Phalacropsylla paradisea Rothschild, 1915, Paradise [Cochise County], Arizona, off Epimys sp. [= Rattus], Mus sp., and civet cat, collected in September, October, November, and December 1913 by Otto C. Duffner. [Note: Early collectors often referred to small sylvatic rodents as “Mus” and reference to the “civet cat” in southern Arizona likely refers to the ring-tailed cat (Bassariscus astutus, Lichtenstein) and not to skunks of the family Mephidae.]

Frons broadly rounded, without frontal tubercle. Inter-antennal suture (falx) well developed in both male and female. Antennal groove shallow, opened posteriorly. Antenna asymmetrical, extending onto prosternosome in male, female antenna shorter. Margin of pedicel with short setae, none extending much beyond base of clavus. Occipital area with three oblique rows of setae. Pre-antennal area (anterior to eye) with two rows of setae. Head lacking setae below or posterior to eye. Eye elliptical and pigmented; central unpigmented sinus present. Eye contiguous with two overlapping, darkly pigmented spines; lateral anterior spine broader and shorter than longer narrow mesal spine. Maxilla very elongated, extending half the length of forecoxa. Labial palpus long, extended to or beyond apex of trochanter. Pronotum with complete row of long setae anterior to 14–18 broad, bluntly pointed ctenidial spines. Mesonotal collar with several pseudosetae per side. Pleural arch well developed [an unusual characteristic for a true nest flea, Eads and Maupin (1982), Lewis and Maser (1978), and Barrera and Traub (1967)]. Suture dorsad to lateral mesonotal area expanded into a distinct rounded incrassation at posterior margin abutting pleural arch. Meso- and metasterna protruding downward producing a lobe between coxae (especially so in metasternum). Meso- and metacoxae with three long stout setae at apico-caudal margin. Fore tibia with six dorsal notches; mid- and hind tibiae each with seven dorsal notches. Distotarsomeres each with four pairs of plantar bristles with a fifth pair shifted onto plantar surface between first proximal lateral pair. More anterior terga with small marginal pigmented spinelets. Terga with two rows of slender setae; anterior row small and posterior main row long with intercalaries. Abdominal spiracles blunt at apex. Three long antesensilial bristles; middle bristle longest of three. Sensilium slightly convex; with 12 sensilial pits per side. Eighth tergum of female with caudal lobe bearing marginal row of five to seven stout long setae and six to eight short stout, more anterior setae. Eighth tergum reduced in male. Caudal margin of female S-VII with rounded dorsal lobe subtended by broad sinus. Bulga of spermatheca pyriform with slender hilla much longer than bulga. Duct of bursa copulatrix narrow, longer than spermatheca, sclerotized, with apical hyaline bursa copulatrix. Male distal arm of S-IX club-shaped; broader at apex than proximal portion. Always bearing various arrangements of spiniform setae near apex. Male basimere very broad and robust; manubrium narrow and elongated with parallel sides. Basimere not divided into lobes but may or may not possess a sinus on ventral margin. Telomere narrow (at least five times as long as widest dimension); fovea variously placed on dorsal margin. Aedeagus structurally narrower at apex than at middle. Crochet and dorsal armature on sclerotized inner tube present.

Phalocropsylla allos males lack a well-defined sinus in the apico-ventral margin of the basimere above acetabulum (Fig. 2). This feature is shared only with P. oregonensis (Fig. 4) and P. morlani, but may be distinguished from the former by the absence of long bent spiniform setae on the apical margin of the distal arm of S-IX and from P. morlani by the presence of a hyaline lobe on anterior margin of distal arm of S-IX (Fig. 3) and an acutely pointed crochet. The lobe on the caudal margin of S-VII of the females of P. allos and P. nivalis is each longer than broad, whereas the lobes of other species are broader than long. The ratio of P. allos is 1.9 times as long as broad, whereas P. nivalis is only 1.5 times as long as broad.

Figures 2–5. 

