a few accidental hosts, includingsmall mammals. For example, deer mice (Peromyscusmaniculatus) frequently inhabit old bank swallow nestburrows and may subsequently become infested.

Ceratophyllus (Ceratophyllus) rossittensis Dampf,1913

(Figures 22, 42, 53, 95)

Ceratophyllus rossittensis. Dampf, 1913, Schr. Phys.-Ökon. Ges. Königsb. 53: 345, Figures 1-3, 6.Germany, East Prussia, 8 km from Rossiten Bird

Observatory, environs of Ulmenhorst, ~50.13N06.59E, from nest of Corvus [corone] cornix,8.IV.1912, A. Dampf leg. Current location of typesunknown.

Ceratophyllus swansoni. Liu Chi-ying, 1935, Ann.Entomol. Soc. Am. 28: 121, Figures 1-3. USA,Minnesota, Polk County, Fertile, 47.31N 96.19W,from Asio [otus] wilsonianus, 8.V.1931, G.Swanson leg. Holotype male, Allotype female,deposited in the University of Minnesota, St. Paul,MN [UMSP].

Diagnostic CharactersMale One long antepygidial seta per side arising

on a slight caudal projection, bordered dorsally andventrally by single minute setae. Spiculose area poorlydeveloped, restricted to the dorsocephalic third of t VIIIwhich also bears ~12 marginal and submarginal bristles.Sternite VIII widest in the middle, tapering apically, itsapex usually with 2 or 3 bristles and a subapical vexillumarising below the apex that is long, narrow, pointed andlacking fimbriation or spicules. Acetabular bristlesarising slightly below the middle of the roughlytriangular fixed process on a distinct marginal convexity.Movable process clavate, 2.6X as long as wide at itswidest point. Marginal denticle poorly developed.Crochet broad, bluntly pointed and lightly sclerotized.

Female Spermathecal bulga cylindrical, slightlywider than the papillate hilla which is set off from thebulga by a shallow constriction. Sternite VII with asmoothly rounded caudal margin lacking a projectionor sinus. Bursa copulatrix unsclerotized. Ductusobturatus well sclerotized for most of its length. Penisrods convoluted about twice 360°. Dorsal margin of tVIII with about 12 marginal and submarginal setae.Three antepygidial setae, the medial 3X longer than thedorsal seta, 3.25X longer than the lower. Spiracularfossa 3.2X as high as wide, surrounded by ~12 fine setae.Anal stylet 2.75X as long as wide at base.

RemarksThere is little literature concerning this species in

North America. It was originally described from Europe,and a form of it from Minnesota was subsequentlydescribed by Liu (1935). It belongs to the gallinae groupof species but as Holland (1985) indicated, males of thistaxon may be separated by the proximal rather thanapical origin of the vexillum on st VIII. While this mayassist in identification, a review of the origin of thisstructure in related species reveals that it is notcompletely apical in a number of species in the gallinaegroup. However, this character, in conjunction withother genitalic features, especially the vestigial spiculose


area on the male t VIII and the contours of the othermodified segments, coupled with its host associationspermit relatively easy identification. On the other hand,females in the absence of accompanying males caneasily be mistaken for females of other members ofthis species group. Accompanying males and corvidhosts provide the best means of determination.

DistributionHaddow (1983) reported this species to be

amphiberingian. In North America it is known fromTraub’s (1983) Rocky Mountain, Alleghenian/PlainsSector, with isolated records from Montana, Minnesota,Manitoba and Ontario.

Host AssociationsThis is a specific parasite of crows, Corvus spp. In

North America the host of choice is the common crow(Corvus brachyrhinus) but it may also be associated withraptors that use abandoned crow nests.

Ceratophyllus (Ceratophyllus) scopulorum Holland,1952

(Figures 7, 43, 61, 81)

Ceratophyllus scopulorum. Holland, 1952, Can.Entomol. 84: 297, Figures 1-7. Canada, YukonTerritory, Rampart House, 67.25N 140. 59W, fromPetrochelidon pyrrhonota nests, 24-26.X.1951, R.L. Rausch leg. Holotype male, Allotype female,CNIC No. 15895.

