Brigham Young University Science Bulletin, Biological Series

Volume 7 Number 2 Article 1

6-1966

Siphonaptera () of the Nevada Test Site

D Elden Beck Department of Zoology and Entomology, Brigham Young University, Provo, Utah

Dorald M. Allred Department of Zoology and Entomology, Brigham Young University, Provo, Utah

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Recommended Citation Beck, D Elden and Allred, Dorald M. (1966) "Siphonaptera (fleas) of the Nevada Test Site," Brigham Young University Science Bulletin, Biological Series: Vol. 7 : No. 2 , Article 1. Available at: https://scholarsarchive.byu.edu/byuscib/vol7/iss2/1

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JUL 28 1966 HARVwr>J UNIVERSITY

Brigham Young University Science Bulletin

SIPHONAPTERA (FLEAS) OF THE NEVADA TEST SITE

by D ELDEN BECK and DORALD M. ALLRED

BIOLOGICAL SERIES — VOLUME VII, NUMBER 2

JUNE 1966 BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN BIOLOGICAL SERIES

Editor: Dorald M. Allred, Department of Zoology and Entomology, Brigham Young University, Provo, Utah

Associate Editor: Eabl M. Christensen, Department of Botany, Brigham Young University, Provo, Utah

Members of the Editorial Board:

J. V. Beck, Bacteriology C. Lynn IIayward, Zoology W. Derby Laws, Agronomy

Howard C. Stutz, Botany

Wilmer W. Tanner, Zoology, Chairman of the Board Stanley Welsh, Botany

Ex officio Members

RudgerRudgerH.H. Walker, Dean, College of Biological and Agricultural SciencesScienci

Ernest L. Olson, Chairman, University Publications

The Brigham Young University Science Bulletin, Biological Series, publishes acceptable papers, particularly large manuscripts, on all phases of biology.

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Science Bulletin

SIPHONAPTERA (FLEAS) OF THE NEVADA TEST SITE

by

D ELDEN BECK

and

DORALD M. ALLRED

BIOLOGICAL SERIES — VOLUME VII, NUMBER 2

MAY 1966 TABLE OK CONTENTS

Page

INTRODUCTION 1

LITERATURE REVIEW 1

METHODS AND PROCEDURES 1

ACKNOWLEDGMENTS 3

SCHEME OE CLASSIFICATION 3

FAMILIES AND SPECIES 3

SPECIES PRESENTATION 4 Echidnophaga gallinacea 4

Ptdcx irritans 4 Cediopsylla inaequalis 4 Hoplopsyllus anomalu.i 6

Atyphloceras echti ...... 6

Epitedia tvenmanni ...... 6 Catallagia decipiens ..9 Meringis dipodomys 9

Meringis parkeri .... 11 Meringis hubbardi 11

Jordanopsylla allredi II Stenistomera alpina 13

CaUistopsyllus deuterux .... 13

Megarthroglossus procus .... 13

Anomiopsyllus amphiholus ...... 13 RlmdiiuipM/lIu heiseri 15 Rhadinopsylla sectilk 15 Carteretta carteri 15 Thrassis hacchi .....18 Thrassis aridus 18

Dactyhpsylla bluei ... IS Foxclla ignot/i 20 Diamanus montanus 20 Orchopeas sexdentatus ... 20

Monopsyllm wagneri ...... 22 Monopsyllus eumolpi 22

Malaraeus telchinum .... 22 Malaraeus sinomus 22 Malaraeus euphorbi 24 Amphipsylla neotomae 24 OdontopsyUus dentatus 24 Peromyscopsylla hesperomys ...... 24 Nyctcridopsylla oarwouverensU ... 24

CONCLUSIONS 26

LITERATURE CITED 20 MSI (il III IMIi VI IONS

Figure Page

1 Map of the .southeastern corner ol Nevada showing the regional location of thi Nevada rest Sit< 2

2 Extent "i thi majoi plant communities ol the Nevada ["esl Site 2

igraphii distribution nl Echidnophaga gullinacca, Pulex irritant, and Cediopsylla inequalix 5

i Geographii distribution ol Hoplopsyllus tmomalits 7

Geographic distribution ol Uyphlocera echis Epiteiliu wenmanni, and Catallagia tlecipieru H

i' Geographic distribution ol \feringw diftodoniys and Weringh ptirkeri 10

7. Si-. ivun.il occurrenci ol WeringAi dipadamys I]

s Geographic distribution ol Weringw huhbardi, lordanopsyUa allredi, and Stenixtomera alpina 12

I Geographic distribution ol Cullistopsyllm deuterus, Anomio)>xyllus amphiholws, Wegarthroglossus procus, .iiul Rhadiuopsylla lu iseri 14

10. Geographic distribution nl Rbadinopsylla sectilia and Carteretta carteri 16

11 Geographic distribution ol Thrtuma hucchi IT

12 Seasonal occurrence "I Tbrassis bacchi .... is

Geographic distribution ol Thrassu aridi-i, Foxella ignota, Dactylopsylltt bluet, and Dianianin mon- tanus 19

It Seasonal occurrence ol Thrassis aridh 20

IS Geographic distribution ol Orchopeat texdentatti* 21

16. Geographic distribution ol Monopsyllua wagneri, Malaraetm tclchinum, Monopsyllui eumolpi, and

Malaraeus rinomus 2 ;

!". Geographic distribution ol Malaraeus euphorbi, Odontapvyllm dentat us, Nycteridopsylla oancou- oerensis, Amphipsylla neotomae, and Peromyscopsylla hesperomys .

SIPHONAPTERA (FLEAS) OF THE NEVADA TEST SITE 1

by

D Elden Beck and Dorald M. Allied

INTRODUCTION

Ecological studies at the Nevada Test Site 1963; Beck, Allred & Brinton, 1963; Allied, 1963; near Mercury, Nevada, were begun in 1959 as Allred & Beck, 1963. 1964; and others—see list a cooperative project between the United States of references ) Atomic Energy Commission and the Brigham This report deals with data on the fleas Young University Department of Zoology and collected at the Nevada Test Site. Geographi- Entomology. Initial research was directed to cal and ecological distribution, host relation- mammals, birds and reptiles (Jorgensen & Hay- ships, and seasonal occurrence are emphasized. ward, 1965; Hayward, Killpack & Richards, 1963; Tanner & Jorgensen, 1963). As the investiga- The biotic communities and areas of study were tions continued, however, parasites and other designated by Allred, et al. (1963 a & b), and eonsortes were collected. Reports on some of our references generally follow their classifica- these collections have been published ( Goates, tion (Figs. 1 and 2).