2 Phalacropsylla allos male basimere and telomere. Arrows define ventral margin of basimere without sinus 3 Phalacropsylla nivalis male basimere, telomere, distal arm of S-IX, and aedeagus. White arrows define margin of sinus in ventral margin of basimere. AA = angular apex of basimere; HL = hyaline lobe of distal arm of S-IX 4 Phalacropsylla oregonensis, male holotype basimere, telomere, and distal arm of S-IX. Arrow indicates small sinus in ventral margin of basimere 5 Phalacropsylla paradisea, male basimere, telomere, and distal arm of S-IX. Arrows define margin of deep sinus on ventral margin of basimere. Scale = 0.2 mm

USA: Arizona, San Francisco Mts., 18 X 1989, G.E. Haas, 4♂, 4♀ (BYUC). California, Siskiyou County, N. cinerea (Ord) nest, 9 XII 1976, B.C. Nelson, 4♂, 2♀; N. cinerea, 31 I 1980, C.R. Smith, 1♂ (USNM). Idaho, Idaho Falls, National Reactor Testing Site (NRTS), Bonneville County, N. cinerea, 20 VIII 1967, D.E. Beck (1♂, 1♀); same data except rodent nest, 17 II 1967, (1♂); Onychomys leucogaster (Wied-Neuwied), 22 IX 1966, 2♀ (BYUC). Montana, Jefferson County, Morrison cave near White Hall, Neotoma sp., 31 XII 1940, H.B. Mills, 1 ♂; Ennis, in cave, Neotoma nest, II 1941, W.L. Jellison and G.M. Kohls, 1 ♂, 2♀ (CMNH); 1♂, 1♀ (BYUC). Nevada, Clark County, Spring Mts., N. cinerea nest, 5 VI 1985, 2♂, 2♀ (BYUC). New Mexico, Sandoval County, W edge Valle Grande, 2637 m, Jemez Mts., N. cinerea nest, 18 IX 1970, Animal Ecology Research Unit (AERU), 20♂, 16♀; Cibola County, N side Ice Cave Canyon, 2424 m, Jemez Mts., N. cinerea nest, 21 IX 1970, AERU, 2♂, 1♀; Sierra County, SW side Cerro del Medio, 2652 m, Jemez Mts., N. cinerea nest, 21 IX 1970, 1♂, 2♀; W edge Valle Grande, 2622 m, Jemez Mts., N. cinerea ♂, 25 IX 1970, 1♂; SE corner Baca location no. 1, Line 17, rocks with Dome Meadow, Neotoma mexicana Baird ♀, 16 X 1970, 1♂; Lincoln County, midden mix, 6 VIII 1991, G.E. Haas, 2♂, 1♀ (BYUC). Utah, Beaver County, Delano Ranger Station, Peromyscus maniculatus (Wagner), 25 VI 1957, D.M. Allred, 1♂, 1♀; Utah County, Provo, woodrat nest, 13 XI 1948, N.C. Acraia, 11♂, 13♀, (BYUC); same data except 2♂, 3♀ (CMNH), N. cinerea, 12 XI 1949, Allan Dotty, 1♂, 1♀ (USNM); Provo, Rock Canyon, N. cinerea nest, 15 IX 1949, V.J. Tipton, 3♂, 1♀; Provo, woodrat nest, 13 XI 1948, N.C. Acraia, 1♂, 2♀, (CMNH); N. cinerea nest, 12 XI 1949, A. Doty, 14♂, 21♀; Provo, Rock Canyon, N. cinerea nest, 13 XI 1948, 1♀; Provo, woodrat nest, 6 XI 1948, N.C. Acraia, 1♀; Provo, Rock, Canyon, N. cinerea nest, 24 XI 1949, D.M. Allred, 4♀; Provo, Rock Canyon, N. cinerea nest, 24 II 1951, D.E. Beck and D.M. Allred, 3♂; Provo, Buckley’s Mine, N. cinerea nest, 21 X 1950, D.M. Allred, 1♀; East of Provo, N. cinerea nest, 30 III 1951, D.E. Beck and D.M. Allred, 4♂, 3♀; Provo, N. cinerea, 25 XI 1948, N.C. Acraia, 2♂, 3♀ (BYUC).