Diagnostic CharactersMale Antepygidial setae 1 per side arising on a

slight pedistal and bordered dorsally and laterally bysingle minute setulae. Spiculose area well developedand occupying most of the dorsal margin of t VIII whichalso bears 12 or 13 marginal and submarginal setae.Margins of st VIII parallel, its apex with 4 or 5 longbristles and a long, blade-like vexillum the dorsal marginof which is fimbriated from its base to apex. Penis rodscompleting about 1.5 complete convolutions.Acetabular setae arising very low on the fixed processfrom a distinct convexity. Movable process 3X as longas wide at its widest point, fusiform in lateral view andbearing a denticle about 1/5 down the cephalic marginthat engages in a fovea on the apex of the upper lobe ofthe elongated fixed process. Fixed process columnar,extending almost to the apex of the movable process.Upper lobe of st IX with a distinct subapical concavityin the cephalic margin above which the structure isbroadly rounded. Penis rods convoluted ~1.5X 360 °.Crochet well sclerotized, narrow and pointed.

Female Spermatheca sausage-shaped, the bulga setoff from the slightly narrower, strongly papillate hillaby a slight constriction. Bursa copulatrix unsclerotized,ductus obturatus with at least the basal half sclerotized.Caudal margin of st VII with a broad lobe subtended bya deep sinus the margin of which is strongly sclerotized.Three antepygidial setae, the median 4.2X the length ofthe upper, 6.3X the length of the lower seta. Spiracularfossa on t VIII 3X as high as wide and surrounded by~8 small setae. Anal stylet 2.25X as long as wide atbase, with 1 long apical seta and 2 subapical bristles onthe outer surface, the upper only about 3/4 as long asthe ventral.

RemarksAmong the North American ceratophyllids, this is

one of the most easily recognized, at least in the males.The fusiform configuration of the movable process isduplicated nowhere else in the genus. The dorsal lobeof the fixed process is unlike any other in the genus, asis the vexillum of st VIII, the apex of st IX and the shapeof the crochet. In females, while this is not the onlyspecies in which the caudal margin of st VII is lobedand incised, it is the only species in which the margin ofthe incision is so strongly incrassate. The larva of thisspecies was described by Pilgrim and Galloway (2000).

DistributionHolland (1985) listed a number of collections from

Alaska and the Yukon Territory and along the border ofthe latter with British Columbia. He also reported thespecies from Kent Island, Grand Manan, NewBrunswick. His map number 61 shows records fromManitoba. Eads et al. (1979) also listed collections fromnear Hallock, Kittson County in extreme northwesternMinnesota, a record not mentioned by Benton & Timm(1980), and Larson (1997) reported the species fromPembina County, North Dakota.

Host AssociationsAlthough the New Brunswick records are from both

Petrochelidon pyrrhonota and Hirundo rustica, exceptfor occasional accidental associations, this is a specificparasite of the former. The lack of additional publishedrecords from central Canada is certainly an indicationof lack of collecting, and further collections may beexpected from the northern tier of the United States.

Ceratophyllus (Ceratophyllus) vagabundus(Boheman, 1866)

(Figures 6, 44, 62, 92)

Pulex vagabunda. Boheman, 1866, Ofvers. K. Vet.-


Akad. Förh., Stockh. 22: 576, pl. 35, Figures 1, 1A.Norway, Svalbard Archipelago, Spitzbergen, CapTodsen, ~78.30N 15.00E, jumping about on theground, 11.VII.1864, D. Malmgren leg. Typedeposition unknown.

Diagnostic CharactersMale One antepygidial bristle per side bordered

dorsally and ventrally by single small setulae. Spiculosearea well developed along entire dorsal margin of t VIIIwhich also bears ~15 marginal and submarginal setae.Sternite VIII with parallel margins, its apex bearing acluster of ~4 long bristles and a fimbriated, cone-shapedprojection subtended by a blade-like vexillum the dorsalmargin of which is also spiculate. Acetabular setaearising from a slight convexity below the middle of thecaudal margin of the oblong-oval fixed process, the apexof which is roundly pointed. Movable process roughlyrectangular, 3X as long as wide at its widest point, itsapex obliquely angulate, its cephalic margin with a welldeveloped denticle slightly below the middle. Apex ofapical arm of st IX gently curved caudad, its marginssubparallel. Penis rods convoluted about 1.5X times360°. Crochet narrow, pointed and well sclerotized.