LITERATURE REVIEW

Previous records of fleas from the Nevada eus sinomus from the Deer Mouse ( Test Site are unknown, although there are re- maniculatus) at Searchlight. Hystrichopsylla ports for the state of Nevada. Every record gigas dippiei (most likely //. dippiei dippiei) from Nevada is not made here, but some gen- was collected from the Sonoran Deer Mouse eral ones are worthy of mention. (P. maniculatus sonoriensis) at Charleston Park

Most reports of fleas for Nevada are the on Charleston Mountain near Las Vegas. Orch- result of the extensive work by C. A. Hubbard opeas sexdentatus nevadensis was taken from the Desert Wood Rat (Neotoma lepida) from (1947). Practically all references made by him Clark County. Hopkins and Rothschild (1962) are tor collections made in the northern half of reported a contribution to the British Museum the state near or above the 39th parallel al- of Anomiopsyllus amphibolus taken from the there are some collections southern though from White-throated Wood Rat (Neotoma albigula— Nevada. For example, Hoplopsyllus anomalus most likely .V. lepida) in Nye County. All lo- was taken from the White-tailed Antelope Squir- calities are in the southeastern corner of Ne- rel (Ammospermophilus leucurus), and Malara- vada, relatively near the test site.

METHODS AND PROCEDURES

Most of the fleas were taken from the bodies the houses or other recesses of the Desert Wood of host which were trapped or shot Rat, and the eonsortes extracted by the use of specifically for the puqjose of collecting ecto- a modified Berlese funnel. Data contained with parasites or selected from and birds mammals each collection included date, host or source, which were collected for other purposes. In and the biotic community where found. These each case the host was placed immediately into were coded for computer analysis. a paper sack which was then sealed, data writ- ten on it, and returned to the laboratory. Fleas were retrieved by brushing the fur of In some instances nests were removed from the host which was held under a 75-watt lamp

'1! Y U -A E ( publication No. C00-1355-1+. This work was support ed (in part) by funds from the United States Atomic Energy Commission, department of Zoology and Ento logy, Brigham Young Uni ty, Provo, Utab Bhirham Young University v m s. i Bulletin

SOUTHERN NEVADA

of Map the southeastern comer of Nevada showing the regional location ol the Nevada Test Site. ) ) )

Fleas of the Nevada Test Site

in a deep, white enamelware pan. All con- Specimens were identified from the slide pre- sortes were preserved in 707c ethyl alcohol un- parations, all data then placed on collection rec-

til thev were mounted individually on micro- ord forms, and IBM punch cards prepared to slides to which were applied the collection data. enable computer analysis.

ACKNOWLEDGMENTS

Many technicians were involved in the field Health Service, San Francisco, California. Dr. collection and laboratory preparation of hosts C. Lvnn Havward, Department of Zoology and and parasites. Without their careful work this Entomology, Brigham Young University, Provo, study would not have been as readily accom- Utah, helped with the specific identification of plished. We are most grateful for their unself- mammal and bird hosts. ish, enthusiastic participation. We are especially grateful for the excellent Dr. William L. Jellison, Rocky Mountain laboratory services and transportation facilities Laboratory, Hamilton, Montana, corroborated which were provided by the Civil Effects Test and assisted in most of the determinations. Operations (CETO), Division of the Atomic Dr. Robert Traub, School of Medicine, Univer- Energy Commission, at sity of Maryland, also helped to untangle some Mercury, Nevada. We specific problems in flea determinations. This are likewise grateful for similar conveniences was likewise the case with Frank M. Prince, provided by Brigham Young University, Provo. Communicable Disease Center. U. S. Public Utah.

SCHEME OF CLASSIFICATION

In general we have followed the taxonomic been judiciously inserted in the Hopkins-Roths- and phvlogenetic arrangement described by Hop- child arrangement. The species of fleas from kins and Rothchild (1953, 1956, 1962). Where the Nevada Test Site are listed below. special taxonomic studies have recently been made on specific groups, such as the genus For the hosts we used the "Vernacular Names Thrassis discussed bv Stark (1958) and unpub- for North American Mammals North of Mexico" lished information on Malaraeus by Frank M. as approved by the American Society of Mam- Prince, information as deemed appropriate has malogists (Hall, 1957).

FAMILIES AND SPECIES

Pulicidae Megarthroglossus procus Jordan & Rothschild Anomiopsyllus amphibolus Wagner Echidnophaga gallinacea (Westwood) Rhadinop.ii/lla heiseri (McCoy) Ptdex irritans Linne Rhadinopsylla sectilis Jordan & Rothschild Cediopsi/lla inaequalis (Baker) Carteretta carteri Fox Hoplopsi/llus anomalus (Baker)

Hystrichopsyllidae Ceratophyllidae

Atyphloceras echis Jordan & Rothschild Thrassis bacchi (Rothschild) Epitedia wenmanni (Rodischild) Thrassis aridis Prince Catallagia decipiens Rothschild Dactylopsijlla bluet ( Fox Meringis dipodomys Kohls Foxella ignota (Baker) Meringis parkeri Jordan Diamanus montanus (Baker) Meringis hubbardi Kohls Orchopeas sexdentatus ( Rothschild Jordanopsylla allredi Traub & Tipton Monopsi/Uus wagneri (Baker)

Stcnistomera alpina (Baker) Monopsijllus ettmolpi ( Rothschild Callistopsyllus deuterus Jordan Malaraeus telchinum (Rothschild) BlUGHAM YOUNC 1 'Sl\ I llsIM Si II M 1 I.

Malaraeus tinomus \ [ordan 1

Malaraeus euphorbi i Rothschild Peromyscopsylla hesperomys (Bala

Vmphips> llidae [schnopsyllidae AiiijihijisiiUii neotomae Fox

Odontopsyllus dentatus I Baker) Sycteridopsylla vancouvernensis Wagner

SPECIES PRESENTATION

For each species listed, the following se- Vccording to Stark i L958) and Wheeler and

quence in presentation ol information was fol- Douglas i L945), E. gallinacea has been found

i lowed: a I specific .ind subspecific identity and naturall) infected with plague organisms and other pertinent taxonomic data; (1)) ecological demonstrated a high vector efficiency. ind geographical distribution with maps, (c)

i and

Some hosts at the test site were collected in Most Associations. Most of the 73 specimens greater numbers than others which were taken obtained were collected from the Kit Fox. Other hosts only occasionally during a particular season ol were the Black-tailed Jack Babbit, the Coyote (Cants latrans), and the Western Pipis- the year. Although it is most unfortunate that trelle Bat all hosts at the test site were not collected on a (Pipistrellus hesperus) which repre- sents an unusual collection. Fleas oi this species daily schedule, it was considered not econom- have in ically or conservationallv feasible to do so. been taken most abundance from bur- rowing animals, which may explain the greatest number found on the Kit Fox. Echidnophaga gallinacea Seasonal Occurrence. Apparently fleas ol this (Westwood ). 1875 species are not restricted seasonally, but may be Distribution. A total of 56 specimens was found at any month of the year when their collected at widely separate parts of the site host is active. Fleas were collected in |.muarv. (Fig. 3). Too few specimens were collected to February, May, August, and December. indicate a community predominance. Comments. To better understand the dis- Host Associations. The Black-tailed Jack tribution and seasonal occurrence of /'. irritant Rabbit (Lepus californicus) and the Kit Fox at the lest site, a larger series of predators such

I as the Kit Fox, and Bob ( Li/nx ( Vtilpes maCWtii were the only animals on Coyote, to which fleas ol this species were tound. The rufttS | need be collected. Their dens also fleas wen- about equalh distributed among should be carefully examined. The records those hosts collected. from the bat and rabbit most likely were acci- dental encounters. Seasonal Occurrence. \\\ specimens were found m Vugust and December. CediopsyUa inaequalis (Baker). 1S95 Comments. Sufficient data are not available The male He. is collected indicate that the to ai curatel) delimit geographic boundaries Ex subspecies is C. inaequalis interrupta [ordan tensive rabbit collections were made over sev-

( L925). eral years at various seasons, yet only lour rab- bits were found infested, and then onlj with Distribution. Most of the fleas (45 of 59) a single Ilea ol this species On each rabbit. were taken bom hosts from the Pinv on-|uniper Fleas of the Nevada Test Site

GRAPHIC SCALE. FT.