Phalacropsylla allos is the most widely spread species of Phalacropsylla, occurring in southern British Columbia, Arizona, central to northern California, Colorado, Idaho, Montana, Nevada, New Mexico, Utah, and Wyoming (Fig. 1). It is sympatric with P. paradisea in Arizona, Colorado, and New Mexico. The vast majority of specimens examined were recorded from N. cinerea throughout it range, with single records from N. mexicana (1♂), Onychomys leucogaster (2♀), and P. maniculatus (1♂, 1♀). No other species of Phalacropsylla have been collected from N. cinerea (Table 1). Most of the specimens from nests of N. cinerea were collected during the cooler fall and winter months from September through February. Only one collection of three males was reported in a warmer period (June) and this site was inside a cool mine shaft (Buckley Mine) located at an elevation of 2896 m. We did not examine specimens of P. allos reported by Eads and Campos (1982) from N. mexicana (1♂), Peromyscus difficilis (J.A. Allen) (3♂, 1♀), P. maniculatus (1♂, 1♀), Reithrodontomys megalotis (Baird) (1♀) from Larimer County, Colorado; however, these specimens reported from the eastern slopes as P. allos by Eads and Campos (1982) were collected from March through August during the warmer months at elevations from 1600–1900 m. Although they collected during all months of the year, P. allos specimens were not collected from September through February. This seasonal disparity is enigmatic and warrants future collecting and studies.

Figure 1. 

Distribution of Phalacropsylla species in the Canada, Mexico, and the Western United States. Arrows define same locality for two species.

The apico-ventral margin of the basimere is entire, without a sinus, a feature shared only by P. allos and P. oregonensis (Figs 2, 4). Readily distinguished from both by the shape of the distal arm of S-IX which is wider basally than at apex. The apex of the distal arm of S-IX (DA9) also lacks modified long spiniform setae, as in P. allos, whereas P. oregonensis possesses modified setae (Fig. 4). The female of P. morlani has not been described.

New Mexico: Santa Fe [Santa Fe County, elev. 3048 m], Ochotona princeps (Richardson), 10 X 1958 [10 XI 1958 was recorded by Eads and Campos (1982:241)], H.B. Morlan, holotype (USNM).

The description of P. morlani was based on one male from Santa Fe County, although Eads and Campos (1982) indicated that a second male had been lost. The discrepancy in the month the holotype was collected (October or November) was not resolvable. Eads and Campos (1982) stated correctly in their diagnosis that P. morlani has a pronotal comb of 18 teeth but erroneously recorded 20 in their key (couplet 6.). Although P. allos is documented from adjacent Bernalillo and Sandoval counties, P. allos has not been found in Santa Fe County. Dedicated collection efforts are needed to determine if P. allos and P. morlani are sympatric at high elevations in Santa Fe County and to collect and describe the female and additional males of P. morlani. Since P. morlani was collected from a pika (O. princeps), additional collecting might best be directed at collecting both O. princeps and N. cinerea in Santa Fe County.

Males of P. nivalis are separable from other species of Phalacropsylla except P. paradisea by the lack of a sinus in the apico-ventral margin of the basimere above the acetabulum (Figs 3, 5). Males are further distinguished from P. paradisea by the absence of long curved modified spiniform setae on the apex of DA9 (Figs 3, 5). See diagnosis of P. allos to differentiate females.

Mexico, State of Mexico, [Mirador del Poeta, N slope Mt.] Popocatépetl, 300 m SW Tlamacas, [~19.02°N, 98.38°W] 3900 m, s/Neotoma [Neotoma mexicana torquata], 19 IX 1963, A. Barrera, holotype ♂, allotype ♀ (USNM). Río Frío, 3100 m, N. mexicana, 17 I 1965, T. Álvarez and A. Barrera, paratype ♂; Popocatépetl, Mirador del Poeta, 3900 m, Peromyscus melanotis J.A. Allen and Chapman, 5 VII 1964, A. Barrera and T. Álvarez, paratype ♀, (MZFC-S).

Known only from type material from Popocatépetl mountain, Mexico. Specimens were taken from two different hosts: Neotoma and Peromyscus. Phalacropsylla nivalis is the most extreme southern species of the genus, occurring many hundreds of kilometers from its closest allied species, P. paradisea.

Males differ from P. paradisea and P. nivalis by the absence of a sinus on the apico-ventral margin of the basimere. A small sinus is indicated, but its depth is much less than its width (Fig. 4). Readily differs from P. allos and P. morlani by the lack of modified long spiniform setae at the apex of DA9 (See diagnosis for P. allos). Of the two species whose lobes on the margin of S-VII are wider than long, the lobe of P. oregonensis is more triangular, bluntly pointed, and curved downward than that of P. paradisea. The latter is broadly rounded at apex.