Female Spermathecal bulga curved, sausage-shaped, slightly narrowed at duct end, set off from thestrongly papillate hilla by a slight constriction. Bursacopulatrix and ductus obturatus both sclerotized. Caudalmargin of st VII smoothly rounded, lacking a lobe orsinus. Three antepygidial setae, the medial 2.75X aslong as the upper, 3.1X as long as the lower. Spiracularfossa 3.5X as high as wide, surrounded by ~7 fine setaeAnal stylet 2.75X as long as wide at base, with 1 longapical seta and 1 subapical seta which is over 1/2 aslong as the apical bristle.

RemarksAlthough reported from a number of host species,

this is a flea that primarily attacks hosts which nest onthe ground, rocks or cliff faces. It is sometimes taken inthe nest burrows of sea birds and may occasionally beencountered in rodent burrows, although there are noreports of its breeding there. More probably these adultswere searching for suitable avian hosts and their presencein rodent burrows was accidental.

DistributionThis circumpolar species is widely distributed along

the northern coasts and on the islands of the northernseas, as well as in many mountain systems in theNorthern Hemisphere. Ioff & Scalon (1954) attributedthe Asian form of this flea to C. v. orientalis Wagner,1930a, a name later synonymized with the nominate

taxon by Smit & Wright (1965). Ceratophyllus v.insularis Rothschild, 1906, and C. v. alpestris Jordan,1926, are forms still recognized by some workers. Theformer has been applied to material from the British Isles,Belgium and northern Germany, the latter to thepopulation found in Switzerland. In Asia, the species isknown from the Caucasus, Tyan Shan and Altaimountain ranges, as well as Transbaikalia and Mongolia.In the Western Hemisphere it seems to be limited to thehigh Arctic of Alaska, Canada and Greenland accordingto (Holland,1985).

Host AssociationsThis species prefers birds that nest in colonies on

the ground or on rocky cliffs along the northern coastsin the Northern Hemisphere, or at high elevations in theinland mountain ranges. As Holland (1985) pointed out,the primary hosts include gulls, ducks, certain sea birdsand raptors such as hawks, falcons and eagles.

Subgenus Emmareus Smit, 1983Ceratophyllus Curtis, 1832 - Emmareus subgen. n. Smit,

1983: 14.Ceratophyllus (Emmareus) Smit, 1983. Smit, 1983: 36.

The subgeneritype is C. (E.) columbae (Gervais,1844).

Diagnostic CharactersHead Frontal tubercle present, more conspicuous

in the males. Eye well developed and heavily pigmented.Frontal setal row of 4 to 6 setae extending up the cephalicmargin of the antennal fossa in males, sometimes fewerin females, and never extending as high. Ocular row of3 setae in both sexes. Postocular seta present butsometimes difficult to see in darkly pigmented species.Females with a few of the setae that arise on the antennalpedicel extending beyond the apex of the clavus. Maleswith 15 to 20 small setulae extending along the dorsalmargin of the antennal groove, females with 10 to 15.Preoccipital setae usually 3, arranged in 2 rows of 1 and2. Occipital row of 4 or 5 long setae per side plusintercallaries.

Thorax One to 4 pseudosetae per side under themesonotal collar, usually 3. The mesepisternum usuallywith 1 or 2 long setae per side plus 5 to 12 fine setulaescattered irregularly over the surface, especiallydorsally. The mesepimeron bears 4 to 7 long bristlesarranged in 2 irregular rows. The metanotum usuallybears 1 small but clearly visible marginal spinelet perside. The lateral metanotal area usually bears 2 setaeper side, the dorsalmost the longest. Metepisternumwith 1 seta per side. Metepimeron with 5 to 8 longsetae arranged in 3 more or less vertical rows.


Legs Outer surface of forefemur with a numberof small setae and the basal 1/2 to 2/3 of the mid- andhindcoxae is devoid of setae. There is a small seta nearthe base of the forefemoral guard seta, 5 notches in thecaudal margin of the foretibia bearing heavy bristles,and 5 pairs of lateral plantar setae, none of which isnoticeably shifted on to the plantar surface. Thiscondition is found in both sexes on all 3 pairs of legs.Midtibia with 5 caudal notches, hindtibia with 6.

Abdomen Abdominal tergal spinelets per side:males usually I, 2; II, 2; III, 1-2; IV 0-1. Females usuallyI, 1-2; II, 2; III, 2; IV, 1. The remaining characters aresexual in nature, involving the genital segments.

