HERRIN, Charles Selby, 1939
This dissertation has been microfilmed exactly as received 6 9 -2 2 ,1 4 4
HERRIN, Charles Selby, 1939- A SYSTEMATIC REVISION OF THE GENUS HIRSTIONYSSUS (ACARI - MESOSTIGM AT A) OF THE NEARCTIC REGION, BASED ON THE PRINCIPLES OF NUMERICAL TAXONOMY. The Ohio State University, Ph.D., 1969 Entomology
University Microfilms, Inc., Ann Arbor, Michigan A SYSTEMATIC REVISION OF THE GENUS HIRSTIOUYSSOS
(ACARI - ?-SS03TIGHATA) OF THE NEARCTIC REGION,
BASED ON THE PRINCIPLES OF NUMERICAL TAXONOMY
DISSERTATION
Presented in Partial Fulfillinent of tho Requirements for the Degree Doctor of Philosophy in the Graduate School of The Ohio Stato University
b y
Charles Selby Herrin, 3.S., M.S,
******
Tho Ohio Stato University 1969
Approved by
Academic Faculty of Entom ology ACKNOWLEDGMENTS
Grateful acknowledgment is made to E. I V. Baker and the D . S .
National Kuseum, G. 0 , Evans and the British I-Iuseum of Natural History,
D. M, Allred, R, W. Strandtmann, E, W. Jameson, Jr., F. J , R adovsky,
C. D. Radford, and G. W, Krantz for the loan of study specimens. Tho
Acarology Laboratory provided additional study specimens, laboratory space, equipment, and supplies. Special acknowledgment is made to
Dr. D. E. Johnston, ray adviser, and Dr. G. W. 'Tharton, Director of tho
Acarology Laboratory, for their assistance, valuable suggestions, and criticism of the manuscript.
Acknowledgment is made to Dr. F. J, Rohlf and the University of
Kansas Computation Center for processing the data using their Numerical
Taxonomy System, NTSYS. The Ohio Stato University Computer Center pro
vided froe computer time for the preliminary statistical computations.
This investigation was supported by a predoctoral traineoship
from the Rational Institutes of Health Training Grant 2TX-AI-00216
to tho Acarology Laboratory.
ii VITA
Kay 16, 1939...... Born - Kechanicsburg, Yazoo Co., Mississippi
I 96 I-I 963 ...... , Research Assistant, Department of Zoology and Entomology, Brigham Young University, Provo, Utah
1 9 6 3...... B.S., Brigham Ycrung University, Provo, Utah
1 9 6 3 -1 9 6 4...... Teaching Assistant, Department of Zoology and Entomology, Brigham Young U niversity, Provo, Utah
196^-1965 ...... Research A ssistant, Department of Zoology and Entomology, Brigham Young University, Provo, Utah
1965-196 6 ...... Teaching Assistant, Department of Zoology and Entomology, Brigham Young University, Provo, Utah
1966 ..,..,, ...... M.S., Brigham Young University, Provo, Utah
1966 -I 969 . , * ...... Acarology Predoctoral Trainee, Acarology Laboratory, The Ohio Stato University, Columbus, Ohio
PUBLICATIONS
Herrin, C. Selby. 1964. Notes on the Distribution of Otobius lagophilus. Proc. Utah Acad. Sci., Arts, Letters 42(1) :l65, November, 1964.
Herrin, C. Selby, and D Elden Beck. 1965. Observations on the Biology, Anatomy, and Morphology of Otobius lagophilus Cooley and Kohls. Brigham Young Univ. Sci. Bull., Biol. Ser. 6(2 ) t 1-19, 1 table, figs. 1-32, February, 1965#
iii Herrin, C, Selby. 1963. Dsrmacentor andersoni in National Forest Recreational Sites of Utah. Great Basin Natur. 28(l):30-43» 2 tables, figs, 1-3? March, 1968 B
FIELDS OF STUDY
Major Field: Entomology
Studios in Acarine Systematics. Professors D. E, Johnston and G, W. Wharton
iv TABLE OF CONTENTS
Pago ACKNOWLEDGMENTS ...... i i
VITA ...... i l l
LIST OF TABLES ...... v i i
LIST OF ILLUSTRATIONS ...... v i i i
INTRODUCTION ...... 1
Application of Numerical Taxonomy to tho Acari
The Genus Hirstionyssus
MATERIALS AND METHODS...... 9
M aterials Utilized
Characters Utilized
Computational and A nalytical Methods
RESULTS OF NUMERICAL TAXONOMIC ANALYSES ...... 16
Sim ilarity Coefficients
Cluster Analysis
Graphic Analysis
DISCUSSION...... 39
Establishment of a Classification and Correlation of the Numerical Taxonomic Analyses with Biological Data
Groups and Taxospocies Formally Recognized
v Page SYSTEMATICS OF TH3 GENUS HIRSTI0NY55US ...... 75
Subfamily Hirstionyssinae Evans and T ill, 19 66
Genus Hirstionyssus Fonseca, 19*+S
Key to Females of the Genus Hirstionyssus
Recognized Taxospecies of the Genus Hirstionyssus
Hosts of Hearctic Hirstionyssus Taxospecies
SUMMARY AND CONCLUSIONS ...... 181
APPENDIX
I ...... 185
II ...... 193 III ...... 200
REFERENCES CITED ...... 297
vi LIST OF TABLES
Table Page
1. Partitioning of characters ...... 12
2. Cophenetic correlation coefficients between sim ilarity matrices and phenograms obtained from the three trans formed data matrices ...... 18
3« Phenogram groupings obtained from six sim ilarity m a tric e s ...... 19
h-. F k tr ix o f c h a r a c te r s t a t e v a lu e s o f 30 qualitative characters for the 25 recognized taxospecies ...... M
vii 1
LIST OF ILLUSTRATIONS
Figure Page
1. Phenogram obtained from the Condensed Ratio Correlation (CRC) matrix ...... 233
2. Phenogram obtained from the Condensed Basic Correlation (CBC) m atrix ...... 235
3. Phenogram obtained from the Condensed Ratio D ista n ce (CRD) m atrix ...... 237
4. Graph diagram of the STAF cluster; relationships based on a revised Condensed Ratio Correlation (revised-CRC) matrix ...... 239
5. Graph diagram of major groups "G" and "H"; relationships based on a revised Condensed Ratio Correlation (revised- CRC ) m a trix ...... 239
6 . Graph diagram of major groups "I” and "S-K"; relation ships based on a revised Condensed Ratio Correlation (re v ise d -C R C ) m a t r i x ...... 241
7. Semi-diagramatic representation of the dorsal chaetotaxy in tho females of Hirstionyssus ...... 243
8 . Hirstionyssus isabellinus (Oudemans). Paraxial view of chelicera in the female ...... 245
9. Hirstionyssus isabellinus (Oudemans). Ventral viow of gnathosona in the female ...... 245
10. Hirstionyssus isabellinus (Oudemans). Ventral view of tarsus II in the female ...... 245
11. Hirstionyssus isabellinus (Oudemans). Ventral view of tarsus IV in the female ...... 245
12. Hirstionyssus femuralis Allred. Ventral view of the female ...... 247
viii Figure Page
13• Hirstionyssus femurails Allred. Dorsal view of the female ...... 2*4-7
1*4-. Hirstionyssus femu rails Allred. Ventral view of tarsus II in the female ...... 2*4-7
15. Hirstionyssus femur alls Allred. Ventral view of genu II in the fem ale ...... 2*4-7
16 . Hirstionyssus geomydis (Keegan). Ventral view of tho female 2*4-9
17. Hirstionyssus geomydis (Keegan), Dorsal view of the female ...... 2*4-9
18. Hirstionyssus geomydis (Keegan). Ventral view of tarsus II in the female ...... 2*4-9
19. Hirstionyssus thomomys Allred and Back. Ventral view of the female ...... 251
20. Hirstionyssus thomomys Allred and Beck, Dorsal view of the female ...... 251
21. Hirstionyssus thomomys Allrod and Beck. Ventral view of tarsus II in the female ...... 251
22. Hirstionyssus longlcholae Allred and Beck. Ventral view of the fomale ...... 253
23. Hirstionyssus longicholao Allred and Beck. Dorsal view of the female ...... 253
2*4-. Hirstionyssus longichelae Allred and Eeck, Ventral view of tarsus II in tho female ...... 253
25. Hirstionyssus hi 111 (Jameson), Ventral view of the female ...... 255
26. Hirstionyssus hi111 (Jameson). Dorsal view of the female ...... 255
27* Hirstionyssus "C" n. sp. Ventral view of the fem ale ...... 257
28, Hirstionys sus "G" n, sp. Dorsal view of tho fe m ale ...... 257
ix Figure Page
29. Hirstionys sus incomptis (Eads and Hightower). Ventral view of the female ...... 259
30. Hirstionyssus incomptis (Eads and Hightower). Dorsal view of the female ...... 259
31. Hirstionyssus triaoanthus (Jameson). Ventral view of the female ...... 261
32. Hirstionyssus triacanthus (Jameson). Dorsal view of the female ...... 261
33* Hirstionyssus "B" n. sp. Ventral view of the fem ale ...... 263
34, Hirstionyssus "B" n. sp. Dorsal view of the fem ale ...... 263
35* Hirstionyssus "B" n. sp. Ventral view of tarsus II in the female ...... 263
36 . Hirstionyssus talpae Zemskaya. Ventral view of the female ...... 265
37. Hirstionyssus talpae Zemskaya. Dorsal view of the female ...... 265
39. Hirstionyssus talpae Zemskaya. Ventral view of tarsus II in the female ...... 265
39* Hirstionyssus affjnis (Jameson). Ventral view of the female ...... 267
40. Hirstionyssus affinis (Jameson). Dorsal view of tho female ...... 267
41. Hirstionyssus affinis (Jair.eson). Ventral view of tarsus II in the female ...... 267
42. Hirstionyssus affinis (Jameson). Ventral view of tarsus IV in the female ...... 267
43. Hirstionyssus "D" n, sp. Ventral view of the fem ale ...... 269
44. Hirstionyssus "I)" n. sp. Dorsal view of the fem ale ...... 269
x Figure Page
45. Hirstionyssus "D" n. sp. Ventral vierw of tarsus II in the female ...... 269
46. Hirstionyssus "D" n. sp. Ventral view of tarsus IV in the female ...... 269
47. Hirstionys sus occldentalis (Ewing). Ventral view of the female ...... • 271
48. Hirstionyssus occldentalis (Ewing). Dorsal view of the female ...... 271
49. Hirstionyssus occldentalis (Swing). Ventral view of tarsus II in the female ...... 271
50. Hlrstionyssus occldentalis (Swing). Ventral view of tarsus IV in the female ...... 271
51. Hlrstionyssus eutamlae Allred and Beck. Ventral view of the female ...... 273
52. Hirstionyssus eutamlae Allrad and Beck. Dorsal view of the female ...... 273
53. Hirstionyssus eutamlae Allred and Beck. Ventral view of tarsus II in the female ...... 273
54. Hirstionyssus eutamiae Allred and Beck. Ventral view of tarsus IV in the female ...... 273
55* Hirstionyssus c.ynomys (Radford). Ventral view of the female ...... 275
56 . Hirstionyssus cynomys (Radford). Dorsal view of the female ...... 275
57. Hirstionyssus cynomys (Radford). Ventral view of tarsus II in the female ...... 275
58. Hirstionyssus cynomys (Radford). Ventral view of tarsus IV in tho female ...... 275
59. Hirstionyssus utahensls Allred and Beck. Ventral view of the female ...... 277
60. Hirstionyssus utahensis Allred and Eeck. Dorsal view of the female ...... 277
xi Hirstionyssus utahensis Allred and Beck. Ventral view of tarsus II in the female ......
Hirstionyssus utahensis Allred and Beck. Ventral view of tarsus IV in the female ......
Hirstionyssus latiscutatus (de Keillon and Lavoipierre). Ventral view of the female ...... 279
Hirstionyssus latiscutatus (de Keillon and Lavoipierre). Dorsal view of the female ...... 279
Hirstionyssus latiscutatus (de Keillon and Lavoipierre). Ventral view of tarsus II in the female ...... 279
Hirstionyssus "A" n. sp. Ventral view of the fem ale ...... 281
Hirstionys sus "A" n. sp. Dorsal view of the fem ale ...... 281
Hirstionyssus torus Allred and Beck. Ventral view of the female ...... 283
Hirstionyssus torus Allred and Bock. Dorsal view of the female ...... 283
Hirstionys sus torus Allred and Beck, Ventral view of tarsus IV in the female ...... 283
Hirstionyssus obsoletus Jameson. Ventral view of the female ...... 285
Hirstionyssus obsoletus Jameson, Dorsal view of the female ...... 285
Hirstionyssus isabellinus (Oudemans). Ventral view of the female ...... 287
Hirstionyssus isabellinus (Oudemans), Dorsal view of the female ...... 28?
Hirstionyssus breviseta Strandtmann and I-Iorlan. Ventral view of the female ...... 289
Hirstionyssus breviseta Strandtmann and Morlan. Dorsal view of the female ...... 289
xii Figure Page
77. Hirstionyssus neotomae (Eads and Hightower). Ventral view of the female ...... 291
73, Hirstionyssus neotomae (Eads and Hightower). Dorsal view of the female ...... 291
79. Hlrstionyssus bisetosus Allred. Ventral view of the female ...... 293
80. Hirstionyssus bisetosus Allrod, Dorsal view of the female ...... 293
81. Hirstionyssus Stafford! Strandtmann and Hunt. Ventral view of the female ...... 295
82. Hirstionyssus staffordl Strandtmann and Hunt. Dorsal view of the female ...... * ...... 295
83. Hirstionyssus Stafford! Strandtmann and Hunt. Ventral view of tarsus II in the female ...... 295
xili INTRODUCTION
Tho genus Hjrstionyssus comprises a cosmopolitan group of nidicolous, haematophagous ectoparasites of small mammals. Careful study of the type specimens of previously described taxospecies of
Hirstionyssus of the Kearctic region, examination of numerous other specimens from many localities and hosts, and a review of the litera ture has revealed tho immensity of our ignorance of the systematica of this genus. Previous workers have largely ignored such biological information as host and locality data, resulting in numerous misidonti- fications and meaningless classifications.