N 800,000

Tonopah

1 50 miles from Mercury

N 650,000

r I ig. 3. Geographic distribution of Echidnophaga gallinacea -At Pulex irritans and Cediopsylla inequalis £^ Bingham \m si. University Scienci Hi i i i i in communit) on Rainiei Mesa Thirteen were liais Dipodomys microps) Most specimens t.tki-M hum the vicinit) "I Cane and Tippipah 260 ol 286) ol // anomalus were obtained Springs which are in mixed vegetational com- From the White-tailed Antelope Squirrel. munities. One collection was mad* in Seasonal ( kcurrcncc. Most Ilea collections og) in- community i Fig were made in June, xsith lesser encounters in April, collections Host Associations. I In- Rainiei Mesa col Max, and August. Two were In linns were From Nuttall's Cottontail Rabbil made m September and one in Dea mbei (Sylvilagus nuttallii), Thirteen specimens were Comments. Collections ol //. anomalus

.' < < 1 collected from the Desert > 1 1 < > 1 1 . i i 1 Rabbit show them to OCCU] in |une. ssith a total ab- \\ i Sylvilagus audubonii I and one from the hite sence in fuly, and reoccurrence in August. This tailed Antelopi Squirrel, ssas true regardless ol elesation Or host. These Seasonal Occurrence. The greatesl number data should not be interpreted .is conclusive, is seasonal ol specimens was From a Nuttall's Cottontail for there a difference in the activity hosts Rabbit collected in April on Rainier Mesa. All of the from which Fleas ol this species others were collected in November From the have been taken. For example, one should not species Desert Cottontail Rabbit at lower elevations. Compare the seasonal occurrence for a ol Ilia on the Rock Squirrel xsith the- same ( collections omments Extensive of mam- species on the White tailed Antelope Squirrel. at the test site have not been made at mals While the latter host is active during the- win- and collections lower the higher elevations, at ter in the valleys, the Former max be hiberna- i li s limited in some instances. val have been ting in talus covered In snow at a much higher studies ol the abundant mammalian Vlthough elesation. All specimens of S. VariegOtUS xx ere Fauna in the valleys and Foothills have been taken during April. Max', and |unc, xsith the made, collections of cottontail rabbits carefully exception of one collection in August at the desert living in the vicinity of springs on the west side ol Frenchman Flat along the loot- their elimination have been limited to avoid hills. lis The vast areas ot trapping and shooting. According to Stark (1958), lleas of this desert clothed with sage brush (Arte- highland species in New Mexico have been tound natural- misia tridentata) in the western and northwest- Is infected xsith plague organisms. ern part of the test site, and the extensive f*in- von juniper woodland to the north and north- Atyphloceras echis could afford a close natural history west well Jordan and Rothschild, 1915 scrutin) These undoubtedly provide For an The subspecies ol our collections is \. echis extended range For rabbits as well as other mam- mals and their ectoparasites, echis. Distribution. Hosts and their parasites were

Hoplopsy litis anomalus (Baker), 1904 confined primarily to the Coleogyne commun- ity. One specimen was taken in a mixed vege- Distribution. Fleas were taken from hosts tational situation and one in a Pinyon-Juniper in the valleys and foothills, and to a limited ex- community ( Fig. 5). tent on Rainier Mesa. They were most Fre- species quently encountered in Grayia-Lycium, Larrea- Host Association. Eleven lleas of this Franseria, and Coleogyne biotic communities. were taken from the Desert Wood Rat. taken plant Nevertheless, they were in all ol the Seasonal Occurrence. Collections wore made communities except Atriplex-Kochia. This is in [anuary and March, ssith most encounters in unusual, lor the most common host, A. leUCtirUS, December. was Frequently collected from this latter com- Comments. Fleas ot several species are munit) i Fig 1 I. known to live on A. lepida and in its nests. Host Associations. ( >iil\ a Few Rock Squirrels The natural history of the Desert Wood Rat (Spermophilus oariegatus) were taken on Rain- and its parasites at the test site should he cue- ier Mesa in the Pin) on- |nniper community, but lullx studied, for the' rats have a wide range ol .ill possessed lleas. A Roi 11 id tailed Squirrel distribution at the site. Spermophilus tereticaudus) ssas collected in ssest Frenchman Flat, and two (leas were ob Epitedia wenmanni (Rothschild), L904 tamed One Flea was removed From the Little tsso collected, a male Pocket Mouse (Perognathus longimembris) , and Only specimens were eight specimens From several Chisel-toothed and a Female. On the- basis of the male, this Fleas of the Nevada Test Site

Kawich Valley Gold Flat GRAPHIC SCALE, FT. N 950,000 L !

N 800,000

N 750,000

Tonopah

1 50 miles from Mercury N 700,000-^

N 650,000

Fig. 4. Geographic distribution of Hoplopsyllus anomalus BRICHAM VOUNC I'mmiimm Si li. si I Hi ill lis