Male Wagner’s organ present. In males of thissubgenus the most important characters involve theapophysis of st IX and the penis rods, both of which areshorter than in the nominate subgenus and do notcomplete more than a 360° convolution. With 3antesensilial setae, the median long, the laterals minute.The spiculose area on t VIII is weakly developed in C.diffinis and C. garei, better developed in C. borealis.There are 2 acetabular setae but there is no welldeveloped acetabular proceass.

Female Caudal margin of st VII smoothly rounded,lacking a lobe or sinus. In females of this subgenus thebursa copulatrix is not sclerotized but the ductusspermathecae is dilated and wrinkled basally. Theductus obturatus is only weakly sclerotized, if at all,and the spermathecal duct inserts into the bulga at anangle of at least 45° to the long axis. The bulga isfusiform, widest in the middle, tapering both apicallyand distally. Hilla and bulga about equal in length, thehilla lacks a well developed apical papilla. Anal stylet~2X as long as wide at base with 2 subapical setae whichare almost as long as the apical seta.

RemarksThis is one of the eight new subgenera erected by

Smit (1983) and one of the six subgenera assigned byhim to Ceratophyllus. Three of these we believe shouldbe returned to or elevated to the status of full genera,but Emmareus, Celeophilus and Ceratophyllus areretained here in their original context. Eight species ofCeratophyllus are assigned to this subgenus, 3 of whichare known from the area under consideration. All areparasites of birds and are usually associated with speciesthat nest on or near the ground and have moist to wetnests.

Key to the species of the subgenus Ceratophyllus(Emmareus)

(Modified from Holland, 1985)

1 Male st VIII as broad or broader than movableprocess (Figure 26); vexillum of st VIII a broad lobebearing a ventral, finger-like projection (Figure 68);crochet with a sharp-pointed apex; female st VII with abroadly rounded apex with or without a subtending,shallow sinus (Figure 75) (Holarctic) ............. garei

Male st VIII narrower than movable process (Figure25); vexillum of st VIII vestigial (Figure 67) or semilunar(Figure 66) crochet less sharply pointed; female st VIIwith a slight to conspicuous, pointed caudal lobe (Figures73, 74) .................................................................... 2

2 Spiculose area of male t VIII narrow, less than halfthe width of the movable process, vexillum of st VIIIusually small (Figure 67); caudal lobe of female st VIIrounded but more pointed apically and subtended by ashallow sinus (Figure 74) (Nearctic) ........... diffinis

Spiculose area of male t VIII broad, more than halfthe width of the movable process; dorsal lobe of fixedprocess with a distinct subapical, caudal lobe (Figure24); vexillum large, semilunar (Figure 66); aedeagalcrochet narrow with a rounded apex (Figure 66); caudallobe of female st VII conspicuous, pointed, triangular,subtended by a shallow sinus (Figure 73) (Holland’s1949 figure of the female st VII of C. diffinis moreclosely approximates that of C. borealis and vis. vis.)(Holarctic) .................................................. borealis

Ceratophyllus (Emmareus) borealis Rothschild, 1907(Figures 24, 46, 66, 73)

Ceratophyllus borealis. Rothschild, 1907,Entomologist’s Mon. Mag. 43: 11, 1 Figure.Scotland, St. Kilda Island, Outer Hebrides, 57.49N08.34W, from gannet nest [=Sula bassanus],VII.1906, N. H. Joy leg. Holotype female, bymonotypy, BMNH.


dorsally and laterally by single minute setulae. Movableprocess elongate-clavate, 3.1X as long as wide, with awell developed denticle. Fixed process columnar, itsapex oblique, 3.6X as long as wide at its widest point,its dorsocaudal angle forming a pronounced lobe in thearea of the fovea. Acetabular bristles arising low on thecaudal margin of the fixed process. Spiculose area welldeveloped on dorsocephalic half of t VIII. Dorsal marginof t VIII with 3 to 5 marginal bristles. Sternite VIIIwith subparallel margins, apex usually with 1 to 3bristles, 1 longer than the other 2. The apex also bearsa large, crescentric, spiculose vexillum. Crochet curvingto a narrow, rounded point. Apical lobe of distal arm of


st IX with parallel cephalic and caudal margins, curvedcaudad.