This paper is tho result of an investigation into the systera- atics of fomales of the genus Hirstjonyssus of the Kearctic region, utilising the principles and methods of numerical taxonomy. Tho first part of this study deals with the numerical treatment of tho data.
Several methods of coding and transforming the basic data and soveral methods of analysis wore used, and the results are herein compared.
Attempts are made to determine tho phonetic interrelation ..hips of membors of tho gonus, and to construct an objective natural cla ssifi cation. The socond section is concerned with a taxonomic revision of tho genus Hirstionyssus. The taxonomy of tho gonus is reviewed, an identification key is constructed, and each recognised taxospecies is given a complete taxonomic treatment. The morphological terminology and chaototactic signatures used are from Evans and T ill (1965! 1966). Tho classification and technical names of North American mammals aro from M iller and Kellogg (1955)»
The numerical taxonomic treatment and terminology used in this study are basically that proposed by Sokal and Sneath (1963)* Subse quent to this presentation, numerous papers have been published on various phases of numerical taxonomy, of which only a few of the major works of pertinence to the present study wl.ll be cited.
Rohlf and Sokal (1965) made a study of the properties of two
coefficients of sim ilarity used in numerical taxonomy; Pearson*s
.product-moment correlation (Michener and Sokal, 1957) and Sokal*s taxonomic distance (Sokal, 1961). Their study was based upon analyses
of 97 species of bees by Michener and Sokal (1957) a^d h-8 species of mosquitoes by Rohlf (1962). They found that best results are obtained when all characters in a given study are standardized (Sokal, 1961)
(transformed so that each character w ill have a mean of zero and a
standard deviation of unity, thus attempting to relievo character
state values of the unequal v;eighting introduced by the use of differ
ent units of measurement). They also concluded that when most charac
ters used in a study aro measurements of various parts of an organism
and if tho OTU's differ much in over-all size, then correlation
coefficients should be used rather than distance. They suggested that
characters might be converted to ratios independent of size and then
distance coefficients used with good results. Sokal and Kiehoner (I967) conducted a sim ilar investigation using the Hoplitjs complex of bees which was originally analysed by
Kichener and Sokal (1957)* Their results support those of Rohlf and
S o k a l ( o p . c i t . ), but in addition they found that differences in clustering procedures affect the resulting classification. They observed differential weighting of stems during clustering to cause distinct effects, particularly with matrices of distance coefficients.
Only minor differences existed between pair-group and variable-group clustering methods.
Crovello (1968) investigated the effect of change of technique at two stages of a numerical taxonomic study based on data from 30
species of willow, Salix, for 131 morphological characters. The first
stage studied was the formation of the basic data matrix, a character by taxon table, by two methods, Tho first method takes into account
intrataxon variation of a character (two-dimensional character state
distance using both tho moan and standard deviation), while the second
method estimates only contral tendency (onc-dimonsional character stato
distance using the mean). Tho socond stage of his study involved tho
comparison of two methods of transforming characters: standardization
versus condensation (a transformation of character states to range
from zero to one). Crovello's results indicate that changes in tech
nique do mako a difference in the final taxim etric outcome, but the
magnitude of difference varies at the difforont levels of taxonomic
organization. He noted that tho agreement of his classification of Salix by numerical methods with that arrived at by conventional means was good.
Application of Numerical Taxonomy to the Acarl
To date there have been few applications of numerical taxonomic methods to acarine systematics. Preliminary papers by Funk (1963*
196*0 and Johnston (196*0 present discussions on the pertinence of numerical taxonomy to tho classification of the Acari. In a study by
Gheals (1965)» an analysis of the Hypoaspis-Androlaelaps complex obtained by numerical methods was contrasted with a conventional classification. His simple matching sim ilarity coefficient matrix was analyzed by means of both single linkage cluster analysis and vector (principal components) analysis. Although there was some agroemont between the results of these two methods, the overall pic ture was different. Results of the vector analysis were closer to conventional interpretation but differed sufficiently to suggest that these conventional interpretations were in need of re-ovaluation. He made no attempt to propose a classification based on his results.
Funk ( I 968 ) has applied numerical taxonomic methods to the
Celaenopsoidea, a large group of trigynaspid m ites. Funk used distance and correlation coefficients based on standardized data, and analyzed by moans of the unweighted pair group method (UFGKA) of cluster analy
sis . His study provided evidence in favor of tho exemplar method at the genus and family levels and the use of many equally weighted charac ters In numerical taxonomy (see Sokal and Sneath, 1963 ). This study also indicated that OTU relationships in a phonogram must bo carefully- analyzed with reference to the original sim ilarity matrix boforo final
classificatory conclusions can be drawn. Funk’s studies have led to
the recognisation of four distinctive new fam ilies and several new
genera in the Colaenopsoidea.
Recently Moss (1966j 196?; 1963) concluded a numerical taxonomic
study of the family Dermanyssidae (£. s tr . ), comprising 17 OTD’s
(Operational Taxonomic Units), including the females of a ll described
species of the genus Dermanyssus and one representative of tho genus
Lipon.yssoides. He computed correlation and distance coefficients based
on standardized data of 135 characters from all areas of the body, and
analyzed these coefficient matrices by means of cluster analysis,
graphic analysis, contour analysis, and centroid factor analysis. Ko
compared groupings formed by five methods of cluster analysis on corre
lations and four methods on distances with rospect to their congruoncios
and differences. These various clustering techniques were compared by
computing their correlations with each other, using cophonotic values
obtained from thoir respective phonograms (method of Sokal and Rohlf,
1962). The Unweighted Pair Group Method (UPGKA) and tho V.'oighted Pair
Group Method (l/PGMA), both with average linkage, wore closest to each
other in their placement of OTU’s. Moss found the UPGKA cluster analy
sis based on distances to have the highest cophenetic correlation
coefficient of any method. Graphic and contour analysis aidod only
in providing alternative ways of vievring the OTU clusters. The
centroid factor analysis gave pleasing results, and it was on this 6 that Moss based his final classification although relationships expressed in graphs and phonograms of the sim ilarity coefficients were taken into account. As a result of his study, Moss proposed several taxonomic changes in the family, including the recognition of a new subgenus and one new species.
The Genus Hirstionyssus
The species now comprising this genus of parasitic mites were first recognized as belonging to a distinctive homogeneous group by
Fonseca (19^8), who erected the genus and relegated 18 species to it.
Prior to this time the several described species were placed in various related genera.
Tho first taxonomic treatment of the North American species of the genus Hirstionyssus was by Jamoson (1950b). Jameson considered
Hirstionyssus as a subgonus of Nooichoronyssus Fonseca, 19^1, on morphological and ecological grounds. This arrangement was not accepted as valid by subsequent workers. Jameson suggested a classifi cation based on morphological characters but supported by biological characters (host distributions and geographical origins). He recog nized that some species from related hosts show relationships in certain morphological characters, and attributed this host specificity to "a coeval evolution of host and parasite." Jameson attempted to show
those relationships between species in a key, which included eight
s p e c ie s . Strandtmann and Korlan (1953) rejected tho generic and sub- generic classification of Jameson (1950b) in their treatment of tho known species of the world. They presented 2h species in a key to females, 12 of which were reported from North America, Their key was based on the most obvious morphological characters and is very inaccurate.
In her book treating the Gamasid Mites of the U.S.5.R., Bregetova
(1956) presented keys to known males and females of the U.S.S.R., w ith accompanying illustrations. There is no indication that biological data were taken into consideration in the construction of tho keys.
In their studies on tho British Dermanyssidae (s^. la t. ), Evans and T i l l (1966) erected a separate subfamily, Hirstionys sinae, for the gonus Hirstionyssus and four related genera. In their treatment of this new subfamily they gave diagnoses of the subfamily and the genus
Hirstionys sus, and descriptions of the six species "occurring in Britain.
They also included koys to males and females, and detailed illustrations
of each species.
The most recent work restricted to North American Hirstionyssus
is contained in Allred and Bock's (1 9 6 6 ) "Mites of Utah Mammals." This
treatment deals only with those species occurring in Utah. A key to
the females of 19 species, 10 of which are described as new, is given
along with host, locality, and seasonal distribution data. No attompt
was made to develop a natural classification based upon both morpho
logical and biological data. During the early stages of this study as the data were being recorded, definite relationships were observed between morphological characters of similar taxospecies and the hosts from which they were collected. As a result, the individual specimens (OTU's) used for the numerical taxonomic analyses were arranged alphabetically within major host groups and are listed in this same order throughout the first part
of this study.
The recognition and interpretation of phonetic relationships within the gonus Ilirstlonyssus and the comparison of this with bio
logical data is far overdue. Such a study would seem to be a most
worthwhile task and one well suited to the techniques of numerical
taxonom y. MATERIALS AND METHODS
J Cate r i a l s U t i li z e d
Preserved material of a ll the described Hoarctie taxospecies of Hirstionyssus, and a considerable amount of unidentified or tenta tively Identified material was obtained from several museums and private collections. Representative specimens of all previously described taxospecios and specimens of some undescribed material
occurring in North America were examined and included in the study.
Holotype specimens of a ll but four North American taxospecies were
examined and included. The exceptions were a cotypo of H, occidentalis
(Ewing), and paratypos of H, cynomys (Radford), H. obsoletus Jameson,
and H, staffordi Strandtmann and Hunt, Representative specimens of
species with a Holarctic or cosmopolitan distribution were examined.
Specimens of _H, isabellinus (Oudemans) and H. latiscutatus de Keillon
and Lavoipierre (= musculi Auct. ) from various parts of the world were
obtained. One specimen of H, carnifex (Koch) from the U.S.S.R., and
several specimens of H. talpae Zemskaya (= arcuatus Auct.) from Europe
and Asia vie re included to compare with North American material previ
ously identified as "H. carnifex" and "H. arcuatus". Duplicates of
most taxospocios were included in order to provido an estimate of tho
effect of intrataxon variation (a test of the "exemplar" method). In 10 many cases these duplicates were from different hosts and localities.
Twenty replicates of H. isabellinus from throughout the Holarctic region and from various hosts were included to give a more detailed examination of intrataxon variation, and to provide some estimate of the effect of different hosts and localities. Specimens were chosen for inclusion in the numerical taxonomic analysis also on the basis of tho quality of the slide preparations. Only specimens for which a ll characters could be scored were chosen, that is, in the basic data matrix no NC values appear. A list of tho 105 OTU's, with their collec tion data, is given in Appendix X. The OTU codes consist of the first four letters of a preliminary taxospecies identification plus an arabic numeral to distinguish between replicates of a single taxospecies,
Kost of the material examined in this study had been previously mounted on slides. Unmounted specimens and slide specimens requiring
remounting were mounted in Hoyer’s medium and ringed with clear finger
nail polish. 'Kites wore examined with both American Optical and Wild
phase contrast microscopes. Keasurements wore mado using a micromotor
disc in the eyepiece of an American Optical microscope equipped with a
floating stage which facilitated fast, easy orientation of specimens.
Tho micromotor units wore immediately transformed into microns with tho
use of a circular slide rule and then recorded on data sheets. Illus
trations wore prepared with tho aid of a Wild drawing tube, Treffonborg
type, and then photographically reduced to tho desirod size. 11
Characters U tilized
In choosing the final set of characters for this study, repeated reference was made to the definition of a character as given in Sokal and Sneath (1963) i.*®.* • a feature that varies from one individual (OTU) to the next and cannot be logically subdivided further. An attempt was made to obtain as many qualitative characters as possible but quantita tive characters proved to be much easier to find, thus a majority were
of this type. Characters from all areas of the body were utilized and no attempt was made to test the hypothesis of non-specificity. An
effort was made to obtain approximate equal numbers of characters from
four major areas of the body, but this proved to be impossible in the
case of gnathosomal characters. Originally 184 characters were con
sidered, but this number was finally reduced to 152 by eliminating
invariant characters and those shovring groat variability, i.o_., mea
surements between body parts which wore obviously dependent upon degree
of engorgement. The final list of characters recorded are contained in
Appendix II, and aro summarized in Table 1, Tho character state values
were punched on computer cards for processing. This basic data matrix
of character state values for the 105 OTU's is presented in
Appendix III,
Computational and Analytical Kothods
Tho U niversity of Kansas Numerical Taxonomy System, N'TSYS, was
used in most of the treatment of tho data, Tho preliminary programs
which included the computation of ratios and the condensation 12 transformation wore written by the author and processed at the Ohio
State University Computer Center. All other processing of data was carried out at the University of Kansas Computation Center under the direction of Dr. F, J. Rohlf.
Table 1, Partitioning of characters.
Quantitative
Area of body Qualitative I-jBristlc Continuous Totals
I. G nathosoma 1 1 15 17 (1 1 .2 $ )
II. Idiosoma (ventral) 10 0 34 44 (28,936)
Sternal area 3 0 14 17
Genital shield 3 0 7 10
Anal shield area 1 0 12 13
P e ritre m o 3 0 1 4
III. Idiosoma (dorsal) 12 1 32 45 (29.856)
IV. Logs 14 0 32 46 (30.1J6)
T o ta ls 2 113 37 113 152 (2 4 ,3 $ ) (7 5 .7 * )
The basic (raw) data matrix (character X OTU) was transformed in
three differont ways prior to the computation of sim ilarity coefficients,
resulting in throe separate transformed data matrices. Thoso three matrices and the computations involved in producing them were as follows: (1) Standardized Basic Data (SBD) matrix generated by the standardization computations as given in Sokal and Sneath (1963)|
(2) Condensed Basic Data (CBD) matrix computed by Crovello's (1968) method of condensation; and (3) Condensed Ratio Data (CRD) matrix which was generated by first converting a ll continuous quantitative character state values to a ratio of the length of the dorsal shield
(character 62), then condensing as in the CBD matrix above. For each of these three transformed data matrices, coefficients of Pearson's product-morcent correlation (r) and Sokal's taxonomic distance (d) were computed. The resulting matrices of OTU sim ilarity values were ana lyzed by the UPGKA (Unweighted Pair-Group Kethod with Average linkage)
.cluster analysis and graphic analysis. The results of cluster analysis
of both correlation and distance coefficionts for the three data matrices yielded a total of six phonograms (dendograms of phonetic
resemblance). From each phonogram, matricos of cophenetic values were
computed and compared with the original matricos of sim ilarity values.