Geographic distribution ol Atyphloceras

is most likely of the subspecies E. wenmanni site. This is due to the wide distribution and wenmanni. abundance of their common hosts, kangaroo rats. These mammals were under continuous study Distribution. The two fleas were taken at for several years, primarily in the valleys and Tippipah Spring, a mixed type of biotic com- foothills where they are most usually found. munity ( Fig. 5 ). From the standpoint of biotic community dis- Host Association. Both specimens were col- tribution, many collections were from Larrea- letted from a Western Pipistrelle Bat. Franseria with about equal numbers from Cole- ogyne and Grayia-Lycium. The next ranking Seasonal Occurrence. The fleas were collect- community was Salsola, with few collections ed in November. from mixed vegetative communities. Two col- Comments. Main specimens of Peromyscus lections were in Pinyon-Juniper, and one was in spp. and N. lepida were collected during this Atriplex-Kochia (Fig. 6). survey. It is unusual that we did not collect Host Associations. As the specific name of fleas of this species from these hosts. More the flea indicates, the most common hosts are unusual is that the only specimens taken were species of Dipodomys. The Chisel-toothed from a bat. In the original descriptions made Kangaroo Bat is the species on which these in 1904, the male was taken from a White- fleas were most often encountered. The next footed Mouse (Peromyscus leucopus) and the was Merriam's Kangaroo Bat (Dipodomys mcr- female from a Bushy-tailed Wood Hat ( Neotoma riami). Only two collections were taken from cinerea). Most published records show fleas of the large Desert Bat Kangaroo ( Dipodomijs this species to occur on species of Peromyscus deserti ) which is so restricted in its distribution and Neotoma. that not main collections were expected. A single collection was made from Ord's Kangaroo Catallagia decip Both ild, 1915 Bat (Dipodomys ordii). Other animals from which fleas were taken, in order of abundance, Distribution. Hosts of two different species are the White-tailed Antelope Squirrel, the were collected, and one flea was collected from Southern Grasshopper Mouse (Onychomys tor- each. One was in a mountainous vicinity, the ridus), the Desert Wood Bat, the Great Basin other in the Pinyon-Juniper community on Rain- Pocket Mouse (Perognathus parvus), the Deer ier Mesa (Fig. 5). Mouse and the Kit Fox. Host Associations. One host , a Pin- Seasonal Occurrence. Fleas of this species yon Mouse (Peromyscus truei) was from Rain- were not taken in May, July, or August, and ier Mesa; the other, a Desert Wood Bat, was in only one collection was made in June when the vicinity of Tippipah Spring. three males were taken from D. microps. Most Seasonal Occurrence. The collection from collections were made in October and Novem- /'. truei was made in March, and from .V. ber. The common hosts. Dipodomys spp., were lepida in December. present during these months, and many were trapped and examined during May, fune, fuly, few collections of this Comments. The and August as well as in other months. Two species perhaps can be explained on the basis collections were made in September, 32 in Oc- that fewer hosts have been collected at higher tober, and 86 in November. A relatively high elevations at the test site compared to the num- incidence was maintained throughout the fall, ber made in the valleys and foothills. In other winter, and early spring until May (Fig. 7). surveys during the past 20 years and in liter- ature references, the Deer Mouse has been the Comments. Kangaroo rats of several species main host. Others mentioned in the literature and the White-tailed Antelope Squirrel were are mainly mammals whose habitats art 1 at high studied extensively at the test site. This pro- elevations or in cool environs. Peromyscus man- vided an opportunity to examine their ectopara- iculatus at the test site is most abundant in the site fauna on a year-round basis. Pinvon-Juniper community which has not been The disappearance of adult flea populations extensively surveyed by us. during Ma\-, June, July, and August is an enig- ma. The host is active during the summer months, but fleas were not found on those Meringis dipodomys Kohls, 1938 examined during that period. One might sur- Distribution. Fleas of this species were the mise that fleas of this species are sensitive to ones most commonly encountered at the test the high temperatures during the hot summer Ill BnicHAM Younc University Scienci Hi i i i i in

GRAPHIC SCAL£, FT N 950,000 LEGEND Ar»o Boundary. Paved Rood Dirt Rood

Tonopoh

1 50 miles from Mercury nzoo.ooo-^^^

N 650,000

Fig, 6 Geographic distribution i>l Meringis dipodomysQ and Meringis parkeri Jf Fleas of the Nevada Test Site I linn .11 \ m l"i\i. NivEBsm Science Hi i i i i is

Fig. 8. Geographic distribution ol Meringia liuhlumlig^ JordanopsijUa ullndij^ and Stenistomera o/pi Fleas of the Nevada Test Site 13 the summer of 1965 without obtaining a single 281) stated: "The two species (of Callistop- flea of this species. Collections of nests were syllus ) come consistently off deer mice, occasion- made in the same locality where fleas of this ally off other rodents." This has been our ex- species previously had been collected. The studv perience in many years of collecting in Utah. of Howell (1955) revealed a very low popula- The host, /'. crinitis, has been collected in most tion of all species of fleas in the nests of N. months of the year and at widely-separated lepida in Utah during the summer periods. parts of the test site, mainly along the foothills The specimens from which the original descrip- in mixed communities; yet this is the only flea tions were made were collected in December of this species we have taken (Fig. 9). and November. Megarthroglossus procus Stenistomera alpina (Baker), 1895 Jordan and Rothschild, 1915

Distribution. Fleas of this species have been Insofar as we can determine, our single spec- considered by some workers as rare in occur- imen belongs to the subspecies M. procus procus. rence. In our studies at the Nevada Test Site, The male was collected from a Desert Wood they occurred commonly on the Desert Wood Rat in the vicinity of Tippipah Spring in a Rat at some seasons of the year. This host is Coleogyne biotic community in November (Fig. not restricted to any one biotic community at 9). the test site, but is found in the Pinyon-Juniper, Coleogyne, Salsola, Larrea-Franseria, and Gray- Anomiopsyllus amphibolus ia-Lvcium communities. About the only restric- Wagner. 1936 tive influence affecting its distribution is tin' Distribution. This species was taken at wide- absence of house-building materials, rocky led- ly-separated points at the test site, but most ges, or large shrubs in which a house may be frequently at the northwestern part along the constructed with appropriate situations for nest- foothills or at higher elevations. The biotic ing either in the house or in close association community association was mainly with Pinyon- with it. Most of the fleas were collected from Juniper, Grayia-Lycium, and in mixed vegetative hosts trapped in the mixed biotic communities types ( Fig. 9). (Fig. 8). Host Associations. collection was Host Associations. The Desert Wood Hat One each made from a pocket mouse. Deer Mouse, Pinyon is the principal host. Six fleas were taken from Mouse, and Cactus Mouse. Five separate en- a Canyon Mouse (Peromyscus crinitis), and counters were with the Desert Wood Rat. three from a White-tailed Antelope Squirrel. Seasonal Occurrence. collections were Seasonal Occurrence. Extensive collections Two made in March, one in April, six October, of the Desert Wood Rat were not made con- in and one in December. tinually throughout any one year. A collecting schedule set up to include the months of Oc- Comments. In Utah studies, hundreds of tober, November, December, and January re- specimens ot A. amphibolus were found in the sulted in thirty collections in December com- nests of N. lepida, whereas few were taken pared to only four encounters each for January, from the host's body (Beck et ah, 1953; Howell, October, and November. During the summer LS55). During June, July, August, and Septem- months of 1965 (June, July, August), collection ber collections in Utah, nests were relatively of N. lepida and its nests was emphasized. Sev- free of fleas of this species compared to other en fleas of this species were taken from one months of the year. Nests were not examined nest in June. None was found in 58 other during fall, winter, and spring months at the nests collected during the summer. test site. The collection of this flea from the host's Comments. Although the principal host, the body indicates a similarity of occurrence at the Desert Wood Rat, is widely distributed, there test site and in collections made in Utah. Addi- seems to be some relationship between the pres- tional support to this view are our studies dur- ence of fleas of this species and the seasonal ing the summer of 1965. More than fifty nests occurrence of the rat. of N. lepida were collected, but not a single flea of any species was found. We did take Callistopsyllus deuterus Jordan, 1937 seven specimens of Stenistomera alpina from The only specimen of this species taken is a one host, but A. amphibolus was not encoun- male from a Canyon Mouse. Hubbard (1947: tered. I Si BniCHAM V.i si. SIMI1SII-, || s. f |,