Female Spermathecal bulga fusiform, widestsubapically, hilla slightly shorter than bulga and bearinga slight apical papilla. Sternite VII with a distinct lobeon its caudal margin. Antepygidial setae 3, the medial3.6X the length of the upper, 4.4X the length of thelower. Two or 3 fine setae adjacent to the spiracle on tVIII. Spiracular fossa ~2.7X as high as wide at widestpoint. Anal stylet 2.25X as long as wide.

RemarksIn many respects, this species is similar to C. (E.)

garei, both morphologically and biologically. The malegenitalia are similar, differing mainly in the form of thefimbriated apex of st VIII. In C. borealis, this structureis crescentric and conical while in C. garei it is almostsquare and bears a finger-like ventral appendage. In C.garei, the fixed process of the clasper is more or lessconical, while in C. borealis, there is a subapical bulgein the dorsocaudal margin. Also, in the latter speciesthe movable process is widest subapically, the apex ofst IX is obliquely squared, its caudal margin stronglyconcave and the spiculose area along the dorsal marginof t VIII is well developed. In C. garei, the apical halfof the movable process is of uniform width, the apex ofst IX less curved and the tergal spiculose area is poorlydeveloped. In the females the spermathecal bulga tapersgradually into the hilla and lacks a conspicuousconstriction between these 2 structures in C. garei whilea constriction is present in C. borealis. The caudalmargin of st VII in C. borealis usually bears a short,pointed lobe that is subtended by a shallow sinus, whilein C. garei the ventrocaudal margin of this sternite isstraight or slightly convex. Haddow et al. (1983)indicated the presence of intergrades between these twospecies in Greenland, Norway, Sweden, Holland,Germany and Great Britain, and the species is figuredand discussed in detail by M. Rothschild (1948, 1955).In the latter work Rothschild suggested in a footnotethat C. frigoris Darskaya, 1950, was morphologicallyvery close to C. garei. Darskaya’s material, collectedin the Caucasus Mountains and Isti-su, Azerbaidjan,“appears to be very similar, if not identical with, theintermediates between C. borealis and C. garei” thatRothschild had just discussed. However, there are sofew collections from North America that little light canbe shed on the subject of interbreeding with otherspecies, much less the validity of C. frigoris as a separatespecies.

Biologically, the two appear to share similarecological requirements. Both parasitize a broad rangeof passerine birds that nest on or near the ground, in

crevices or under rocks and thus have moist to wet nests.Species of Oenanthe (wheatears) seem to be thepreferred hosts of this species although it has been takenfrom many other birds, particularly those nesting nearwater. Ceratophyllus borealis, like C. garei and C.gallinae, tends to disperse widely from the nest of thehost where it has overwintered. As a result it has alsobeen reported from a number of ground-feeding species.

DistributionThis species is distributed in Traub’s (1983)

Forested and Northern Canadian Sector of the NearcticRegion. While it is common in the Northern Hemisphereof Eurasia, it is known only from Greenland and theYukon Territory in the Western Hemisphere.

Host AssociationsThis is a parasite of birds which nest on rocks, cliffs

or under stones in the northern parts of the world. Whileit has a distinct preference for passerine hosts such aswheatears (Oenanthe oenanthe), pipits (Anthusspinoletta) and snow buntings (Plectrophenax nivalis),it is also associated with sea birds such as gulls, ternsand gannets.

Ceratophyllus (Emmareus) diffinis Jordan, 1925(Figures 25, 47, 67, 74)

Ceratophyllus diffinis. Jordan, 1925, Novit. Zool. 32:111, Figure 44. Canada, British Columbia,Okanagan Falls, 49°21’N 119°33’W, fromColymbus holboelli [=Podiceps grisegena],20.IV.1913, G. Garrett leg. Holotype male, bymonotypy, BMNH.

Ceratophyllus rileyi. Liu, 1935, Ann. Entomol. Soc. Am.28: 123, Figures 4-6. USA, Minnesota, Pine County,from Bonasa umbellus, 16.X.1931, H. E. Wallaceleg. Holotype male, UMSP. Synonymized byJordan, 1937: 263.


dorsally and laterally by single minute setae. Movableprocess 3X as long as wide, clavate, widest subapicallyand with a denticle in the cephalic margin. Fixed processcolumnar, its apex rounded, 3.8X as high as wide.Acetabular bristles arising low on the caudal margin ofthe fixed process. Spiculose area poorly developed onthe dorsocephalic half of t VIII. Only 3 or 4 marginalbristles along the dorsum of t VIII. Sternite VIII withsubparallel margins, tapering to a point apically, narrow,with 2 or 3 long apical bristles per side, of unequal lengthand arising 1 behind the other. Vexillum vestigial,


usually a low, triangular, convexity at the apex of st VIII.Crochet deflected downward, tapering to a roundedpoint. Apical lobe of distal arm of st IX with parallelmargins, its caudal margin only slightly concave.