USA: Idaho, Bonneville County, NRTC, Idaho Falls, P. maniculatus, 21 X 1966, [D.E. Beck], coll. code: 36HF, 1♂ (BYUC). Oregon, Malheur County, Succor Creek State Park [~43.28°N, 117.08°W], Neotoma lepida Thomas, 15 V 1975, C.O. Maser, holotype ♂ USNM No. 75247; same data except P. maniculatus, allotype ♀ [MWH re-mounted holotype and allotype, as original medium was crystalized] (USNM).

Allred (1968) recorded one male from P. maniculatus collected at the National Reactor Testing Station in southern Idaho as P. paradisea but was misidentified and is herein referred to P. oregonensis. This specimen and the type series from eastern Oregon are the only known representatives of this species. In addition to the specimens examined, P. oregonensis was also collected from Peromyscus crinitus (Merriam) (Lewis and Maser 1978) (Table 1). The medium in which the holotype and allotype specimens was originally mounted was crystalized, obstructing the specimens from view. The specimens were removed from the slides with xylene, remounted in Canada balsam, and are now adequately preserved.

Males of P. paradisea and P. nivalis each possess a deep sinus on the ventral margin of the basimere (at least as deep as wide) that separates both from other species of Phalacropsylla. Further separated from P. nivalis by the presence of long modified spiniform setae on DA9 which are absent in P. nivalis (Figs 3, 5). See diagnostic features of females for P. oregonensis above.

Mexico: Nuevo León, Cerro Potosí, 3050 m, rodent nest, 20 IV 1964, V.J. Tipton et al., P. hamata holotype ♂ (USNM). USA: Arizona, Apache County, Neotoma mexicana, 13 XI 1973, W. Begay, 1♂ (USNM). Cochise County, China Point, Dragoon Mts., Neotoma stephensi Say and Ord nest, 1 X 1993, G.E. Haas, 1♂; China Point, Dragoon Mts., 19 VI 1994, G.E. Haas, 1♂, 1♀; Chiracahua, nr Paradise, N. albigula nest, 23 IX 1989, G.E. Haas, 8♂, 14♀; Dragoon Mts., N. albigula nest, 26 IX 1989, G.E. Haas, 1♂; Paradise Cemetary, Chiracahua Mts., N. albigula nest, 19 X 1994, G.E. Haas, 1♂, 5♀ (BYUC); Paradise, Mus sp., XI 1912, O.C. Duffner, 1♀, R. Traub no. B-1330 (CMNH); Paradise, Mus sp., IX 1913, O.C. Duffner, 1♂ lectotype; Paradise, Epimys sp. = Rattus sp., XI 1913, O.C. Duffner, 2♀ paralectotype; Paradise, Mus sp., 12 III 1913, O.C. Duffner, 2♂ paralectotypes; Paradise, “civit cat”, 10 IX 1913, O.C. Duffner, 1♀ paralectotype (BMNH). Coconino County, Bixler Mt., N. mexicana, 23 IX 1993, G.E. Haas, 3♂, 4♀; Williams, Neotoma nest, 13 IX 1981, G.E. Haas, 1♂, 2♀; Ben Williams, Neotoma nest, 20 IX 1981, G.E. Haas, 1♀; Site W-3, NNW Williams, N. stephensi nest, 16 X 1989, G.E. Haas, 3♂, 1♀; SE Flagstaff, Neotoma nest, 19 XII 1981, G.E. Haas, 3♂, 3♀; Haulapai, host unknown, I 1986, G.E. Haas, 3♂, 5♀ (BYUC). Graham County, Pinaleno Mts., host unknown, 18 V 1990, G.E. Haas, 2♂, 2♀; Pinaleno Mts., vole nest, 20 X 1990, G.E. Haas, 11♂, 10♀; Shannon Park, Pinaleno Mts., N. mexicana nest, 10 XI 1989, G.E. Haas, 1♀; Stockton Pass, Pinaleno Mts., host unknown, 22 XI 1989, G.E. Haas, 1♀; data missing except leg. G.E. Haas, Pinaleno Mts., 1♀ (BYUC). Greenlee County, Dipodomys merriami Mearns 16 XI 1938, 1♂, 1♀ (CMNH), 1♂, 1♀ (USNM). Navajo County, north of Show Low, N. albigula nest, 30 IX 1989, G.E. Haas, 2♂, 1♀ (BYUC). Colorado, Montezuma County, Mesa Verde National Park, Peromyscus truei (Shufedit), 20 X 1961, C. Douglas, 3♂, same data except 25 X 1961, 1♀, 26 X 1961, 2♂, 3♀, 24 XI 1961, 1♂, P. truei/maniculatus, 13 X 1961, 1♀ (BYUC). New Mexico, Bernalillo County, N. albigula, 20 II 1981, Curt Montman, 1♂; same data except 4 XI 1981, 1♀ (USNM, previously identified as P. hamata); Catron County, Ben Lilly camp ground, Mogollon Mts., N. mexicana nest, 23 IX 1991, G.E. Haas, 2♂, 1♀; Snow Canyon, N. mexicana nest, 28 IX 1996, G.E. Haas, 15♂, 18♀; Bear Wallow, Tamiasciurus hudsonicus (Erxleben) nest, 1 X 1998, G.E. Haas, 1♀ (BYUC). Guadalupe County, 6.5 km S of Santa Rosa, N. albigula, 9 X 1951, 1♂ (USNM). Hidalgo County, Peloncillo Mts., N. albigula nest, 23 III 92, G.E. Haas, 1♀; Peloncillo Mts., N. albigula nest, 24 III 92, G.E. Haas, 1♀; the Pass, Peloncillo Mts., N. albigula nest, 25 III 92, G.E. Haas, 3♂, 10♀ (BYUC). Texas, Brewster County, Big Bend National Park, 1737 m, Peromyscus pectoralis Osgood ♂, 2 XI 1963, V. J. Tipton et al., 2♀ (BYUC).