GotdFlot

N 950,000 . S

N 650,000

Fig. '>. Geographic distribution ol Callistopsytttu deuterus^^ inomiopsylhis nmphibolusjf Vfegarthroglossus /inninjfc .mil Rhadinopsylla heiseri Fleas of the Nevada Test Site 15

Rhadinopsylla heiseri (McCoy), 1911 Rhadinopsylla sectilis Jordan and Rothschild, 192.) This species has been listed as Actenopthal- mus heiseri in some publications. We follow the Rhadinopsylla sectilis was listed by Hubbard taxonomic placement by Hopkins and Roths- (1947) and Holland (1949) as Micropsylla sec-

is tilis. child ( 1962) in which Acte'nopthalmus listed Hopkins and Rothschild (1962) listed as a subgenus and the generic status is Rhadi- Micropsylla as a subgenus of Rhadinospsylla. As nopsylla. According to personal cor- far as we can determine, Jellison ( our specimens are of

respondence ), there is a close relationship be- the subspecies R. sectilis sectilis. tween heiseri and multidenticulata of Morlan Distribution. Most hosts were taken in the and Prince (1954). Nevertheless, we feel these Grayia-Lycium community. A few encounters specimens more closely fit the description for were from Larrea-Franseria and Coleogyne. The heiseri. geographical distribution was generally in Yuc- ca and Frenchman Flats Fig. Distribution. Most of the 127 fleas collected ( 10). of this species were taken from hosts found in Host Associations. The Chisel-toothed Kan- mixed vegetation communities in Jackass Flats garoo Rat is the host on which fleas of this at the southwestern part of the test site. The species were most commonly encountered. The next most often encountered collections were in next most common host is the White-tailed An- Larrea-Franseria communities in the southeast- telope Squirrel. One specimen was removed ern section. Another community with about from O. torridus, and three were from Merriam's equal encounters is the Coleogyne community Kangaroo Rats. in the northeastern part. All other communities except the Pinyon-Juniper produced occasional Seasonal Occurrence. About equal numbers collections (Fig. 9). were collected during January, February, March, and December. One specimen each was taken Host Associations. The host on which most in October and November. fleas of this species were taken is the White- tailed Antelope Squirrel. Occasional collections Comments. Apparently fleas of this species are were also made from the Desert Kangaroo Rat, not abundant <>n a variety of hosts. Collec- tions ot /'. Chisel-toothed Kangaroo Rat, Southern Crass- such hosts as maniculatUS at higher hopper Mouse, and Southern Pocket Gopher elevations may produce a greater number of specimens. (Thomomys umbrinus). Onychomys torridus is a predator and thus may be expected to have fleas from various rodent victims. Carteretta carteri Fox, 1927

Dr. William L. Jellison (personal corres- Seasonal Occurrence. Most collections were is pondence ) of the opinion that these fleas are in January, with December, November, and of the subspecies C. carteri clavata. February ranking in relative sequence for other collections. Only one collection was made in Distribution. The geographical and ecolog- March, one in April, and one in October. Fleas ical distribution is throughout Frenchman and were not found in other months of the year. Yucca Flats. Most encounters were in the Lar- rea-Franseria community, followed by Coleogyne Comments. The original description of this and mixed vegetative communities (Fig. 10). species made by McCoy (1911) was of a female taken at Mojave, California. The host is un- Host Associations. Three collections (total of known for that collection, but in later collections seven fleas) were taken from the Long-tailed by several workers, the most common host en- Pocket Mouse (Perognathus formosus). One countered was the White-tailed Antelope Squir- specimen each was from a Merriam's Kangaroo rel. Hubbard contributed a male and female of Rat, a Canyon Mouse, and a White-tailed Ante- this species which he had collected in Decem- lope Squirrel. ber of 1949 at Carson City, Nevada, to the Seasonal Occurrence. Specimens were taken Rritish Museum in 1950. These were from A. in March, April, October and November. leucurus. The great majority of our collections were likewise from this same host. No doubt Comments. The occasional appearance of the instances of occurrence on hosts other than this flea on Merriam's Kangaroo Rat and the A. leucurus are accidental, for several species White-tailed Antelope Squirrel may be accident- of animals live in close association with this al, for only a few of several hundred of these squirrel. hosts that were examined were infested with Si III BnlCHAM YOUNC L'NIVEBSITl ii \. I I I I MS

GRAPHIC SCALE, FT

LEGEND Area Boundary Pov«j Rood Dirt Rood

Fig 10 I ctpliii distribution "I Rhadinopsijlla sectllisQ find Carteretta carteri^ Fleas of the Nevada Test Site 17

Gold Flat N 950,000 l

Fig. 11. Geographic distribution of Thrassis hacchi liim.iiwi YOL'NC I \nnwn Si n \i i Bmiiiis

fleas ul tins spei ies, < >l tin- man) mice examined Antelope Squirrels wen- collected during the during tin' five-veai period, onl) six specimens simimer hut the) did not possess lleas.

a/in clavata wire taken. It is possible Comments Fleas oJ this species reflect the ih.il Hi. is

Thrassis hacchi i Rothschild I, 1905 where the incidence ol occurrence is greatl) reduced during the summer months. Burrow \ total nl 1,491 specimens of tins species examination and nesting site collections would uas collected during the period <•! lliis study. be a worthwhile endeavor, lor thev might help making it the must abundant flea .it the test determine what happens to the adult Ilea popu- sit.- I sing Stark s L958 publication on i tah lations during the summer months. Siphonaptera .is .1 guidi we conclude that the subspecies is /' hacchi gladiolus

1 Thrassis

1 inhabitants. were found in all commu- ' The) I j~>s . our specimens are T. aridis hoffmani.

'/'. nities relegated (<> these elevations ( Fig. 11 ). This species ranks next to bacchi gladiolus in abundance at the (est site. Most Associations. I In- White-tailed Ante- lope Squirrel was most often encountered as the Distribution. There was widespread distri- dost. The Southern Grasshopper Mouse was bution at the test site in the biotic communities the next most prominent host, with the Chisel- of the valleys and foothills. Most of the Heas toothed Kangaroo Ral about equal in host pref- came from the Grayia-Lycium, Coleogyne, Lar- erence. Other hosts on which specimens were communities Fig. rea-Franseria, and Salsola j 13). infrequentl) taken are Merriam's Kangaroo Rat, Host Associations. Thrassis aridis hoffmani the Desert Kangaroo Rat, Little Pocket Mouse. was most encountered on the ('hisel- Canyon Mouse. Pinyon Mouse, and Coyote. often toothed Kangaroo Hat. Merriam's Kangaroo Hat Seasonal Occurrence. There were about was tin next most frequently infested, with equal numbers of collections made during most the White-tailed Antelope Squirrel producing ol the months ol the \ear except the summer occasional specimens. One lo live encounters

ol 1 il\ (Fig. There months 1 and AugUSl 12). were made with the Southern Grasshopper was an absence ol fleas in July, and only one Mouse. Kit Fox, Canyon Mouse. Southern Pocket specimen was taken in August. White-tailed Gopher, Long-tailed Pocket Mouse, and Hock Squirrel.