Female Spermatheca fusiform, widest in themiddle, its dorsal margin straight or slightly concave,set off from the apapillate hilla by a conspicuousconstriction. Three antepygidial setae, the medial 5Xthe length of the upper or lower. Caudal margin of stVII with a distinct tringular lobe subtended by a shallowsinus. With 5 or 6 small setulae adjacent to the spiracularfossa on t VIII which is 3.5X as high as wide at its widestpoint. Anal stylet 2X as long as wide at base, conical inform.

RemarksThere is little in the literature about this species.

As can be seen from Holland’s (1985) host list it hasbeen collected from a number of piciform andpasseriform birds and their nests. He stated that in spiteof the fact that the type was taken from a grebe, thespecies seldom occurs on water birds [unlike C. garei]but prefers hosts which nest on or near the ground.Haddow et al. (1983) pointed out that another bird flea,Dasypsyllus gallinulae perpinatus (Baker, 1904) alsooccurs in nests in bushes and trees in open woodlandfrom sea level to ~600m and seems to replace C. diffinison the west coast and in the Rocky Mountains. Thelarva of this species was described by Pilgrim andGalloway (2000).

DistributionBenton and Timm (1980) note records from

Hubbard and Pine counties in Minnesota, and also listeda record from Hubbard County, Minnesota. Except forone record from Rockingham County, Virginia andanother from Long Island, New York, all of Benton’sremaining records are from northeastern New York andNew England. Traub et al. (1983: map 37) showedBenton’s records, plus additional collections fromQuebec, Ontario and Newfoundland on the east, andthree records from British Columbia in the west. Chilton& Galloway (1995) reported the species fromsouthwestern Alberta and Holland (1985: 298) reportedadditional collections from southeastern Canada,including Ontario. He also showed additional collectionsfrom British Columbia and a number of records fromAlaska. He went on to state that this species and C gareihave similar distributions in North America, but C.diffinis tends to occur farther south.

Host AssociationsThis species prefers birds that nest on or near the

ground in bushes and trees in open woodlands. Beyondthat, it tends to be somewhat catholic in its tastes andhas been taken on a broad range of hosts.

Ceratophyllus (Emmareus) garei Rothschild, 1902(Figures 26, 48, 68, 75)

Ceratophyllus garei. Rothschild, 1902, Entomologist’sMon. Mag. 38: 225, pl. IV, Figures 1-3. England,Hertfordshire, Tring, 51.48N 00.40W, 200 m, fromnest of Gallinula chloropus, VII.1902, N. C.Rothschild leg. Lectotype male, BMNH,designated by Smit & Wright, 1978b: 20.

Ceratophyllus utahensis. Chapin, 1919, Bull. BrooklynEntomol. Soc.14: 60. USA, Utah, mouth of BearRiver, ~41.29N 112.10W, from Steganopus[=Phalaropus] tricolor and Spatula [=Anas]clypeata, 22.VII.1916, A. Wetmore leg. Holotypemale, Allotype female, USNM No. 29372.Synonymized by Jordan & Rothschild, 1920: 69.

Ceratophyllus quebecensis. I. Fox, 1940a, Proc.Entomol. Soc. Wash. 42: 65, pl. 10, Figures 3-4.Canada, Quebec, St. Mary’s Island, 50.12N59.46W, from eider down, 30.VI.1938, H. S. Petersleg. Holotype male, USNM No. 53587.Synonymized by Holland (1949).

Diagnostic CharactersMale One antepygidial seta per side arising on a

slight caudal projection of t VII and bordered dorsallyand laterally by minute setulae. Movable process 3.5Xas long as wide at its widest point. A denticle is presenton its cephalic margin. Fixed process columnar, roundlypointed apically, 3.75X as high as wide. Acetabularbristles arising on a projection low on the caudal marginof the fixed process. Spiculose area well developed onthe dorsocephalic half of t VIII, its dorsal margin bearing3 to 5 long setae. Sternite VIII with cephalic marginalmost straight, caudal margin strongly convex, its apexwith 3 setae, 1 of which is longer than the other 2.Vexillum enormous, and roughly oval or quadrate inshape, its dorsal surface spiculate, its ventral margin witha narrow, appendiculate appendage. Crochet with asharp apex, not as deflected as in the preceding species.Anterior and posterior margins of apical lobe of distalarm of st IX subparallel, its posterior margin mildlyconcave.