This comparison, togother with the computation of a cophenetic corre
lation coefficient for each phenogram, provided an estimate of the
proportion of information transferred to the phenogram from the original
sim ilarity matrix (Sokal and Rohlf, 1962).
Subsequent to the above numerical taxonomic analyses, character
correlation coefficients (r) were computed for the 113 continuous
quantitative characters. This restriction to 113 characters was
partially imposed by the size lim itations of the particular program • 14 utilized, the BMD02D correlation with transgeneration program propared by the U.C.L.A. Health Sciences Computing Facility, and adapted for use on th e I,B,M. 7094 computer at the Ohio State University Computer Cen ter. Character correlation coefficients were computed for a 112 charac ter by 105 OTU ratio data matrix. This matrix was examined and analyzed for "highly correlated" characters, "Highly correlated” characters were defined to be those pairs of characters with correlation coefficients greater than 0.700. From a list of pairs of highly correlated charac t e r s , 37 w e r e chosen to be eliminated before recomputing OTU correla tion coefficients, A revised Condensed Ratio Data matrix of 115 charac ters by 105 OTU's was used in computing the OTU correlation coefficients.
The revised Condensed Ratio Correlation (revisod-CRC) matrix was then compared with the NTSYS Condensed Ratio Correlation matrix to dotermine what effect, if any, tho elimination of the highly correlated characters may have had on OTU sim ilarity values.
This revised-CRC matrix was used as the basis for a graphic analysis. This analysis provided a two-dimensional representation which aided considerably in the visualization of OTU relationships.
The graphic analysis in Moss’ ( 1967 ) numerical taxonomy study was used as a guide. It consists of constructing a graph diagram by Jotting down the OTU codes of each highly correlated group and drawing an arrow
(vector) from each OTU to that OTU with which it has the highest simi
larity value. When all highest sim ilarity values have been plotted then
second and third-order vectors are added where possible. The resulting network of arrows, produced by a process of triangulation, is of considerable use in determining the lim its of taxonomic groupings. A group of OTU's with high sim ilarity values among themselves are not connected by vectors to other such groups and thus form a closed sys tem. After groups of OTU's have been formed then the sim ilarity matrix is scanned to determine which OTU's serve as "links" between groups. Group connecting vectors are then added to complete the two dimensional graph diagram. Thus the graphic analysis is actually a two-dimensional representation of single linkage cluster analysis. In this study each OTU is represented by a circle containing the OTU code.
A ll v e c to r s betw een OTU's are drawn p r o p o r tio n a l in le n g th to th e magnA tude of the correlation coefficients. First-order vectors were repre sented by solid straight lines, second-order by dashed straight lines, third-order by dotted straight lines, and group connecting vectors of lower-order by solid wavy lines. RESULTS OF NUMERICAL TAXONOMIC ANALYSIS
Sim ilarity Coefficients
The computation of Pearson's product-moment correlation coefficient (r) and Sokal's taxonomic distance coefficient (d) from each of the three transformed data matrices resulted in six simi larity matrices. Those are designated as follows: (1) Standard ized Basic Correlation (SBC)j (2) Condensed Basic Correlation (CBC)i
(3) Condensed Ratio Correlation (CRC)| (4) Standardized Basic Dis ta n c e (SBD); (5) Condensed Basic Distance (CBD)i and (6) Condensed
Ratio Distance (CRD),
Because of the large size of each of these sim ilarity matrices it would be impractical to reproduce them here. All of the basic and transformed data matrices, the sim ilarity matrices, and the computer- produced phonograms are stored on magnetic tape and computer punch cards, and may be obtained in the form of computer print-outs or punch cards from the author or the Acarology Laboratory, College of Biological
Sciences, Ohio State University, Columbus, Ohio 43210.
Cluster Analysis
A phenogram was produced from each of the six sim ilarity matricos using the UFGMA cluster analysis (Unweighted Pair-Group Method with
16 Average linkage). The closest approximation to a representation of the relationships among the OTU's that can be obtained is the matrix of sim ilarity. Relationships in a sim ilarity matrix are n-dimensional where n represents the number of characters over which the sim ilarity values are computed. In the present study the 105 OTU's are in a 152- dimension character space. Since a phonogram is only a one-dimensional re presentation of these multi-dimensional relationships, it is impossi ble for such a diagram to give an accurate representation of the simi larity matrix. For this reason it is imperative that the phonograms be
carefully analyzed with reference to the sim ilarity matrices from which
they vrere generated, Sokal and Rohlf (1962) described a method for
estimating the proportion of information transferred from a sim ilarity
matrix to the phenogram. It consists of computing a correlation (the
cophenetic correlation coefficient or cophenetic r) between the original
matrix of sim ilarity values and the new matrix of OTU relationships read
directly from the phenogram. It should be kept in mind that the magni
tude of the cophenetic r is only a measure of the information transfer
that has taken place as a result of the cluster analysis. It dees not
provide information on the content of the actual clusters or their
arrangement in the phenogram. Cophenetic correlations between the six
phenograms and their respective sim ilarity matrices wero computed
(Table 2). These results will be discussed further in the following
s e c t io n s . 18
Table 2, Cophenetic correlation coefficients between sim ilarity matrices and phenograms obtained from the three transformed data m a t r i c e s .
Coefficient of sim ilarity
Data matrix r d
Standardized Basic 0 .7 8 9 0 .8 0 2
Condensed Basio 0 .8 8 7 0 .7 6 2
Condensed Ratio 0 .9 0 7 0 .9 1 0
Comparison of Phenograms
Upon inspection of the six phenograms it was obvious that they
shared a number of sim ilarities in their major groupings. The extent to which groups were shared depended to a large degree on the clustering level under consideration. In order to make an objective comparison of
these major groupings among the phenograms, it was necessary to choose
an arbitrary, but consistent, level of the phenogram sim ilarity scale
at which a group defining line could bo drawn. In the present study the
criterion of group recognition was a line drawn across each phonogram at
the mid-point of the observed ranges of sim ilarity values. Any stem cut
by such a line was assigned a group number and the presence or absence
of this group noted in the different phenograms. With the aid of this
arbitrary criterion the phenogram groupings from the six sim ilarity
matrices were analysed (Table 3)» It should be noted here that when
the condensation transformation is used the value of the sim ilarity 19 coefficient usually lies between zero and one, whereas when standard ization is employed the range is from minus one to plus one for corre lation and usually from about zero to about three for taxonomic distance (Crovello, 1968).
Table 3* Phenogram groupings obtained from six sim ilarity matrices. Criterion of group recognition was a line drawn across each phonogram at the mid-point of the observed ranges of sim ilarity values (in parenthesis below sim ilarity matrix abbreviation). Presence of a group is denoted by a plus (+) sign and absence by a minus (-) sign. Recognition of group combinations is indicated by a bracket and an "X",
Sim ilarity matrix
Group Components SBC CBC CRC SBD CBD CRD ( 0 . 4 0 0 ) ( 0 . 490) ( 0. 500 ) ( 1 . 190) ( 0 . 252 ) (0 . 256 )
I FEMU + X> II THOM I + III GSOM + +
IV LONG + +
V HILL - i i :co + + +
VI TRIA + + +
VII AFFI 1 +
VIII AFFI 2 - AFFI 4 +
IX SUTA 1 - PUNC 4 +
X CYNO 1 - AHGU 1 +
XI MUSC +
XII ISAB 3 + - >X 20
Table 3. (continued)
Sim ilarity matrix
G roup Com ponents SBC C3C CRC SBD CBD CRD (0.400) (0.490) (0.500) (1.190) (0.252) (0.256)
X III ISAB 4 + - — — . - - 1 ► X >x XIV ISAB 5 + •a* --- - >x >x XV ISAB 1 - 1 4 + — - — — -
XVI ISAE 9-17 + - - -- -
XVII 03S0 + - - - - -
XVIII TARS .1 - UTAH 7 + - - - -
XIX UTAH 6 + - - - --
XX TORU + -- - - ► X -
XXI BREV + -- - - -
XXII BISE - TJEOT + -- + - +
XXIII CARN + + + + + +
XXIV HSPB 1 - TALP 8 + + + + + •h
XXV NSPA + + + + +
XXVI STAF + + + + + +
Total number of groups recognized 25 7 8 10 10 12
All the phenograms, with the exception of the SBC phenogram, were fairly consistent in thoir formation of major groupings at the recogni tion level chosen. In the SBC phenogram 25 separate groups were recognized while in the others the number ranged from 7 to 12. Four groups (XXIII, XXIV, XXV, and XXVT) were consistently recognized in a ll phonograms. Groups I through IV ranged from being recognized separately to being combined into one large group. The most common situation was the combination of group I with II and group III with
IV, Groups V and VI were recognized as separate in four phenograms but were combined in the other two. Groups VII through XXII were variously combined in a ll except the SBC phonogram. The sim ilarity between the major phonogram groupings and the major host groups designated in the beginning of this study was quite striking. Pheno
gram groups I-IV and V-VI were synonymous with the Geomyid host group and the Heteromyid host group, respectively. Group XXVI was the
Carnivore host group, and the Insectevoro host group corresponded to
phenogram groups XVII, XXIII, and XXIV. In the latter case group
XVII was removed from the other two. Congruencies within the remainder
of the phenogram groupings were not as evident—members of the Scinrid
and Ihiroid host groups wore variously grouped together.
Clustering within the major groups and the joining of one group
with another was somewhat variable in the phenograms. V.Tith respoct to
the placement of OTU's and OTU groups, the SBC phonogram was the most
variable, and differed considerably from a ll others. The S3D and CBD
phenograms wore quite sim ilar to each other, differing in the placement
of several OTU groups, but were quite different in overall structure
from the other phenograms. The close relation between the SBD and the
CBD phonograms may bo explained by a sim ilar comparison made by Rohlf 22 and Sokal (1965)* They found a close relation between taxonomic dis tance based on standardized and unstandardized characters. The explana tion given was that the main effect of standardization is to change, usually raise, both the mean and standard deviation of the distance coefficient, and changing the origin of the character axis does not change the distance between OTU's, Condensation would be expected to have a sim ilar effect on the distance coefficients.
The SBC, SBD, and CBD phenograms gave poor representations of
OTU relationships as evidenced by the rather low cophenetic correlation
coefficients (see Table 2), Clustering within many OTU groups was quite
sim ilar to that of the other three phonograms, but the joining of OTU
groups was quite different and variable. There is considerable evidence
that overall size differences among OTU’s accounted for the distortions
and OTU misplacements in these three phenograms. According to the
results of Rohlf and Sokal's (19 65 ) investigation of the properties of
correlation and distance coefficients, the ovorall size variability of
organisms in a given study affects the sim ilarity coefficients, espe
cially taxonomic distance, in giving certain groups of characters
excessive weights. They found that whon most of the characters used
in a study are measurements of various parts of an organism, and when
the OTU's differ much in overall size, the distance coefficient w ill
be more a measure of sim ilarity in size than a moasure of overall simi
larity. Sokal and Michener (196?) also found the correlation coeffi
cient to have advantages over distance coefficients as measures of
sim ilarity for the Hoplitis complex of bees. In the present study 76 per cent of the characters were measure ments of body parts, and there -were considerable overall size differ ences in the OTU's included. This would surely tend to cause excessive weighting of somo groups of characters and result in distorted simi larity values. Observations in the present study support those of
Rohlf and Sokal (19&5) and Sokal and 1-h.chener ( 1967 ), and suggest that, under the conditions noted above, correlation coefficients may also be
more a measure of size than overall sim ilarity. For these reasons the
SBC, SBD, and CBD phenograms were considered to portray OTU relation
ships poorly, and thus were not included in any further analyses.
The CRC, CBC, and CRD phenograms were very sim ilar to each other
in all respects, differing only in the placement of two or throe OTU
groups and in the clustering of OTU's within several groups. These
three phenograms yielded quite high cophenetic correlation coefficients
(0.907, 0.887i and 0.910 respectively). Based on these results, as well
as the judgement of the author arrived at by a thorough examination of
the basic data matrix and the specimens (OTU's) themselves, the CRC,
CBC, and CRD phenograms were considered to give the best ono-dimonsional
illustrations of OTU phenetic relationships (Figures 1, 2, and 3)» From
a visual inspection of these throe phenograms it \ms observed that the
CBC and CRC phenograms were more sim ilar in overall structuro and place
ment of OTU's than either were to the CRD phenogram. This very close
correlation between the CBC and CRC phonograms indicates that, for
correlation coefficients, condensation of basic data alone produces
nearly the same results as calculating ratios for continuous quantitative characters prior to the condensation transformation. The computation of ratios seems to have little or no effect on correlation coefficients. From these observations it might be suggested that when most characters are measurements and the OTU’s differ in overall size, either correlation coefficients based on condensed data should be used, or the characters should be converted to ratios of some sort prior to the condensation transformation and then good results may be obtained from both correlation and distance coefficients,
OTU Placement in Phenograms
It should be kept in mind that a phenogram is only a one- dimensional illustration of OTU relationships. The vertical order of
OTU placement in a phenogram is therefore to a large extent arbitrary,
somewhat dependent upon the order in which the OTU’s were arranged in
the original input data matrix and necessitated by the requirements of
a two-dimensional diagram. Thus it would be possible to rotate the
stems of clusters without altering the accuracy or efficiency of the
phenogram .
In considering the placement of OTU's and OTU clusters in the
phenograms, the CRC, C3C, and CRD phonograms w ill be emphasized
(Figures 1, 2, and 3). OTU clusters w ill be referred to by the taxo-
speclos abbreviation or combinations thereof, i.e ., F3MJ group and FEMU
THOM group. Upon visually scanning the phenograms, six to eight major
clusters could be recognized. These major groups were designated by a
large capital letter which refers to the major host group(s) in which
the OTU's were placed in the beginning of this study (See Appendix I). Using the CRC phenogram as a starting point, those groups were as follow s: "G" group (FEMU 1 - LONG 3): "H” group (HILL 1 - TRIA 3 ) i
"S-M" group (AFFI 1 - NSOT 2); NSPA 1-2 cluster; "I” group (CARN 1 -
TALP 9)j and "C" group (STAF 1 - *+). These same basic groups were also recognized in the CBC phenogram, except that the NSPA 1 - 2 cluster was placed immediately following and joined to the "H" group*
The principal differences noted in the CRD phenogram was the recogni tion of the TRIA cluster (placed near the bottom) and the CARN 1 OTU
as separate and distinct. The CARN 1 OTU could be considered as
separate in the other phenograms except that it joined the main stem
of the "I" group.