Seasonal Occurrence. The greatest numbers were collected in November, followed by Oc- tober. December, and January. Incidental col- lections made at other times ol the year were one each in June and August (Fig. 14).

Comments. The same picture for seasonal distribution as seen in others is also reflected m this species. Summer incidence is low or lacking, with the highest being late fall to mid- w inter,

1 I'M*) DactulojMijlla blut I Fox 1,

Foxella and Dactylopsylla are variously used bv workers in generic designation lor pocket gopher lleas There is sufficienl difference in genital anatom) alone to separate them as two Jan F.b Ma Aug S«p. Oct Nov D«<. distinct genera, and we follow Prince (1945) hacchi. Fig. I- Seasonal occurrence of Thrassis The and Holland I' 19 m lliis arrangement. Speci- 111411ns on do not represent the total incidence mens o| /). IiIiki wen submitted to Dr. W, I.. numbers ol specimens taken during .1 month, hill |< IlisiMi who recommended the subspecific des- ' 11,, li, .a. ili.- total numbci "l 1 ollei ignation ol n bluri ftsilm 1, „!, . hi 1, 11, Fleas were found. Fleas <>e the Nevada Test She 19

Kawich Valley Gold Fiat GRAPHIC SCALE, FT. N 950,000 I §

N 900,000

N 800,000

Tonopah

1 50 miles from Mercury

N 700,000-^

N 650,000

Fig. 13. Geographic distribution of Thrassis aridis £ FoxeUa ignota aft Dactylopsylla hluei^^ and Diamanus mdntanus -^. Hlili.ll \M Yoi'NC I'NIVKIlsm Si II \i I Hi I I I I IN

sixteen fleas. Vn unusual record was thi ex- traction "I 121 lie. is from a Southern I'ocket Gophei which was collected from a Pinyon

|uniper communit) I Fig

Collections were made in |anuar\ and Jul\.

/ oxella ignota should he further studied, es- peciall) in the Pinyon-Juniper communit) and tin Vrtemisia associations found at higher ele- vations.

Diamanus montanus > Bakei . 1895

In our surveys in Utah we found the Rock Squirrel to lie a common host lor this flea. Stark (1958) likewise reports this host prefer- ence. Literature records lor other hosts indi- F«b Mar Apf May Ju Jul Aug. Sop Oct cate their habitats as foothills and median mon- tane- elevations. At the test sit. s variegatus

1 I Seasonal occurrence "I Thrassis aridis. The Fie. was collected in the Pirn on- Juniper communit) figures on incidence not represent total num-

on Rainier Mis,, i Fig. 13 . In our studies over bers ot specimens taken during tl»- month, but usual cast- was indicate the total number of collections (encoun- the years, especiall) in Utah, the wherein fleas were found. to find the host heavil) infested, but at the test ters I site onl) twelve fleas were collected. These

Distribution. Most of the specimens were were taken during April. May, and June. From gophers trapped in Grayia-Lycium com- Obviously a more extensive surve) is needed. munities. A few were from Coleogyne and some especially at higher elevations, to determine the- from mixed Mutational types. One (lea was extent ol distribution and seasonal occurrence collected from an area ol much gopher activity Of this Ilea. in the west-central area of the test site which is The Ilea is ot public health importance in Artemisia association at a higher elevation in an that it has been considered moderately effective

i. I ig 13 transmission Stark. m plague I L958

Host Associations. The Southern Pocket

l H>~> Gopher was the principal host. Collections were Orchopeas sexdentatus (Rothschild), l also made trom the Southern Grasshopper We consider this Ilea to he ol the subspecies Mouse and liom an unknown species of pocket 0. sexdentatui agilis. The taxonomic characters mouse. which have been used to distinguish the sub- Seasonal Occurrence. Collections were made species of 0. sexdentatus are variously inter- during fanuary, February, March. May. June. preted by different workers. A rather careful |uh. and September. Most of them were en- Stud) needs to be made- ol this species over a the basis ol lit- countered in March. wide geographical range. On erature descriptions and accompanying illustra- Comments. The principal host lor this flea tions, our specimens .uc nearer agilii than a is T. umbrinus, although it is occasional!) found closely related form, nevadensis. Hubbard in the same vicinity. on other mammals living i 1957) stated; "During 1938 this flea, }i

is i )~> include Pinyon-Juniper, Salsola, Grayia-Lycium, Foxella ignota | baker |,

and several mixed vegetative types i Fig L5). ()nl\ two hosts were' found infested with ol separate col- fleas oi tins species \ Deseit Cottontail col- Host Associations. Four 85 lections were from the Canyon Mouse. The re- lected in a 1 .atrea I'Yanseii.i community vicldcd Fleas of the Nevada Test Site 2!

N 650,000

Fig. 15. Geographic distribution of Orchopeas sexdentatus 6

I 1 |ISI Sc I I VOUNC M\ II II S( I. hi I I I IN mainder were taken from the bodies ol Desert Seasonal Occurrence. Specimens were col- Wood Rats. lected about equalh during \pril and Novem- ber. Two encounters wen m |uly, and on< ea< h Seasonal Occurrence. Twenty-three collec- in VugUSl and < >< toller .2* tions were made in [anuary, » in November

I l I Vi < >< m embei , 9 in tober and 9 in Man h. Comments. Extensive year-round surveys During February, Vpril, fune, and Septembei need to be in. nil m tin Pinyon-Juniper and one to three em ountei were made in eat li "I \rtcmisia associations in the western and north the months. Western parts ol the test site.

Comments. Fleas ol O. iexdentatus (no Monopsyllus eumolpi (Rothschild), L905 subspecific designation | have been found to plague in nature are considered ef- harbor and Distribution. All specimens were collected ficient vectors ol the disease (Eske)

Mm mil; the L965 s ner surveys of wood Host Associations. This Ilea is commonly rat habitats .it the test site, we were impressed termed the "chipmunk" Ilea. Most collections l>v the similarity "l conditions encountered in at the test site were from the C.'litt Chipmunk surveys iii Othei parts ol the western United (Eutamias clorsalis). Additional fleas were tak- States. We found situations in which whole en from the Deer Mouse and Great Basin Pocket colonies ol wood rats apparently had died. Mouse. \ests and houses showed varying degrees ol n .. lit us.' and dead hudles weir found ill till' Seasonal Occurrence. Most collections were ". \s stated . . the in |ulv and houses. b) Beck I L955), in April with minor occurrences disappearance ol once abundant rodent popu- \d\ ember. lations has been too consistent to he happen- ( lomments. Eighty-three specimens w ere col- stance. These extreme fluctuations in rodent lected. Although this ma) be considered com- populations may he due in part to man's inter- paratively lew. all collections were in the I'm ference with environmental conditions, or per- yon-Juniper comn ity, a biotic situation where haps it is a reflection "I inherent population we have done relativel) little surve) work with rhythm. ... It is possible that the almost, and. the vertebrate fauna m sonic cases, complete disappearance is due to a disease agent and the vectors involved." Malaraeus telchinum (Rothschild), 1905