Tipton and Mendez (1968) described P. hamata from one male from Cerro Potosí, Nuevo León, Mexico. Eads and Maupin (1982) described the female of P. hamata from two specimens collected from Bernalillo County, New Mexico and considered an additional four males as P. hamata. These were collected from Peromyscus leucopus (Rafinesque) and N. albigula. With the recent accession of the Glenn E. Haas flea collection (now part of the BYUC), many specimens of Phalacropsylla were available for study from the vast areas of Arizona, Colorado, New Mexico, and Texas. These specimens were identified as either P. hamata or P. paradisea. While studying this material, it was impossible to distinguish females accurately from one or the other. To date, males have been distinguished primarily by the presence of various numbers of long spiniform setae on the apico-ventral margin of the distal arm of S-IX. The diagnosis of the male of P. hamata provided by Tipton and Mendez (1968) included: 1) A deep sinus in the caudal margin of the immovable process of the clasper (basimere), 2) four long bristles on the apical and subapical portion of basimere, 3) two hook-like spiniform setae near the apex of the distal arm of S-IX, and 4) the bifid portion of the proximal arm of S-IX as angulate. The sinus on the caudal margin of the basimere, long bristles on the apex of basimere, the number of hook-like spiniform setae present at the apex of the distal arm of S-IX, and shape of the bifid portion of proximal arm of S-IX, each proved to be quite variable within series from the same study sites and even among specimens from the same host. Based on these comparative studies, we concluded that P. hamata is not a valid species and consider it to be a junior synonym of P. paradisea. Phalacropsylla paradisea is representative of the genus in the southern portion of its distribution with records ranging from central Arizona, southern Colorado, through New Mexico, Texas, and into northern Mexico. Phalacropsylla nivalis is the only species occurring further south than P. paradisea. Although P. allos is the most commonly collected species of Phalacropsylla, P. paradisea has been collected from a much more diverse group of rodent host species (n = 10) (Table 1).

During studies on plague in the Western United States by the U.S. Army in the mid-1970s, the junior author (MWH) identified two specimens (previously unreported) of P. paradisea that were collected on N. albigula (one specimen among 37 hosts examined) and Peromyscus boylii (Baird) (one from 10 hosts examined) from Fort Huachuca, Cochise County, Arizona. Although the whereabouts of these two specimens are unknown, they are documented in unpublished reports of the U.S. Army Environmental Hygiene Agency-Regional Division West, Aurora, Colorado.

In the latter years of his life, Dr. Glenn Haas concentrated his studies on the fleas in nests of small mammals, primarily the nests of Neotoma and arboreal Tamiasciurus. He placed the nests in breathable paper grocery bags, maintained humidity with moist paper towels, and meticulously hand-picked the emerging adult fleas over a period of weeks and months. Thus many of his mounted specimens were teneral and often not yet expanded from their recent pupal state. These “rearing” studies document the importance of species of Neotoma, particularly N. albigula, as the primary hosts of P. paradisea.