Clustering within the "G" group was almost identical in a ll
three phenograms. The only difference was in the level of clustering,
which would be expected to vary a little from one phenogram to another
in all groups. The FEMU cluster joined the THOM cluster and the GEOM
cluster joined the LONG cluster consistently prior to the formation
of the major "G" group. This was also the case In the SRC phenogram
(not illustrated). In the SBD and CBD phonograms (not illustrated)
the FE?>TJ - TITOM cluster and the GEOM - LONG clustor were formed, but
they failed to join each other as in the other phonograms. These
four OTU clusters ropresent four previously described taxospccies
which were tentatively placed togother in the Geomyid host group in
the beginning of this study.
Within the "II" group the HILL and IMCO clusters wore formed
and joined each other in exactly tho same mannor in all six phonograms. 26
The TRIA cluster joined the HILL - IKCO cluster in the CRC and CBC phenograms, but in the CRD phenogram was placed just above the "C" group. Upon close examination of the sim ilarity matrices of all three phenograms it was quite evident that the joining of the TRIA
cluster with the HILL - IIICO cluster was the most accurate repre
sentation of the phenetic relationship. These three OTU clusters
represent the three taxospecies originally placed in the Heteromyid
host group.
It should be noted that within the HILL cluster OTU's 4 and 5
consistently clustered together in all six phenograms before joining
the HILL 1-2-3 cluster. In the beginning of this study these two
OTU's were observed to differ from the other throe in at least two
readily recognizable qualitative characters as well as some quanti
tative characters. This situation w ill be analyzed and discussed in
more detail later.
Because of the averaging effect in the clustering procedure,
the phenograms gave little indication of the affinity which the "G"
and "H” groups had to each other or to other major groups. The "H"
group consistently joined the "S-M" group, whereas the "G" group
joined the main phenogram stem just pi’ior to the joining of the "C"
group. Upon visually scanning the CRC, CBC, and CRD sim ilarity
matrices, however, it became quite apparent that the "G" group and
the "H" group had members with mutually high sim ilarity values for
each other, thus indicating a definite affinity between these two
major groups. The joining of the "H" group with the "S-M" group 27 was due prim arily to the averaging effect of the UPGKA cluster a n a l y s i s .
The "S-M" group, which contained a majority of the individuals, was a conglomerate of OTU's, some of which were originally placed in
the Scuirid host group and some in the Muroid host group. The first
definite subgroup within this major group consisted of the AFFI 1 -
AUGU 1 cluster. This cluster was recognized and occupied the same
position in all three illustrated phenograms. It is composed of
several previously described, closely related, taxospecies originally
grouped together in the Sciurid host group. OTU clustering within
this group varied considerably from one phenogram to another, with,
but two exceptions. The PUNC 1-2-3 cluster and the cluster comprising
CYNO 1, IMVA. 1-2, and AHGU 1 wore recognized in a ll throe phenograms.
Formation of the latter cluster was identical in all six phenograms,
and it consistently joined the remainder of the group in the same
manner. The FUKC 1-2-3 cluster varied from one phenogram to another
in its joining of other OTU’s. Because of the clustering inconsis
tencies from one phenogram to another in tho greater part of this group,
further detailed studies were necessary. The phonetic relationships of
this group will be dealt with in later analyses.
Two other taxospecios were originally relegated to the Sciurid
host group, but clustered elsewhere in all the phenograms. These were
the TORU and the ITS PA clusters. The TORU cluster consistently joined
the BRBV cluster in all tho phonograms. An inspection of the simi
larity matrices on which tho three phenograms were based revealed that 28 the TORU OTU's were secondarily most sim ilar to members of the CYNO -
IITVA - AKGU cluster discussed above. The placement of the TORU - BRSV cluster was the same in the CRC and the CRD phenograms — immediately following ORSO 1 and 2, whereas in the CRC phenogram it was placed between the UTAH and the ISAB OTU groups. Further analyses wore neces sary to ascertain the most correct placement of the TORU cluster, thus further discussion of its phenetic relationship with other OTU's w ill be deferred until later.
The two NSPA OTU's which represent an undescribed taxospecies, were also placed in the Sciurid host group in the beginning of this
study. This two-OTU cluster joined the "H" group in tho C3C phonogram, but in all others was placed near the bottom, joining one of the last
major stems. Its varied placement in each of the six phenograms raised
a question as to its nearest phenetic relationship. Upon examining the
CRC, CBC, and CRD matrices it was found that the NSPA OTU's had a
greater affinity for the TORU cluster than to any other. This illu s
trates very well the lim itations of cluster analysis in expressing such
relationships in one-dimension. Many misplacements of OTU groups in
phenograms are due, at least in part, to the averaging effect of the
UPGKA cluster analysis.
The second recognisable subgroup within the "S-M" group con
sisted of the LATI and TARS - UTAH clusters. It was with this sub
group that the AFFI 1 - ANGU 1 group, discussed above, consistently
joined. Clustering of the OTU's within both the LATI and the TARS -
UTAH clusters varied a little from one phenogram to another, but in 29 a ll three phenograms the two clusters joined each other prior to the joining of any other stem.
The group consisting of the TORU and the BREV clusters followed the LATI - UTAH group in the CRC pehnograra, but could have, just as accurately, been placed following the OBSO cluster as in the CBC and
CRD phenograms. The CBC phenogram differed most in the placement of the TORU - BREV group. In this phenogram the group joined tho BISE -
NSOT group rather than the main stem of the "S-K" group.
The largest subgroup within the "S-M" group was the ISAB
clusters. As noted earlier, this group consisted of 20 replicates
of a single taxospecies from various hosts and localities throughout
tho Holarctic region. Clustering within the ISAB group varied from
one phenogram to a n o th er. The placem ent o f some OTU's was c o n s is t e n t
whereas others varied considerably. A more detailed analysis and
discussion of the intrataxon variability of this group w ill be deferred
until later.
In a ll phenograms the two 0BS0 OTU's, which were originally
placed with the Insoctivore host group, clustered with the ISAB group
just before joining tho main "S-M" stem. Upon examining the sim ilarity
matrices it was found that the 0330 OTU's had their greatest affinity
for some TSAD OTU’s, but were secondarily most sim ilar to members of
the "I" group. The placement of the 0330 OTU's w ill bo discussed
further in the graphic analysis.
The BISE and IIEOT clusters formed a rather distinctive subgroup,
b e in g th o l a s t t o jo in th e la r g e "3-1!" group. C lu ste r in g w ith in th e 30
BISE - NEOT subgroup was identical in a ll phonograms, and was con sistently the last subgroup to join tho "S-M" group,
Tho "I" group was recognized in a ll phenograms as a rather homogeneous group in which several clusters were consistently formed.
The CARN 1 OTU was recognized as separate and distinct from a ll other
OTU's, In the CRC and CBC phonograms it joined the main stem of the
"I" group, but in the CRD phenogram it joined the main phonogram stem
just prior to the NSPA and TRIA stems. This placement of the CARN 1
OTU was not surprising, because it was early recognized as a very different OTU, It represents an old world taxospecies somewhat
related (phenetically) to some members of the "I" group. It was
included in this study to confirm the fact that, this taxospecies does
not occur in the Nearctic region, contrary to what some authors have
in d i c a t e d .
The RSFB cluster was recognized as separate in a ll phenograms,
and it joined the TALP - PALU clusters in the same manner. The NSPB
cluster represents an undescribed taxospecios closely related to the
TALP - PALU group. Clustering of tho TALP and PALU OTU's was somewhat
variable in the thi'ee phenograms. The TALP i, 2, 3» and PALU 1 OTU's
clustered together in all three phenograms, and tho TALP 5» 6> 7» and
8 OTU's clustered together. Talp b and 9 were placed differently in
each phonogram, V/ithin the TALP - PALU group were 10 very closely
related OTU's, some of which were Ilearctic in distribution, some
Palearctic and one Oriental, Generally tho Nearctic and Palearctlc
OTU's clustered separatoly, with but two exceptions — PALU 1 and 31
TALP 9. The TALP 1, 2, and 3 OTU’s were Palearctic in distribution*
TALP 4 was Oriental, and all others were Rearctic, Relationships within this group w ill bo dealt with in more detail in the graphic
a n a l y s i s .
The final major group to be considered was the ”C” group or
3TAF clustor. These four OTU's represent a single taxospecies para
sitic on Kephitine Carnivores, This group clustered vrith no other
group, and, as Indicated in all the phenograms, is only distantly
related (phene tic ally) to the other members of tho genus in the ITearctic
region. This conclusion was supported by examination of the sim ilarity
matrices, in which sim ilarity values between the STAF OTU's and a ll
other OTU’s indicated very low affinity.
Graphic Analysis
Subsequent to the classical numerical taxonomic analyses
discussed above, character correlations were computed for the con
tinuous quantitative (measurement) characters in the manner described
earlier. Upon analysis of the character correlation matrix several
groups of highly correlated characters (pairs of characters with
correlation coefficients greater than 0,700) were noted. The three
major groups were as follows: ( 1) length of body setae, ( 2 ) distances
between setae and between setae and other body parts, and ( 3 ) w id th
and length of various appendages. Of the most highly correlated char
acters, 37 were selected for elimination as follows (see Appendix Tl):
6 , 8 , 26 , 27, 30, 31, 32, 35, 42, 44, 4 7, 4 8 , 4 9 , 55, 56 , 61 , 62 , 75, 32
77, 80, 81, 82, 8^ , 96, 100, U 6 , 122, 125, 135, 1 ^0 , 1^-1, 1^2, lhb,
1^5, 1^7i 1^9» and 151.
A revised OTU correlation coefficient matrix was computed using a revised condensed ratio data matrix of 115 characters by 105 OTU's,
A visual comparison was then made between the revised Condensed Ratio
Correlation (revised-CRC) matrix and the NTSYS Condensed Ratio Corre lation (HTSYS-CRC) matrix. In most cases the differences were very
small; some sim ilarity values were higher and some were lower.
The revised-CRC matrix was used as the basis for a graphic
analysis, which provided a two-dimensional representation of OTU and
OTU cluster relationships (Figures 5 , £nd 6 ) . The methods used in
this analysis have been described earlier. In these two-dimensional
graphic diagrams the OTU's are represented by circles containing tho
OTU codes. Relationships between OTU's are indicated by vectors
(arrows), proportional in length to the magnitude of the correlation
coefficionts read directly from the revised-CRC matrix. This method
of graphic analysis is basically a modified single linkage cluster
analysis with relationships illustrated in two-dimensions, Bocause
of the lim itations imposed by a two-dimensional diagram, the only
significant and reliable relationships between OTU's are those in
which OTU's are connected directly by vectors. That is, there is
usually no significance in the distance between two OTU's not directly
connected by a vector, Tho first-ordor relationship between clusters
of OTU’s is indicated by thoir positioning in relation to each other. 33 Thus tho relative placement of OTU clusters and groups has a certain amount of significance in the diagrams.
F ig u re U- served mainly to illustrate the OTU's with which the
STAF cluster had its highest correlations. The correlation r between
STAF 2 and TALP 3 was 0.215» and that between STAF 3 and PUNC h- was
0.193* The lowest correlation between the STAF cluster and another
cluster was -0,186, which was between STAF 1 and BISB 3. This simi
larity value was only O.hOl units on the sim ilarity scale below the
highest sim ilarity noted above. Thus the indication is that the STAF
cluster is very dissim ilar, in the context of the characters utilized,
to any other members of tho genus in Forth America, It should be
noted that the four OTU's comprising this cluster were very similar
to each other; All correlation coefficients were groater than 0.875*
the highest being 0.913 between STAF 1 and 2.
The graphic representation in Figure 5 aided considerably in
visualizing the relationships within and between the "G" and the "H"
groups. The single OTU's around the bottom and loft side of the
diagram (AFFI h, NSPA 2, ISAB 20, ISAB 18, and NE0T 3) were included
to show the relationship between the OTU's in Figure 5 and those of
F ig u re 6 , The joining of clusters in this graphic analysis differed
from that observed in tho phenograms in some respects, but generally
the analysis yielded sim ilar results. The major difference was in
the joining of the four clusters comprising the "G" group (FEMU, GEOM,
TKOH, and LONG), Tho GE0K cluster was placed closer to the FEKU
cluster than to the LONG cluster duo to the higher sim ilarity value between GEOM 2 and FEMU 2, The LOUG cluster was linked closer to the
THOU cluster than to the GEOM cluster as in the phenograms. The FEMU
cluster was most sim ilar to the TEOM cluster as was the situation in
the phenograms. Those differences are probably a result of the
different clustering methods employed — clustering in the phenograms
was by average linkage whereas clustering in the graphic analysis was
by modified single linkage. Another major difference was in the rela
tionship between the "G" and the "H” groups. In the phenograms no
linkage between these two major groups was evidenced. But in the
graphic analysis the "G" group was quite closely linked to the "H"
group. The THOM cluster had slightly greater affinity for members
of the "H" group than for other clusters in the "G" group. The THOM
cluster then served as tho bridge between the "G" cluster and the "H"
cluster. The three clusters forming the "H" group were more sim ilar
to each other then to any other cluster or group. Thus evidence s till
supports the recognition of the two major groups.
Upon visually scanning this diagram, seven distinct clusters
were recognized. These were identical to those recognized in the
phenograms. Clustering of OTU's within each of these seven clusters
was very sim ilar to that observed in the phonograms. The most out
standing exception was in the HILL cluster. The sim ilarity value
between HILL 1 and 5 was highest in the graphic analysis, whereas in
the phenograms there was a first-order linkage between HILL 1 and 3
and between h- and 5» It should be noted, however, that the sim ilarity
values between 1 and 3 » 1 and 5 » a ?id h- and 5 were very nearly the same, 35 differing by less than 0.015 units on the sim ilarity scale. This difference resulted from the elimination of the 37 highly correlated
characters prior to computing the revised-CRC matrix. Even though in
several clusters (GEOM and THOM) the OTU's were not closely clustered,
they were more sim ilar to each other than to any OTU of another cluster.