Only two specimens of this species were col- Monopsy litis wagneri (Baker), 1904 lected at the test site. One was taken from the

Our specimens arc of the subspecies M. ir

, Fig. 16). hosts located along the foothills or at higher elevations. The majority were from the Pinyon- Malaraeus sinomus (Jordan), L925

[uniper biotic community i Fig. L6). One ol the principal characteristics used tu Host Associations. In our extensive surveys separate M. sinomus from M eremicus is the in Utah and in sin\c\s In other workers, speci- comparative length of the first metatarsal seg- mens ol M. wagneri wagneri are considered to ment. In eremicus it is longer than the com lie the most lrei|ueutK collected fleas. They bined 2nd. 3rd. and 1th tarsal segments, where have been collected at elevations much higher as in sinomus it is shorter. The majority of our than those "I tin desert valleys ami lowlands specimens lit sinomus, but there are borderline at the test site. Most published data list the cases. Nevertheless, the structure ol the ninth Deei Mouse as the preferred host. \t the test Stemite of the male is unniistakahlv that of M. site /'. maniculatus lias been found mainl) at Sinomus It ma) be that specimens at the test highei elevations in the 1'invon Juniper com- sit. arc at the meeting place ol eremicus popu- munity, and has provided some specimens of lations from the southeast and the sinomus pop- \/ u agneri u >agm ri. additional collet ii«>ns wen ulations from the north ami southwest, made from the Pinyon Mouse, Southern Grass- hoppei Mouse and Chisel-toothed Kangaroo Distribution. Must collections at the test site Rat. were made alone the foothills or in basins where Fleas of the Nevada Test Site 23

Kawich Valley

GRAPHIC SCALE, FT.

LEGEND Area Boundary Paved Road Dirt Road

Fig. 16. Geographic distribution of Monopsyllus wagneri^ff Malaraeus telchinum'f^ Monopsyllus eumolpi^^ and Malaraeus sinomus Q i it I Illiir.MAM ^ ni. Nivciisrn SriKNin Hi i i i i is

(In elevation is little highei tlian Fren< hman and Fleas ol tins species likel) are to l» found on Yucca Flats. Vboul e(|tial encounters were madi rabbits at higher elevations and northward into in the Larrea-Franseria and Coleogyne com- the (.real Basin region, (.'ollections ol rabbits

munities, | tins occasional collections from Gra) al higher elevations at the test site need to be

ii i.i Lycitim, Pinyon |uniper and mixed vegetative in. li in eorroborati tins point ol \ iew . So far,

areas Fig, Ifi inosi collections ol rabbits have been made only

III the vallcN s at the lesl site Host Vssociations. Members ol the genus Peromyscus are considered to he the preferred Peromt/scopsylla hosts. Most n| our collections were from the hesperomys Canyon Mouse, Deer Mouse, and Desert Wood Baku .. 1901 Rat with occasional collections from the White- Vccording to ke\s and descriptions given 1>\ Squirrel, tailed Vntelope Southern Grasshopper |ohnson and Traiib 1954 specimens be- Mousi and Pin) i >n Mouse. long to the subspecies P hesperomys adelpha.

Sc.iMni.il ( ollections Occurrence. were made Distribution. Most hosts and their fleas were "I the and even month year except August taken along the foothills or at highei elevations. September, with most encounters in November, The biotic community most commonly repre- December, [anuary, and February, successively. sented was Coleogyne, followed by the mixed

( >i 1 1 \ nr wen one two encounters made during vegetative types, and to a lesser extent the other months ni the yeai Larrea-Franseria and Pinyon-Juniper communi-

Comments. Tins flea is associated with hosts ties i Fig. 17). living along the loiitliills. Most Associations. The Southern Grasshop- per Mouse and C.muiii Mouse were equally en-

< I, Malaraem uphorbi i Rothschild 1905 countered with flea consortes. The next most often encountered was the Deer Mouse. Single Only one collection ol fleas ol this species encounters were made with the Pinyon Mouse. was made. Two males and si\ females were re- Great Basin Pocket Mouse. White-tailed Ante- moved from a Deer Mouse in November from lope Squirrel, and Southern Pocket Gopher. a Pinyon-Junipei communit) on Rainier Mesa Fig. 17). Seasonal Occurrence. January, November, December, and April were the mouths o| the Amphipsylla neotomae Fox, 1940 year when most encounters were made. Single encounters were made in February, Max. fury, \ male and female specimen were taken September, and October. from a Deserl Wood Rat in the eastern end ol |ackass Flats in a Larrea-Franseria community Comments. Records ol this species empha-

I ig 17). size the need for more intensive surveys along Frank VI. Prince (personal correspondence) the foothills and at higher elevations ol the

indicated thai \ neotomae should be listed as test site.

1 '""' neotomae. has been pub- to I Malaraem Nothing According Holdenried and Morlan I lished to this elteet to date, so we are obligated fleas ol this species have been found naturally to leave neotomat in its present generic alloca infected with plague organisms. tion — \mphipsylla. Nycterhlopsylla t>ancouoerensk

Odontopsyllm dentatus i Baker 1904 Wagner, 1936

Three fleas ol tins species were in 1949) collected Both L947) and Holland | Hubbard I \|in| from Nuttall's Cottontail Rabbit from a listed this ilea under the generic name. Eptes- Pinyon-Juniper communit) on Rainier Mesa COpsylUl. Following Hopkins and Rothschild 17). 1953), we arc using the generic name. Nycter- Fleas ol tins species, although not found in idopsylla. great numbers on the preferred hosts of several Five specimens were collected Irom a West- species ol / .i fins and Si/h ilagus and ern shot while | jackrabbits Pipistrelle Bat flying in the vicin- cottontails, respectivel) I, are considered com- i(\ ol Tippipah Spring in November. This area mon. Man) specimens ol the Black-tailed [ack- is characterized as a mixed vegetative type "I rabbit wen- collected during the period ol tins 17 biotic communit) ( Fig, ). study, and large numbers ol cottontails were The flea fauna ol hats at the test site is also taken and checked for ectoparasites, OnK relatively unknown, for few hats have been one collection ol ( >. tlcutiitus was encountered. collected. Fleas of the Nevada Te.si Site 25

Gold Flat GRAPHIC SCALE, FT. N 950,000 L

N 900.000

N 800,000

Tonopah

1 SO miles from Mercury N 700,000--^

N 650,000

Fig. 17. Geographic distribution of Malaraeus euphorbi^L Odontopsyllus dcntatus Nt/cteridopsylla (Epte copsylla) vancouvercnsisJk Amphipsylla neotomae J^. and Peromyscopst/Ila hesperomys A 2<> Riiii.iivM Young University Science Bulletin