Female Spermatheca with a gibbose, fusiformbulga, widest subapically and set off from the apapillatehilla by a slight constriction. Caudal margin of st VIIwith a smoothly rounded lobe subtended at most by ashallow sinus. Three antepygidial setae, the median3.9X as long as the upper, 5.5X as long as the lower.Seven to 10 small setulae adjacent to the spiracular


fossa on t VIII, its fossa 4.7X as high as wide. Analstylet 2X as long as wide at widest point, stronglyconical.

RemarksThe similarities between this species and C. borealis

have been discussed under the account of that species.For a more detailed comparison of the two species thereader is referred to the works of M. Rothschild (1948,1955).

This is another species that has a somewhatconfusing nomenclatural history. A possible explanationis that it is sympatric over much of its range with relatedspecies with which it may hybridize. Specifically, weare referring to C. borealis and C. diffinis within thearea under consideration, both of which are thought tointerbreed occasionally with C. garei. Whether theseare true hybrids in the sense that they are sterile andcannot reproduce, or some interbreeding takes placeamong mixed populations, is still speculative. However,individuals with intermediate characters probablyprovide the basis for most of the junior synonyms. Onlymolecular techniques or breeding studies are likely toresolve this issue, and to our knowledge none of thesethree species has ever been colonized under laboratoryconditions. Benton (1998) reported a similarphenomenon between C. idius and C. niger; C.scopulorum and C. celsus, and C. celsus and C.petrochelidoni. The species accounts of these taxashould be consulted for further details. The larva ofthis species was described by Pilgrim and Galloway(2000).

DistributionThis is an Holarctic, circumpolar species the range

of which extends southward through Canada and Alaskainto the northern and western United States in NorthAmerica. In Eurasia, it is known from France and theCzech Republic, east to Kamchatka, Mongolia, theRussian Maritime Territories and the Japanese islandsof Honshu and Hokkaido.

Host AssociationsThis species has been taken from a broad range of

hosts but shows a distinct preference for birds that neston or near the ground and thus have moist nests. Thespecies has also been taken on a number of small rodents,though we suspect that these were acquired whilesearching for a suitable avian host.

Acknowledgments

We wish to thank the authorities of the

Biosystematics Research Institute, Agriculture -Canada, Ottawa, Ontario, and particularly Dr. EvertLindquist for the loan of material, including primarytypes. The primary type of C. coahuilensis was loanedby Dr. Brian Brown of the Los Angeles CountyMuseum, Los Angeles, California. Ms. Nancy Adamsof the Smithsonian Institution, Washington, D. C., andMs. Theresa Howard of the Natural History Museum,London, also loaned types from their institution’scollections. Additional assistance was rendered by Dr.Thomas Schwan of the Rocky Mountain Laboratory,Hamilton, Montana. We also wish to thank Dr. JoelCoats, Chairman of the Entomology Department at IowaState University for his continuing support. JournalPaper No. J-19011 of the Iowa Agricultural and HomeEconomics Experiment Station, Ames, IA, Project No.3100, and supported by Hatch Act and State of Iowafunds.

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Figures 1-4 Hind femora of C. (Cel.) zhovtyii, C. (Cel.) adustus, C. (Cer.) hirundinis and C. (E.) columbae.Figures 5-7 Male claspers.


Figures 8-16 Male claspers.


Figures 17-26 Male claspers.


Figures 27-38 Apex of male sternite IX.


Figures 39-48 Apex of male sternite IX.


Figures 49-56 Apex of male sternite VIII and crochet.



Figures 71-72 Spermathecae of C. adustus and C. zhovtyii.


Figures 73-75 Sternite VII and spermathecae of C. borealis, C. diffinis and C. gareiFigures 76-77 Sternite variation in C. adustus and C. zhovtyii.


Figures 78-83 Sternite VII and spermathecae.

a taxonomic review of the ceratophyllus curtis, 1832 of north … · 2017-10-11 · december, 2001...

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