This w ill be considered further in the next section when intrataxon
variation is discussed.
Tho graphic analysis illustrated in Figure 6 was in fairly close
agreement with the phenograms, although relationships between clusters
were much better defined. The major phenogram group showing the least
a f f i n i t y in t h i s g ra p h was th e " I " g ro u p com posed o f TALP, PALU, and
FSFB, In this analysis the TALP 1, 2, 3# a^d h OTU's, which were
Falearctic and Oriental (TALP h) in distribution, clustered fairly
close together and somewhat separate from the remainder of the TALP
OTU's which vfere Kearctic in distribution. The Uearctic PALU 1 OTU
formed the major link botweon the two TALP clusters. The 113 PB cluster
was linked to the TALP - PALU cluster through TALP 9* The graphic
clustering of OTU's in this group corresponded quite well with that
observed in the phenograms.
As in the phonograms, the OBSO cluster was placed closer to tho
ISAB group than to the "I" group, although it was secondarily most
sim ilar to the Palearctic TALP OTU's, The CAP.I! 1 OTU, w hich i s
Palearctic in distribution, could not bo accurately and correctly
placed in the graph. The broken wavy vectors indicate only tho direc
tion of highest sim ilarity, the greatest sim ilarity being with the Palearctic TALP OTU's. The main "I" group was most closely linked to the UTAH cluster through NSFB 3» Also, TALP h served as a link between the main "I" group and the LATI cluster.
The five LATI OTU's formed a distinct cluster which was closely- linked to the UTAH - TARS cluster. The sim ilarity between OTU’s within both of these clusters was quite high. The UTAH cluster was linked quite closely to EUTA 2, a member of the phenogram AFFI 1 - AHGU 1 group. The linkage of the LATI, UTAH - TARS, and AFFI - ANGU OTU groups was identical to that observed in the phenograms. The relation ship between the UTAH - TARS cluster and the EUTA 2 OTU was identically expressed in the CRD phenogram in which EUTA 2 was considered to be somewhat misplaced.
It was hoped that the graphic analysis would serve to clarify the relationships within the AFFI 1 - ANGU 1 group, but as is evident
from an inspection of Figure 6 this was not very well accomplished.
This analysis was congruent with the phenograms in separating the
CYNO - INVA - ANGU cluster from the main part of the group. In con
sidering first-order relationships only, the PUNC 1, 2, and 3 OTU's
formed a distinct cluster, with all other OTU's forming a cluster around
AFFI 3. The first-order relationship of FUNC ^ could not be illustrated
because it was with AFFI 3 also. The relative position occupied by the
OTU’s was the only criterion which served to clarify the relationships
at all. The four AFFI OTU's were adjacent to each other, the EUTA 1
and 2 OTU's were relatively close to each other, and the OCCI 1, 37 PURC i , 2 , 3 1 and 4 OTU's wore placed somewhat together. This rela tive placement of OTU's was based on first, second, and third-order
relationships.
Unlike the phenograms, the graphic analysis placed the TORU
cluster between the ISAB and the CY1IO - IRVA - ANGU clusters, but
linked close to the former. In the phonograms the TORU and the BREV
clusters were consistently linked. Upon examining the character state
values of tho 37 characters eliminated in the computation of the
revisod-CRC matrix it became evident that some "length of setae" char
acters was primarily responsible for the relationship between the TORU
and BREV clusters. Elimination of these "length of setae" characters
resulted in this change in phonetic relationship. The relationship
between the TCRU cluster and the CYNO - INVA - ANGU cluster in tho
graphic analysis was supported by the two-way link between TORU 2 and
ANGU 1.
The NSPA OTU's were most sim ilar to TORU 3* This relationship
-was actually second-order, with the third-order being with HILL 5
(see Fig. 5)* This phonetic relationship was not illustrated in the
phenograms because of the effect of average linkage.
The la r g e c lu s t o r o f 20 ISAB OTU's formed more or l e s s th e c e n te r
of the graphic diagram. The relationships among OTU's within this clus
ter were very difficult and sometimes Impossible to illustrate because
of the lim itation to two-dimonsions, Several first-order relationships
could not be illustrated, thus distorting the graph. One such distor
tion was with the OTU's surrounding ISAB 20; the first-order 38 relationship of ISAB 20 was with ISAB 13» and the first-order relation
ship of ISAB 1 was with ISA3 20. If a three-dimensional model of the
ISAB cluster were constructed, the positioning of the OTU's would be more or less in the form of a sphere, rather than spread out as in the
graphic diagram. Thus in examining the graph in Figure 6 this should be kept in mind.
The BRBV cluster was most closely linked to the ISAB group.
The second highest group linkage (which is not illustrated in the graph
diagram) was with the TORU cluster. As noted above, in the phenograms
the BRBV cluster was linked with the TORU cluster consistently. This
difference in order of linkage was due in part to the elimination of
some characters as discussed above, and in part to the difference
between average linkage and single linkage.
The final group to be considered was the IIEOT and BISS clusters,
'•/hen considering only first and second-order relationships, the BISE
OTU's formed a closed cluster and the N30T OTU's clustered together,
'/hen third-order relationships wore added, the two clusters linked
closely. First-order relationships between OTU's within those two
clusters were almost identical with that observed in the phenograms.
The IIEOT - BISE group was linked to the ISAB group, with NEOT 2
serving as the closest link and BISE 2 as tho second closest. DISCUSSION
Establishment of a Classification and Correlation of tho Numerical Taxonomic Analyses with Biological Data
Unfortunately a certain amount of subjectivity enters into the choice of which analysis to use as the basis for the establishment o f a classification. Other numerical taxonomic workers have faced this same problem, and most were influenced in their decision by their Intui tive impressions of relationships within the group which they studied
(Rohlf, I962i 19^3; Funk, 1963; Koss, 1967). The recognition 01* d e f i n i tion of taxospocies by drawing a phenon line across a phenogram was found to bo impossible in the context of the characters and OTU's chosen in this study. A phenon line drawn at the mid-point of the observed ranges of sim ilarity values for each phenogram served to delim it most major groups (see Table 3)» but moving the phenon line to the right would only serve to make the recognition of subgroups and taxospecies extremely difficult if not impossible. Thus the phenograms have been used mainly to aid in the recognition of major groups, and to a much lesser extent in delimiting clusters which represent taxospecies. The classification arrived at in this study was based primarily on the graphic analysis. Some decisions especially with regard, to tho recogni tion of taxospecies have beon made on utilitarian grounds. Host and
39 40 locality data, as well as the authors intuitive impressions of rela tionships within the genus, have surely had an influence on certain d e c is io n s .
In establishing the classification herein presented it was recognized that in order for it to be useful the various taxonomic categories must be defined by readily recognizable morphological characters. Thus in the following analysis of numerical taxonomic results, taxon diagnosis is based primarily on the most definitive qualitative characters (Table 4), In the following discussion the major groups w ill bo considered in the order of Increasing sim ilarity for other major groups. That is, the groups most dissim ilar to all others w ill be considered first.
The "C" group (Carnivore host group). - This cluster is comprised of four OTU's representing four replicates of a single
previously described taxospecies, H. Stafford! Strandtmann and Hunt, which is parasitic on Carnivores of the subfamily Kephitinae (skunks).
STAF 1 and 2 were paratype specimens, both collected from the same
host species and approximate locality (see Appendix l). In a ll analy
ses these two CTU's had the highest sim ilarity value, STAF 3 and 4,
which were from difforent hosts and localities, consecutively joined
the STAF 1-2 cluster. The H, Stafford! taxospecies differs from all
other Fearctic members of the genus in a number of characters. The
most definitive qualitative characters were t
Char. Ho. 16, 17: Doutosternal denticles 11 in number
arranged in single file j Table 4, Katrix of character state values of 30 qualitative characters for the 25 recognized taxospecies, The major groups are designated and the taxospecies are listed in the same order as discussed in the text. Taxospecies abbreviations are used to conserve space. Refer to Appendix II for a definition of the charac- te r s and character state values.
Character Numbers
Taxospecies 17 24 34 40 43 50 58 60 61 66 6? 68 69 70 72
"C" group STAB 1 4 2 4 3 1 1 5 1 4 0 2 2 2 0
i" group o-\ i r CM CM 1 F2MU 2 5 1 2 -4 2-3 1 1 1 3 1 2 2 2 1 G30K 2 4 -5 1 2-3 2 3 1 1 1 3 1 1 1 1 1 THOtf 2 5 1 4 2 3 1 1 1 3 1 2 2 2 1 LONG 2 2 1 3-4 2 2-3 1 1-2 1 2 1 2 2 2 1
"HM group HTI.L 2 6 1 2 -4 ' 2-3 3-4 0 3 2 3 1 2 2 2 1 "C" n sp 2 6 1 3-4 2-3 3 1 3 1 3 1 2 2 2 1 INC0 2-3 7 1 4 2-3 3 1 1-2 1 3 1 2 2 2 1 TRIA 2-3 6-7 1 3-4 2 3 2 1 1 3 1 2 2 2 1
group "B" n sp 2 6 1 3 2 3-4 1 1 1 1 1 0 2 C 1 TALP 2 6 -7 1 4 2 3-4 1 1 1 2-3 1 2 2 0 1 Table if. - continued
Character Numbers
Taxospecies 17 24 34 40 43 50 58 60 61 66 67 68 69 70 72
"S-M" group
"S" subgroup AFFI 2 6 1 if 2 3 1 4 1 2 1 2 2 2 1 "D" n sp 2 6 1 2 -4 2-3 3-4 1 2-3 1 2-3 1 2 2 2 1 OCCI 2 6 1 2-if 3 3—if 1 3 1 2-3 1 2 2 2 1 BUTA 2 6 1 4 3 3 1 3 1 3 1 2 2 2 1 cyno 2 7 1 if 2-3 3-5 1 2 1 2-3 1 2 2 2 1 UTAH 2 6 1 3-4 2 3-4 1 1 1 2 1 2 2 2 1 LATI 2 6-7 1 5 2 3-if 1 1 1 2-3 1 2 2 2 2
"V" subgroup "A” n sp 2 7 1 if 3 3 1 2 1 2-3 1 2 2 2 1 TORU 2 6 1 2-4 2-3 3 1 1-2 1 2-3 1 2 2 2 1 0BS0 2 6 -7 1 5 2 3 1 1 1 1 1 2 2 2 1 ISAB 2 6 1 5 2-3 3-if 1 1 1 2-3 1 2 2 2 1 BRSV 2 5-6 1 2-if 1 3-if 1 2 1 1-2 1 2 2 2 1 N30T 2 6 1 2-3 1-2 3-if 1 1-2 1 2-3 1 2 2 2 1 BISS 2 5-6 1 1 1 3-if 1 1 1 2 1 2 2 2 1 Table 4, - continued
Character Numbers
Taxospecies 73 74 75 109 111 114 117 120 127 128 129 130 131 133 13^
"C" group STAF 0 0 1 1-3 0 1+ 2-3 4 3-4 0 1 0 0 0 1
3" group F2KU 0 1 0 0 0 1-2 2-3 1-2 0 1 1 1 0 1 1 GSOK 0 1 0 0 0 1 1-2 1 0 0 1 0 0 1 1 THOM 1 1 0 0 0 2 2-3 1 0 0 1 0 0 1 1 4 lo ::g 1 1 0 0 0 3 2 1-3 0 0 1 0 0 1 1
i " group ■*-1 CM HILL 1 1 0 0 0 1-2 1-2 1 0 0 0 0 0 1 1 "C" n sp 1 1 0 0 0 2 2 3 0 0 0 0 0 1 3 INCO 1 1 0 0 0 1-2 1-2 1-2 0 0 0 0 0 1 1 TRIA 1 1 0 0 0 1 1-2 1 0 0 0 0 0 1 3
[" group "B” n sp 0 0 0 0 0 1 0 1 1-2 0 1 0 0 1 1 i r cu TALP 0 0 0 0 1 1 0 1 2-3 0 1 0 0 1 1 Table 4, - continued
Character Numbers
73 74 75 109 111 114 117 120 127 128 129 130 131 133 134
1 1 0 0 0 3 0 3 2 0 1 0 1 1 1 1 1 0 0 0 3 0 3 0 0 1 0 1 1 1 1 1 0 0 0 3 0 3 1-2 0 1 0 2 1 1 1 1 0 0 0 2 0 2 2 0 1 0 1 1 1 1 1 0 0 0 1 0 1 1-2 0 1 0 2 1 1 1 1 0 0 0 1-2 0 1 1-2 0 1 0 0-1 1 1 1 1 0 0 0 1 0 1 1 0 1 0 0 1 1
JL 1 0 0 0 0 0 0 3 0 0 0 0 1 1 l 1 0 0 0 3 0 1 1 0 0 0 1 1 1 0 - 1 1 0 0 0 3 0 3 0 0 0 0 0 1 1 1 1 0 0 0 1 0 1 0 0 0 0 0 1 1 1 1 0 0 0 0 0 1 1 0 0 0 0 1 1 1 1 0 0 0 1 - 2 0 1 1 - 2 0 0 0 0 1 o 1 1 0 0 0 1 0 1 0 0 0 0 0 1 o 2b: Posterior margin of sternal shield irregular*
3b: St. 3 setae off sternal shield*
50: Position of paranal setae at anterior end of
anal field*
60: Feritreme extending only to anterior half of
coxa I I I ,
6 6 : Posterior end of dorsal shield forming narrow,
caudal projection,
67: Dorsal setae z 6 absent*
72: Dorsal setae J2 absent,
75s One pair of R series setae on dorsal shield
margin*
109: Blunt to rounded bifid spur on coxa I,
lib, 120, 127: Inner spur on coxae II, III, and IV usually
bifid and blunt,
133: Seta pdg on genu IV absent.