CONCLUSIONS

Naturally the flea fauna is best known from applies to the foothill environs. The test site does those host animals which have been most fre- not contain high mountain ranges, and one would quently collected. Kangaroo rats and \\ bite not expect to find hosts and their ectoparasites tailed Antelope Squirrels were the animals most at the test site characteristic of high mountain often collected. In those anas oi the test site elevations such as the Sheep Range to the east where studies were made over several years and the Charleston Mountains to the southeast. a conspicuous reduction in numbers ol fleas About One-fourth ol the species of fleas re-

d ig the summer months was observed. Geo- ported from the test site ale 1 1 on i the Desert graphically, the fauna ol Frenchman, Yucca, Wood Hat. Nevertheless, tor the most part these

and [ackaSS Mats is liest known, and animals records are not a result ol vear-roimd collecting ol the foothills and mesas are least known. in the various blOtic communities where the rat

Uthough thirty-three species are listed in is Found. Such a studs would assist in resolving this paper, this does not represent all that arc some of the problems in the ol fleas expected to occur at the test site. The extensive ol the genus McHaraeus and related groups. stands ol Artemisia tridentata in the western There is a junction of the Great Basin and the hall ol the test site arc relatively unsurveyed Mojave biota at the test site, and the fleas biologically, and comparatively little has been characteristic of these two provinces may dem-

done with the Pinyon-Juniper community on the onstrate an unusual distribution pattern once it

mesas and elsewhere. To a certain extent this is known.

LITERATURE CITED

Allied, D. M., D E. Beck and C. D. Jorgensen. 1963b. Hayward C. I.. M. L. Killpack and G. I.. Richards, Biotic communities of the Nevada Test Site. Brig- L963. Birds of die Nevada Test Site. Hre4ha.11 ham Young Univ. Sei. Bull., Biol. Ser„ 2(2): 1-5.1. Young Univ. Sci. Bull., Biol. Ser.. 3(11:1-27. Allied, D. M., D E. Beck and C. D. Jorgensen. 1963b. Holdenried, R. and II. B. Morlan. 1955. Plague- Nevada list Site stuck aieas and specimen depos- infected fleas from northern New Mexico wild itories. Brigham Young Univ. Sci. Bull., Biol. Ser., rodents. Jour. Infect. Dis., 96:1.33-137. 2| t):l-15. Holland. G. P. 1949. The siphonaptera of Canada.

Beck I) E. 1955. Distributional studies of parasitic Science Service. Division of Entomology, Livestock in Utah, determined as actual and po- Laboratory, Pub. SIT. Tech. Bull.. 70:1-306.

tential vectors ot Rocky Mountain spotted fevei Hopkins, G. II. E. and M. Rothschild. 19.53. An ill- and plague, with notes on vector-host relationships. ustrated catalogue of tin- Rothschild collection of

Brigham Young Univ. Sci. Bull., Biol. Ser., 1(1): ( ) fleas Siphonaptera in the British Museum ( Na-

I i.l tural History). Univ. Press, Cambridge, Vol I. Beck D E., A. H. Barnum and L. Moore. 1953. Hopkins. G. H. E. and M. Rothschild. 1956. An illus- consortes found in the nests of Neotoma trated catalogue ol the Rothschild collection of fleas

iicrcnu ( limn a Ord ) and Neotoma lepida lepida (Siphonaptera) in the Britisb Museum (Natural Proc. Ut;ih Sci., Arts, (Thomas). Acad. and Letters History). Univ. Press, Cambridge, Vol. II. 30:43-52. Hopkins C. H. E. and M, Rothschild. 1962. An Beck, I) E., 1) M. Allied and E. I'. Brinton. 1963. illustrated catalogue ol the Rothschild collection of Ticks ol til,- Nevada Test Site. Brigham Vomit; ' fleas (Siphonaptera) in the British Museum Na Univ. Sci. Bull., Biol. Ser., 4(1):1-11. tural History). Univ. Press, Cambridge, Vol. III.

Eskej C H aui\ V. 11. Haas 1939. Plague in the Howell, F 1955. A stuiK ol the- aspcctional var- I western part of tin- United States. Infection in iations of Siphonaptera associated with the nests rodents, experimental transmission of fleas and of the Thomas Wood Rat. Neotoma lepida Upida inoculation tests for Health infection. Public Re- Thomas. The Great Basin Nat.. 15: ( 1-4 ) :35-49. ports, 54:1 167-1481. Hubbard. C. A. 1947. Fleas of western North Amer- Eskey, C. R. ami V. H. Haas. 19KI Plague in the ica. Iowa State College Picss. Ames. western part ot the United Stales Public Health Johnson, P. T. and R. Tiaub. 19.54. Revision of the Bull.. 2,54:1-83. Ilea genus PewmysCOpsylla. Smithsonian Misc. Col-

Goates, M. A. 1963. Mites on kangaroo rats at the lections. 123(4 1 1-68. Nevada Test Site. Brigham Young Univ. Sci. Bull.. ' [orgensen, C. D. and C. L. Hayward. 1965. Mam- Ser., Biol 3(41:1-11. mals ol tin- Nevada lest Site. Brigham Young Sci. Bull., Biol. Ser., 8(3):1-81. Hall. E. It. (Ed 1957 Vernacular nanus „f North Univ. American mammals Inn ol Kansas Museum ol McCo) G. W. 1911 A new flea Ctenophthalmus Natural History. Miscellaneous Publication No. 14: heiseri spec. QOV. (Siphonaptera). Entomol. Neves. 1-16. 22:445-448. Fleas of the Nevada Test Site 27

Morlan, H. B. and F. M. Prince. 1954. Notes on the importance. U. S. Dep. Health, Educ. and Welfare,

subfamily Rhadinopsyllinae Wagner, 1930 ( Siph- Public Health Service, Commun. Dis. Center, At-

onaptera, ) . and description of a lanta, Georgia. new species. Rhadinopsylla multidenticulatus. Tex- Tanner, W. W. and C. D. Jorgensen. 1963. Reptiles as Reports on Biol, and Med.. 12:1037-1046. of the Nevada Test Site. Brigham Young Univ. ' Prince, F. M. 1945. Descriptions of three new species Sci, Bull.. Biol. Ser., 3(3):1-31. of Daetijlopsylhi Jordan and one new subspecies Traub, R. and V. J. Tipton. 1951. Jordanopsylhi allredi of Foxella Wagner, with records of other species a new genus and species of flea from Utah ( Siph-

in the genera ( Siphonaptera ). Canadian Ento- onaptera). J. Washington Acad. Sci., 41(8) :264-270. 15-20. mol., 77(1): Wheeler, C. M. and R. 1945. J. Douglas. Sylvatic Stark, H. E. 1958. The Siphonaptera of Utah: Their plague studies. V. The determination of vector taxonomy, distribution, host relations and medical efficiency. Infect. Dis.. 77:1-12. J. n