Bocauso of these many differences, as well as the dissim ilarity expressed
In the phenograms and graphic diagrams, it Is proposed that H. Stafford! be relegated to a separate subgenus, thus creating tiro subgenera for the genus Hirstionyssus in the ITearctic region.
The "G" and "H" groups. - Tho inclusion of those two rather distinctive groups together is based on the graphic analysis (Fig. 5)-
These two groups together can bo distinguished from all other members of the genus (except H. staffordi) on the basis of one very diagnostic c h a r a c te r : U6
Char, No. 117: Presence of postero-lateral spur on coxa II.
Two other characters not restricted to these two groups but shared by all members of both are as follows i
Char. No. 127: Without spur on coxa IV;
131: S eta av^ on ta r s u s IV norm al, long and
s le n d e r.
The "G" group (Geomyid host group). - As noted previously, the phenograms consistently separated this group from all others. The four clusters of the "G" group, representing four previously described taxo- specios parasitic on rodents of the family Geomyidae, are as follows:
F2MU 1-^ - H. femurails Allred
GEOM 1-2 = H. peonydls (Keegan)
THOM 1-2 = H. thomomys A llred and Reck
LONG 1-3 = H. longichelae Allred and Beck
These taxospecies shared the following characters, which are different from those found in the "H" group:
Char. Mo. Zkx Posterior margin of sternal shield moderately
convex to slightly concave;
129: Setae av- and pv, on tarsus II stout and 1 1 c la w -lik e .
In the phenograms, femuralls and thomomys consistently clustered together as did geom.ydjs and longichelae, whereas in the graphic analysis geonydls clustered closer to fcmuralis. Based onqualitative characters alone, femuralls, poomydis, and thomomys are very similar, with longichelao difforing in four distinctive characters as follows: 1+7
Char. No. 2*+s Posterior margin of sternal shield decidedly
convex |
66 : Posterior margin of dorsal shield in shape
of broad, blunt wedgej
lll+, 115* Inner spur on coxa II short and roundedi
120 -122j Inner spur on coxa III very large and rounded.
II* femuralls and geomydis differ from thomomys and longichelae in one ch a ra c te r*
Char. No, 73* D orsal se ta e SI a b se n t, w ith Z l moved c lo s e r
to edge of dorsal shield.
Two diagnostic characters of femuralls are*
Char. No. 125: .’/ith blunt apophysis on femur III
130: Setae avP and av on tarsus II decidedly c 3 bulbous basally,
H. geomydis may be readily recognized by the following throe characters:
Char. No. 68-70: Setae r2, r3» and rl+ with normal setal bases
but with undeveloped setae.
H* ^murails was represented in the study by four replicates. F2MU 1 was tho Holotype whereas the other three were "representative" speci mens from different hosts and localities, FEMU 1, 2, and 3 were collected from the same general locality (southern Utah) whereas FEMU Ur was from a more distant area (southwest Toxas). Even though FGKU U- differed considerably in collection locality and host species, it clustered quite close to the other three in all analyses. h8
H. geomydis was represented by two OTU's. Both wars from tha same host species but different localities. G30K 1 was the Holotype, collected in east Kansas, and GSOM 2 was from southwest Texas. Tha variability between these two OTU's was greater than that observed within any other taxospecies. One would be tempted to conclude that these two OTU's may represent two separate taxospecies, but on the basis of available qualitative characters these two OTU's could not be separated. Thus they are considered to belong to the single taxo-
s p a c ie s.
The same situation was found in the case of the two replicates
of thomomys. Variability between these two OTU's was nearly as great
as that noted in geomydis. The Holotype, THOM 1, was collected in
northeast Utah while THOK 2 was collected from southeast Idaho.
The throe longichelae OTU's clustered fairly well together.
LONG 3 was the most variable, probably because of the difference in
locality which was nearly the same as that noted for the two THOM
OTU's. LONG 1 and 2 were the most sim ilar even though from completely
different hosts. LONG 1, the Holotype, was from Thomomys talpoides
whoreas LONG 2 was from a Lagomorph, Ochotona princeps.
The "H" group (Ileteromyid host group). - This group can be
readily distinguished from the "G" group on the basis of two qualita
tive characterst
Char. No. 2h: Posterior margin of sternal shield moderately
to deeply concave (second pore to anterior to
seta e s t , 2 ); 49
129s Setae av^ and pv^ on tarsus II normal, long
and slender.
The three clusters within the "H" group represent three previously described taxospecies parasitic on rodents of the family Heteromyidae.
They are as follows j
HILL 1-5 = H. hi H i (James on)*
INCO 1-3 = H. incomptis (Eads and Hightower)*
TRIA. 1-3 = K. trlaoanthus (Jameson).
In all the analyses h illi and incomptis consistently clustered
together, with triacanthus joining at a lower level of similarity.
H, triacanthus can be easily recognised on the basis of several
characters i
Char. No. 5^: Feritreme considerably wider anteriorly than
posteriorly*
77-85: Dorsal setae very long and robust.
A single character readily distinguishes incomptis from the other two
taxospecies:
Char. No. 24: Posterior margin of sternal shield deeply
concave (decidedly anterior to setae st. 2 ).
In hilli the following character is discriminatory:
Char. No. 60: Peritremo extends no further than anterior
half of coxa II,
Within the hilli cluster a variant was recognized both on tho
basis of similarity values as illustrated in the three phenograms and
in two qualitative characters. HILL 4 and 5» "the two variant OTU's, 50 were recognized as different from the other h illi replicates on the basis of the following characters»
Char. No. 58i Feritremo only slightly wider posteriorly
than anteriorly|
6 li Posterior half of perltreme normal, not
broken into small segments.
In the Holotype specimen (HILL 1) and the other two replicates (HILL 2
and 3 ) the peritreme is much wider posteriorly than anteriorly, and
the posterior half of the peritreme is distinctly broken into segments.
HILL 1, 2, and 3 were c o lle c te d from th e same h o st sp e c ie s b a t d if f e r e n t
localities (Nevada, Utah, and Idaho). HILL ^ and 5 were from the same
host genus as the others but different species. These two OTU’s were
collected from separate localities (Nevada and Kansas, respectively),
yet they tended to cluster closer to each other than to the other three
HILL OTU's. However, this separation was not so evident in the graphic
analysis as in the phenograms. Based on the morphological and biologi
cal differences noted above, HILL 4 and 5 will be recognized as a
separate taxospecies, designated as H. "C” n. sp. and will be considered
as such in the systematic treatment which follows.
The th re e INCO OTU's c lu s te re d q u ite c lo se to g e th e r even though
they were collected from different localities. INCO 1, the Holotype,
and INCO 2 were from the same host species but separate localities
(Texas and Utah, respectively). INCO 3 was collected from a host of an
entirely different suborder (Feromyscus manlculatus versus Dlpodomys
ordii for INCO 1 and 2). In spite of the differences in biological 51 data, these three replicates of the incomptis taxospecies were quite similar in all other respects.
The three triacanthus replicates clustered even more closely then the INCO OTU's. All were collected from hosts of the same genu 3, and from three adjacent states (Nevada, Utah, and Arizona).
The "I" group (Insectivore host group). - The third major group to be recognized was the "I" group, originally designated as the
Insectivore group. In all analyses the NSPB OTU's and the TALP - PALU
OTU's consistently clustered together, with the CARN 1 OTU joining at a much lower level of sim ilarity. The CARN 1 OTU represents the paloarctic H. carnifex (Koch) taxospecies. Four characters served to distinguish this taxospecies from all Nearctic Kirstionyssus :
Char. No. 16, 17 > D e u to ste m a l d e n tic le s 8 in number arranged
in single filej
71: Dorsal setae <’5 absent;
111: Antero-ventral seta on coxa II stout, spinose,
and hooked;
132 : Seta pv^ on tarsus IV bluntly spiniform.
H, carnifex differs from all other Nearctic members of tho "I" group in an additional four characters:
Char. No. 2h: Posterior margin of sternal shield only
slightly concave (to second pore);
5 0 : Faranal setae distinctly anterior to
middle of anal field; 73f Setae SI present*
131 1 Seta av^ on tarsus IV bluntly spiniform.
The specimen used as the exemplar of carnifex was collected from Talpa europaea in the U.S.S.R., and was identified by N. G. Bregetova who has examined critically the Palearctic species of Hlrstjonyssus occurring on Insectivores (Bregetova, 1956* Evans and T ill, 1966). Based on the above evidence, it has been concluded that carnifex does not occur in the Kearctic region, and that all records from North American insecti- vores and rodents previously published as carnifex are definitely misidentifications.
In the classification herein established the "I" group i s
composed of the NSPB and th e TALP - PALU clusters. This group i s
recognized by the following combination of characters:
Char. No. ?0 j Setae r*f absent j
73s Setae 31 absent*
7^: Setae S3 absent*
^5» ^6: Anal shield very broad relative to length
and with pair of lateral pore3.
The NSPB OTU's represent an undescribed taxospecies designated as H. "B"
n. sp., parasitic primarily on one host species in the family Soriadae
(Blarina brevicauda), although it is recorded from two species of moles
in the family Talpidae. This taxospecies may be distinguished by two
qualitative characters:
Char. No, ^Oj Line formed by end of genital flap ribs
straight or nearly so* 6 8 j Dorsal setae r2 absent.
The three replicates of this taxospecies clustered quite closely in all the analyses, with NSPB 2 and 3 showing the highest sim ilarity. This correlates well with geographic data — the collection localities of
NSPB 2 and 3 were closer than either were to NSPB 1 (Maryland and east
Pennsylvania versus north Ohio),
The TALP - PALU OTU's c o n s is te n tly c lu s te re d in two groups, although TALP and 9 were placed differently in each of the throe phenograms. The clustering in the CRC pehnogram was most similar to that in the graphic analysis. In these two analyses the Nearctic TALP
replicates clustered together and the Palearctic TALP plus the Noarctic
PALU 1 OTU's clustered together. In all cluster analyses tho PALU 1
OTU joined the Palearctic TALP cluster. In tho graphic analysis PALU 1
and TALP 3 had a slightly higher sim ilarity value than PALU 1 and TALP 7,
l^e., the first-order relationship was with TALP 3 and second-order with
TALP 7, Upon examining the qualitative characters, no reliable defini
tive characters could bo found for separating the Nearctic from the
Palearctic TALP OTU's. Slight differences were noted in the shape of
the postorior margin of the sternal shield and tho posterior ond of the
dorsal shield. The Nearctic TALP and PALU 1 were collected from various
species of insectivores of the family Soricidae, wheroas tho Palearctic
TALP were from Talpa europaea in the family Talpidae, However, this
Palearctic taxospecies has been recorded from various host species in
both the families Talpidae and Soricidae. The host and locality
differences between the Palearctic and the Nearctic TALP OTU's would seem to account for the general separation into the two clusters. This probably also accounts for the varied phenogram placement of TALP
This OTU was collected from a "shrew" in Taiwan (Oriental region), which is quite distant from all other TALP OTU's. The very close clustering of TALP 5, 6 , and 7 correlates well with their geographic distribution - the eastorn seaboard. TALP 8 and 9 clustered together in two phenograms and the graphic analysis} they were collected from not too distant localities (Kansas and V7yoming, respectively). After considering the close correlation between the numerical taxonomic analyses and the bio
logical data (especially collection localities), as well as the lack of
reliable discriminatory characters, it is proposed that all the TALP and
the PALU OTU's be relegated to the single taxospecies, H. talpae
Zemskaya,
The Major "S-H" group (Sciurid-Muroid host group). - This major
group which comprised a majority of the OTU's and taxospecies in this
study, can be readily divided into two subgroups on the basis of one
diagnostic characteri
Char. No. 1291 Form of setae av^ and pv^ on tarsus II.
The first subgroup, which will bo referred to as the "S" subgroup
(basic Sciurid host group), corresponds to the phenogram clusters from
AFFI 1 through UTAH 6 . The second which w ill be referred to as the "M"
subgroup (basic Kuroid host group), corresponds to the remainder of the
CTU's in the "S-M" group plus the NSPA cluster. These two subgroups
were also readily recognized in the graphic analysis diagrams. The "S" 55
(AFFI 1 - UTAH 6 ) subgroup can bo distinguished from the "M" subgroup by the following character state i
Char. No. 1291 Setae av^ and pv^ on tarsus II stout and
c la w -lik e .
In the OTU's of the "K" subgroups setae av^ and pv^ on tarsus II are
normal, long and slender.
Tho "S" subgroup (Sciurid host group). - In the three phenograms
and the graphic analysis this subgroup was composed of four definite
clusters, as has already been discussed. In examining tho qualitative
and quantitative characters which might be used for discriminatory pur
poses, no character or combinations of characters could be found which
would consistently and reliably separate these four clusters. The two
clusters composed of OTU's AFFI 1 through ANGU 1 were found to have one
characteristic in commont
Char. No. 131: Seta av^ on tarsus IV acutely to bluntly
sp in ifo rm .
Eut w ith re sp e c t to t h i s c h a ra c te r th e EUTA OTU's and th e UTAH OTU's
wore very nearly the same. Seta av^ on tarsus IV was somewhat enlarged
basally and shortened in some UTAH OTU's, and in the SUTA OTU's this
s e ta was n o t as d e f i n i t e l y sp in ifo rm as in some o th e r members o f th e
cluster to which it belongs. The graphic analysis showed very well tho
relationship between the EUTA OTU's and the UTAH cluster. In consider
ing qualitative characters alone the EUTA and the UTAH CTU's were more
similar to each other than each was to any other. This was also true .for the OTU's of the two clusters CYNO - INVA - ANGU arl LATI, although
to a lesser degree.
As noted in the previous sections, the relationships within the
large cluster AFFI - OCCI - PUNC - EUTA was not satisfactorily clarified
in the phenograms or the graphic analysis. Although, the graphic
analysis yielded somewhat better results than the three phenograms.
Thus in establishing a classification for the taxospecies of the "S"
subgroup the graphic analysis w ill be used as a very general basis,
but the recognition of separate taxospecies will be based more specifi
cally on an analysis of qualitative characters.
The AFFI 1 OTU was the Holotype of a previously described taxo
s p e c ie s , H. a f f i n l s (Jam eson). The o th e r th re e AFFI OTU's were p re v i
ously identified by other workers as affinis, but an examination of
qualitative characters showed them to differ from AFFI 1 in several
characters. The affinis Holotype (AFFI l) differs from AFFI 2, 3« and
4 in the following two characters j
Char. No, 60: Feritrerae extends no further than middle of
coxa I I |
127: With short acute spur on coxa IV.
The a lte r n a tiv e s ta te s o f th e se c h a ra c te rs as found In AFFI 2-4 a re i
Char, No. 60: Peritreme extends beyond middle of coxa II
but usually no further then posterior part
o f coxa I |
127: Without spur on coxa IV. Thus on th e b a s is o f th ese d iffe re n c e s th e AFFI 2-4 OTU's are h e re in recognized as representing a new taxospecies designated as H. "D" n. sp.
V7ith regard to biological data (host and locality), the affinis OTU was collected from Tamiascjarus douglasi in California. The three repli cates of "D" n. sp. were from two different hosts and three separate localities, although two were in the same state. AFFI 2 and 3 were from
the same host (Eutamias minimus) but from separate localities (Utah and
California, respectively). The third individual was from California, but on an altogether different host (Peromyscus boylil). The host data
for these throe individuals correlates quite well with the graphic
a n a ly s is .
The next most c lo s e ly r e la te d OTU's to th e a f f i n i s and "D" n . sp .
taxospecies were the OCCI 1 - PUNC 1-4 OTU's. OCCI 1 represents a
cotype of H. occidentalis (Swing) collected from Tamiasciurus hudsonicus
in Montana. The PUNC 1-4 OTU's are r e p lic a te s o f H. p u n c ta tu s ,
described by Allred and Beck (1966), with PUNC 1 representing the
Holotype, Three out of tho four PUNC OTU's consistently clustered
together in all analyses, whereas FUNC 4 seemed to be most closely
related to AFFI 3 arid OCCI 1. It is interesting to note that the three
PUNC OTU's which c lu s te re d to g e th e r were from th e same h ost (Sutamias_
quadrlvittatus) and locality (Utah), whereas PUNC 4 was from a host of
a different genus (Glaucomys sabrinns) and slightly different locality
in the same state. On the basis of qualitative characters, punctatus
and occidentalis represent the same taxospecies. Thus punctatus is
heroin synonymized with occidentalis. The lack of close clustering 58 between OCCI 1 and the PUNC OTU's In the analyses seems to be a result of the differences in host and locality, as well as perhaps collection date (OCCI 1 collected h 6 years before the PUNC OTU's), H. occidentalis is very closely related to affinis and "D" n. sp,, and shares a number of characters In common. Two very distinctive characters shared by these three taxospecies are:
Char, No. 31^,120: Inner spur on coxae II and III rounded to
almost truncate,
H. occidentalis differs from the other two taxospecies in one character*
Char. No, 131: Seta av^ on tarsus IV bluntly spiniform.
In affinis and H. "D" n. sp, this seta is acutely spiniform, H,
occidentalis may further be distinguished from affinis by:
Char. No, 60: Peritreme extending beyond middle of coxa II
but no further than anterior margin of coxa II,
And from H, "D" n. sp, by:
Char. No. 127: Nith spur on coxa IV,
The latter character is shared in common with affinis.
A fourth very closely related taxospecies, H. eutamiae Allred
and Beck, Is represented by EUTA 1 and 2. EUTA 1 was the Holotype of
eutamiae, with EUTA 2 a replicate from a widely different locality and
different host species. H. eutamiae differs from the above three taxo
species in two characters:
Char, No. H/1,120; Inner spur on coxae II and III small and
blunt, never rounded. 59
On the other hand, it shares one particular character with all three:
Char. No, 601 Feritreme extending to anterior half of
coxa I I .
H. occidentalis and "D" n. sp, are more similar to eutamiae in this character than is affinis. As noted above, the peritreme in affinis extends only to the posterior half of coxa II, H. eutamiae differs
from occidentalis.j i n in a single character:
Char. No. 131: Seta av-j_ on tarsus IV acutely spiniformj and from H. "D" n, sp, in one character:
Char, No. 127: With spur on coxa IV.
These four taxospecies (affinis, "D" n. sp., occidentalis, and euta—
miae), which consistently clustered together in all analyses, were
found to parasitise basically the same group of hosts, Sciurids of
the genus Butamias, although the Holotype of affinis and cotypes of
occidentalis were from species of the genus Tamiasciurus.
As noted in a ll previous analyses, the CYMO - INVA - ANGU cluster
consistently joined the cluster composed of the four above taxospecies,
CYNO 1 represents a paratype of H. cynomys (Radford), INVA 1 and 2
represent the Holotype and a paratype, respectively, of H. invaginatus
Allred and Beck, and A?!GU 1 represents tho Holotype of H. angustus
Allrod and Beck, Based on the close clustering of these OTU's in all
analyses and on an examination of qualitative characters, those four
OTU's are considered to be replicates of the same taxospecies. No
characters could bo found for separating the throe previously estab
lished taxospecies. Thus invaginatus and angustus are heroin synonymized with cynomys. H. cynomys may be recognized by the following diagnostic charactersi
Char. No. Posterior margin of sternal shield deeply
concave (beyond setae st. 2 );
11^,115,120,121: Inner spur on coxae II and III large and
acute t
129: Stout claw-like setae av^ and pv^ on tarsus II
large, long, and distinctly curved;
131: Seta av^ on tarsus IV bluntly spinifonn.
The latter of the above listed characters is shared in general vrith all
the other taxospecies from sclurid hosts, and in particular with occi
dentalis . The principle host of cynomys seemed to be Citellus armatus,
although the original type specimens were collected from a species of
Cynomys in Colorado.
As noted previously, the OTU’s of the TARS - UTAH cluster wore
found to bo very similar to eutamiae. This relationship was illustrated
in the graphic analysis, and supported by an examination of qualitative
characters. Within the TARS - UTAH cluster are eight OTU's representing
two previously described taxospecies, H, tarsalis Allred and Beck and
H. utahensls Allred and Beck. Only one individual (the Holotype) was
included for tarsalis, whoreas utahensis is represented by seven repli
cates, UTAH 1 being the Holotype. On the basis of the close clustering
of TARS 1 with tha UTAH OTU’s, and tho lack of any diagnostic qualita
tive characters, tarsalis is considered to be a synonym of utahensls■ 61
H, utahensls shares several qualitative characters with eutam iae t
Char. No, 24: Posterior margin of sternal shield
moderately concave (second pore to
s t , 2 s e ta e ) |
40: Line formed by end of genital flap ribs
slightly but distinctly arched;
114, 115t Inner spur on coxa TI small and usually
b l u n t .
However, utahensi s differs from eutamiae in the following characters:
Char, No, 60: Feritreme long, extending to or beyond
middle of coxa I;
120, 121: Inner spur on coxa H I large and acute;
131: Seta av^ on tarsus IV slightly enlarged
basally but not distinctly spiniform.
The utahensls replicates Included in this study were from a vride range
of different hosts and localities. The Holotype was from a sciurid
(Eutamias quadrivittatus) in Utah, whoroas a ll other OTU’s except
UTAH 2, wore from species of tha genus Perom.yscus In widely separated
localities in North America (i.e., Oregon, Kexico, Iowa, Ohio),
Collection records indicate that the principle host of this taxospecies
is tho genus Peronyscus, and its distribution is coincident with that
of its principle host genus.
The final cluster in the "3” subgroup to bo considered is tho
LATI cluster. This cluster of five individual OTU's represents a 62 single cosmopolitan taxospecies, H. latiscutatus (de Keillon and
Lavoipierre). In all analyses this cluster joined the utahonsls cluster quite closely. In the graphic diagram it was placed between the UTAH and the ISA3 clusters. An examination of qualitative char acters revealed that latiscutatus is also very similar to cynomys.
Distinctive characters shared in common with cynomys include the
following t
Char. No. 24: Posterior margin of sternal shield
usually deeply concave;
114,115,120,121: Inner spur on coxae II and III largo and
a c u te .
Several characters may be used to distinguish latiscutatus from cynomys:
Char. No. 40: Line formed by end of genital flap ribs
strongly archedj
127: Spur on coxa IV long, slendor and acute>
131: Seta av^ on tarsus IV normal, long and
s le n d e r.
Diagnostic characters separating latiscutatus from utahensis are as
f o llo w s :
Char. No. 24: Posterior margin of sternal shield usually
deeply concave j
40: Line formed by end of genital flap ribs
strongly arched|
114,115,120,121: Inner spur on coxae II and III large and
a c u te ; 127: Spur on coxa IV long, slender, and acute|
1311 Seta av]_ on tarsus IV normal, long and
s le n d e r.
All the latiscutatus replicates were from very closely related hosts of the family Kuridae (Kus muscuius and Rattus spp.). However, collection localities varied greatly — Britain (the one Old Vorld representative),
Kassachusetts, Georgia, Oregon, and Utah, In spite of these great differences in locality the five replicates clustered very close
together in all analyses, LATI 1, from Britain, was usually the most
dissim ilar as would be expected.
The "If' subgroup (luroid host group). - The second subgroup of
the major "S-K" group consists of several rather diverse clusters of
OTU's,Primarily on the basis of the graphic analysis the following
c lu s te r s are in clu d ed : NSPA, TORQ, OBSO, ISAB, BREV, BISE, and NEOT.
A ll OTU's in th o se c lu s te r s have one c h a r a c te r in commont
Char. Ho, 129: Setae av^ and pv]_ on tarsus II normal,
long and slender.
The only other group with which this character was shared is the "H"
group. In all other taxospecies discussed above setae av^ and pv^ on
tarsus II are stout and claw-like.
The clusters comprising this subgroup could not bo oasily sub
divided further, as was the case in the first subgroup discussed above.
Host clusters were too diverse and shared few diagnostic qualitative
characters in common. Char. Ho. 127 (with or without spur on coxa IV)
could be used to separate the clusters into two groups — clusters 6*+
NSPA, TORU, BREV, and NEOT w ith sp u r on coxa IV ; and OBSO, ISA3, and
BISE without spur on coxa IV. In considering overall phenetic simi larity and all other qualitative characters, this was not considered to be advisable. Thus the seven clusters of this subgroup will be discussed in the order listed above, without any further subdivision.
The NSPA OTU's re p re s e n t a r a t h e r a b e r ra n t undescribed ta x o species designated as H, n. sp. It differs from all other members
of this subgroup in the following characters j
Char. No. 2hj Posterior margin of sternal shield deeply
concave;
11*+-116,120-122: Inner spur on coxae II and III absent,
with indistinct, broad, slightly raised
apophysis in place of spursj
123, 127: Outer spur on coxa III and spur on
coxa IV blunt*
also in a number of quantitative characters relating to length of body
setae — all body setae (dorsal and ventral) unusually small. As noted
in the phenograms this cluster usually joined a major stem and failed
to join any other specific cluster. Although in the graphic analysis
the NSPA cluster joined the TORU cluster first, it is still rather
distantly related to TORU. No exclusive qualitative characters shared
by both could be found, although both share several characters related
to the longth of some dorsal setae. This undescribed taxospecies is
known from o n ly th re e female specimens c o lle c te d from Cynomys
ludoviclar.us In Kansas. It was only on the basis of the host from which it was collected that it was originally placed in the Sciurid host group. Fhenetically it is not particularly closely related to any mem bers of that host group.
The TORU cluster of three OTU's represents a previously described
taxospecies, R. torus Allred and Beck. This taxospecies can be recog
nized by the following characters:
Char. No. Hh-116: Inner spur on coxa II very broad and rounded*
131t Seta av^ on tarsus IV acutely spiniformj
77-^5 J Central dorsal setae and setae of Z and S
serios (except Z5 and S5) minute.
As noted in the "OTU Placement in the Phenograms" section, torus con
sistently clustered with the BREV OTU’s, whereas in the graphic analysis
it clustered between cynomys and some ISAB OTU’s, but closer to the
latter. As noted above, Char. No. 127 (spur on coxa IV) is shared with
the BREV cluster, although this same character is also shared with all
except one taxospecies of the "S" subgroup. In addition, torus shares
two other characters variously with some of the Sciurid taxospecies*
Char. No. llh : Inner spur on coxa II large and rounded*
131*. Seta av^ on tarsus IV acutely spiniform.
Thus it is recognized that this taxospecies is also related to members
of the Soiurid host group. In addition to the morphological characters
notod above, this latter relationship is supported by biological data*
H, torus is recorded only from Abert’s squirrel, Sciurus aberti, in
Utah and Colorado. It was interesting to note that three females in
the collection of the late Dr. H, E. Ewing at the U. S. National Kuseum 66 were originally identified by Ewing as Occidentalls. It is suggested that this may well be an example of phenetic divergence from members of the Sciurid host group and convergence toward certain members of the Muroid host group. After considering all these relationships, torus is provisionally placed in the Muroid host group on the basis of overall phenetic sim ilarity and on utilitarian grounds as exemplified by the diagnostic characters noted in the above discussion.
As discussed previously, in all analyses the 03S0 cluster, representing the 77. obsoletus Jameson taxospecies and which was
originally included in the Insectivore host group, consistently
clustered nearer to the ISA3 group than to any other. Although, it was
found to be secondarily closest to the main "I" group. These relation
ships were confirmed in an examination of the qualitative characters
most useful for discriminatory purposes. Characters diagnostic for
obsoletus are as followsi
Char. No, ^+0: Line formed by end of genital flap ribs
strongly arched;
66 : Posterior end of dorsal shield broadly
rounded;
73: Setae Si often absent or off latoral
margin of dorsal shield;
11^, 120: Inner spur on coxae II and III moderate
in size and rounded;
127: W ithout sprur on coxa IV. 6?
Of the characters just enumerated, characters h-0 and 127 are shared in common with the ISAB cluster. Characters llh and 120 in obsoletus are very similar to that observed in some members of the Sciurid host group
(particularly affinis, "D" n, sp., and occidentalis). H. obsoletus was
observed to resemble the members of the "I" group in the general shape
and form of the sternal shield, genital shield, anal shield, and dorsal
shield, but not in the most definitive qualitative characters noted
above. Originally this taxospecies was thought to share one exclusive
character with the "I" group:
Char. No. 73s Setae SI absentj
but later in examining other specimens, not included in the numerical
taxonomic analyses, it was found that setae Si was present on some indi
viduals. In considering biological data, obsoletus has been collected
only on the Pacific Coast (California and Oregon) from several insecti-
vores (Sorex trowbridgli, Scapanus townsendii, and possibly
Ueutrotrichus gibbsii). In the classification herein established,
this taxospecies is provisionally placed in the Muroid host group near
the ISAB cluster on the basis of phenetic resemblance (overall simi
larity) and the qualitative characters noted above. Similar to the
case of t orus, it is suggested that obsoletus probably represents a
case of phenetic divergence from the "I" group and convergence toward
the ISAB cluster.
The most representative OTU's of the "M" subgroup are those of
the ISAB cluster. These 20 OTU's are replicates of the cosmopolitan
taxospecies H, jsabellinus (Oudomans). Within the large ISAT3 cluster, the OTU's variously formed smaller clusters in all the analyses. This taxospecies can be recognized by the following combination of diagnostic charactersi
Char. TTo. ^Oj Line formed by end of genital flap ribs
strongly arched;
11^,115,120,121: Inner spur on coxae II and III largo and
acu te t
127: Without spur on coxa IVt
129t Setae av^ and pv^ on tarsus II normal,
long and slender.
No one unique character could be found for isabelllnus, but the abovo
combination of characters seemed to define the taxospecies very well.
This lack of uniqueness in any character or even a combination of any
two characters possibly serves to explain why so many other taxospecies
cluster around isabelllnus, as illustrated in the graphic analysis
(F ig . 6 ). The inclusion of the 20 replicates of isabelllnus would
also influence clustering by emphasizing the sim ilarity between it and
other taxospecies. The rolntionships between isabellinns and each other
taxospecies clustering near it was or will be noted in the discussion of
each recognized taxospecies, and thus w ill not be repeated here.
As noted previously, the 20 replicates of isabelllnus were
included in this study to give an indication of the effect of different
hosts and localities on intrataxon variability. In comparing the clus
tering of ISAB OTU’s in the phonograms and graphic analysis with the
biological data, there was no indication of correlation with host species, and only very general correlation with locality data. Almost all the OTU's were from Kicrotine rodents with a majority from the genus Hicrotus. In the case of locality data, those individuals collected from the Palearctic region (ISAB 1, 2, 3» a«d *+) consistently clustered together in the three phenograms although the cluster was not exclusive, i.e..» several Pearctic OTU's joined the Palearctic cluster quite closely. The single OTU from the Oriental region (ISAB 5) clustered quite closely with two Pearctic individuals (ISAB 7 and 1^).
There was a very general indication of a trend for the more northern
TTearctic individuals to cluster together. Thus the effect of host and locality on intrataxon variation in this taxospecies seemed to he rather negligible, except in some of the extreme cases of great differences in colloction locality. In referring to the collection records it was found that isabelllnus parasitizes a vory wido spectrum of small mammals throughout the Noarctic, Palearctic, and Oriental regions.
The last three clusters of this subgroup have soveral morpho logical characters in common. This phonetic relationship is also supported by biological data. These three clusters (F3REV, NEOT, and
BISE) represent three previously described taxospecies, H, breviseta
Strandtmann and Morlan, H. neotomae (Eads and Hightower), and H. bjstosus Allred, respectively. In only one of the illustrated pheno grams did the breviseta cluster join tho PEOT - BISE cluster. In the other two phenograms breviseta and torus clusterod together and joined the main "S-M" group stem prior to the joining of tho PEOT - BISE
cluster. It was interesting to note that in the S3D and CBD phenograms (not illustrated) the BREV, NEOT, and BISE clusters joined each other prior to joining any other group, and the TORU cluster was placed elsewhere. In the graphic analysis breviseta had its highest simi larity to isabollinus but was placed between torus and the neotomae - blsetosus cluster. Cn the basis of qualitative characters, breviseta is most similar to neotomae and bisetosus. These three taxospecies have the following characters in common, which are different from other taxospecies in the Kuroid host group.
Char. IIo. 40: Line formed by end of genital flap ribs
straight or nearly so to curved down;
43: Setae Jvl usually on postero-lateral
margin of genital shield;
114,115,120,121: Inner spur on coxae II and III usually
medium to q u ite sm all in s iz e .
Tho principle hosts on which these three taxospecies are parasitic are species of the genus Neotomae.
H, breviseta differs from neotomae and bisetosus in several definitive characters!
Char. No, 77-85* Central setae of dorsal shield decidedly
smaller than marginal setae;
114-116: Inner spur on coxa II absent, with indistinct,
slightly raised apophysis in place of spur;
120,121: Inner spur on coxa III small with an extremely
acute point j
134: Seta pl£ on tibia IV always present. 71
H. breviseta and neotomae share two characters in common 1
Char. No, 40: Line formed by end of genital flap ribs
straight to slightly arched*
127: With spur on coxa IV.
In neotomae the line formed by the end of the genital flap ribs is never arched but is curved down, and no spur is present on coxa IV.
H. neotomae and bisetosus share the following characters which are different from breviseta:
Char. No. 77-85: All dorsal setae subequal in length*
114,115,120,121: Inner spur on coxae II and III small to
moderate in size and acute to blunt*
134: Seta pl^ on tibia XV absent.
H. bisetosus differs from neotomae in only two distinctive characters:
Char. No. 40: Line formed by end of genital flap ribs
curved down*
127: Without spur on coxa IV.
The three breviseta replicates clustered quite close together in all
analyses, HR1V 1, tho Holotype, and BR'SV 2 were from the same host
species but slightly different localities (New Mexico and west Texas,
respectively). BR/2V 3 was taken from a colony reared from an i n d i v i d u a l
female originally collected in northern California.
Tho neotomae OTU's were c o lle c te d from fo u r d i f f e r e n t h o st
species and localities. In spite of those differences all four OTU’s
clustered quite close in the phenograms and graphic analysis. UEOT 2
was tho most dissimilar, possibly a reflection of the difference in 72 host as well as locality — NEOT 2 was collected In Kansas from a rodent
(Cynomys ludoviclanus) of an entirely different suborder. The other three (NEOT 1 (the Holotype), NEOT 3» and NEOT *0 were collocted from rather widely separated localities (Texas, Utah, and California, respec tively) but hosts of the same subfamily (NEOT 1 and b from Neotoma, and
NEOT 2 from Peromyscus).
The three replicates of bisetosus were collected from the same h o st sp e c ie s. BISE 1, tho H olotype, and BISE 2 were from th e same locality in Utah, while BISE 3 was collected in southeast Idaho. This basically very similar biological data is well correlated with the
clustering in the numerical taxonomic analyses. All three joined each
other at a very high level of similarity.
Groups and Taxospecies Formally Recognized
As a result of the present study, the following groups and taxo
species are formally recognised.
Genus Hirstionyssus Fonseca, 19^8
Subgenus Hirstionyssus str,
Geomyid host group
1. Hirstionyssus (H.) femuralis Allred, 1957
2. Hirstionyssus (H.) geomydis (Keegan, 19^6)
3. Hirstionyssus (H.) thomomys Allred and Beck, 1966
h--. Hirstionyssus (H.) longicholae Allred and Beck, 1966
Heteromyid host group
1. Hirstionyssus (H.) h llli (Jameson, 1950)
2. Hirstionyssus (H.) "C" n. sp. 73
3. Hirstionyssus (H.) incomptis (Eads and Hightower, 1932)
h. Hirstionyssus (H.) trjaoanthus (Jameson, 1950)
Insectivore host group
1. Hirstionyssus (H.) "3" n. sp.
2. Hirstionyssus (H.) talpae Zemskaya, 1955
Sciurid - totroid host group
Basic Sciurid. host group
1. H irstio n y ssu s (H. ) a f f i n i s (Jameson, 1950)
2. Hirstionyssus (H.) "_D" n . sp.
3. Hirstionyssus (JI,) occldentalis (Ewing, 1923)
h. Hirstionyssus (H.) eutamiae Allred and Eeck, 1966
5. Hirstionyssus (H. ) cynomys (Radford, 194-1)
6. Hirstionyssus (H.) utahensis Allred and Beck, 1966
?• Hirstionyssus (H.) latiscutatus (do I'eillon and
L a v o ip ie rre , 1944-)
Basic Kuroid host group
1. Hirstionyssus (H.) "A" n, sp .
2. Hirstionyssus (H,) torus illred and Bock, 1966
3. Hirstionyssus (H.) obsoletus Jameson, 1950
4. Hirstionyssus (H, ) Isabellanus (Oudemans, 1913)
5. Hirstionyssus (H.) breviseta Strandtmann and Morlan,
1953
6. Hirstionyssus (H.) neotomae (Eads and Hightower, 1951)
7. Hirstionyssus (H.) bisetosus Allred, 1957 Subgenus "X" us, new subgenus
Carnivore host group
1. Hirstionyssus ("X") staffordi Strandtmann and Hunt, SYSTEMATICS OF THE GENUS HIRSTIONYSSUS
Family Laelapidae Barlese, 1892
Subfamily Hirstionyssinae Evans and T ill, 1966
Tho classification of the family level used herein is that of
Radovsky (1967, 1969), Radovsky (1967) recognized seven subfamilies
in the family Laelapidae — Laelaplnae Berlose, 1892 1 Hyposaspidinae
Vitzthum, 19^0; Haemogamasinae Oudemans, 1926; Kyonyssinae Bregetova,
1956l Alphalaelapinae Tipton, I960; R aillietiinae Vitzthum, 19^2; and
Iphiopsinae Kramer, 1886, Tho genus Hirstionyssus Fonseca, 19^8, and
four related genera (Echinonyssus H irst, 1925, Patrinyssus Jameson,
1950! Australolae laps ’Nomersley, 1956; and Trichosurolaolaps Womersley,
1956) were provisionally placed in the subfamily Hyonyssinae along with
the genus Fyonyssus Tiraboschi, 1903. Just prior to this proposal,
Evans and T ill (1966) erected a separate subfamily, Hirstionyssinae,
for the genus Hirstionyssus and the four related genera. This was sub
sequently recognized by Radovsky (1969) as an appropriate arrangement.
Thus at present tho family Laelapidae contains eight subfamilies,
Tho classification and diagnosis of the subfamily Hirstionyssinae
are based on Evans and T ill ( 1966 ), Radovsky (1967, 1969), and Evans and
Fain (1968), Six genera of obligatory, small mammal parasites are pres
ently included in this subfamilyt (1) Hirstionyssus Fonseca, 19^-8,
75 containing an undetermined number of species parasitic on small mammals worldwide! (2) Echinonyssus Hirst, 1925» with one species, E. nasutus, parasitic on Malaysian primates of the genus Tupaia; (3) Fatrinyssus
Jameson, 1950* containing ono species, P. hubbardi, parasitic on the mountain beaver, Aplodontia rufa, in the pacific northwest of the
United States| (4) Triehosurolaelaps Komersley, 1956, with seven species parasitic on marsupials in tho Australian region| (5) Aus- tralolaelaps ’.v’oraersley, 1956 , with three species also from marsupials in the Australian region; and ( 6 ) Ancoranyssus Evans and Fain, 1968, represented by one species, A. trichys, collected from an old world porcupine, Trichys lipura, in Baram-Sarawak.
Diagnosis based on Adults
GNATH0S0KA. Chelicera of female slender, rarely fused
proximally (Ancoranyssus), and with edentate chelae j
moveable digit with pointed tip and convex sides; fixed
digit with hyaline process. Chelicera of male slender,
Chelao edentate and suboqual in length; moveable digit
usually simple and without dorsal arm separate from
spormatodactyle. Small pilus dentilAs present in some
females; dorsal seta arising near base of chelae reduced
to microseta or absent; no distinct arthrodial processes
present. Internal malae of hypostome long, slender,
horn-like; com iculi membranous, rarely with horn-liko
apex; tectum capituli tonguo-liko with anterior margin
smooth or fimbriated. Palpal trochanter rarely with ventral process (Trichosurolaelaps); palpal apotele two-
tined or absent (Patrinyssus). Deutosternal denticles in five to 18 transverse rows with one to four denticles
per row. Venter of gnathosoma with normal chaetotaxy
(one pair capitular setae and three pairs hypostomal
s e t a e ) .
VENTER, Tritostemura normal, bipartite, laciniae pilose
or smooth, rarely with armed tritostem al base (Tricho-
surolaelaps). Sternal shield in female with two pairs
of pores and usually with three pairs of setae, rarely
w ith s t . 3 setae off shield (Hirstionyssus); metastornal
setae free. Genital shield broad with broadly rounded
tip, normally bearing only one pair of genital setae;
sometimes with Jvl setae (Hirstionyssus) or Jvl and Jv2
setae (Australolaelaps) on postero-lateral margin. Anal
shield variable from pear-shape to oblong-oval; opistho—
gastric area with variable number of setae, I'etapodal
shields weakly developed or absent. Feritrematal shields
free or fused posteriorly with podal shields; podal shields
poorly developed posterior to coxa IV, Peritrome variable
from well developed to reduced. Kolovontral shield of male
rarely with division or weaker sclerotized region between
ventral and anal portions, not fused vrith peritrem atal
shields; genital orifice prestem al.
DOR.SUH, Dorsal shield entire in both sexes, rarely restricted to anterior half of idiosoma (Ancoranyssus),
and rarely with anterior vertex extending out over
gnathosoma (Echinonyssus). Dorsal shield exhibiting
hypotrichy, bearing 19 t o 27 pairs setae in femalej
setae zl rarely present, j series showing greatest
hypotrichy. Relative lengths of lateral and marginal
setae greater in males than in females.
LEGS. Chaetotaxy of legs normal or with deficiencies
in chaetotaxy of genua and tibiae III and IV. Certain
coxal setae and ventral setae of femur, genu, and tibia
sometimes hypertrophied to form spinose setae, peg-like
setae, claw-like setae, or retrograde spurs. One or
more coxae with distinct, ventral, nonsetigorous spur
or spurs, usually posteriorly directed, besides tho
anterior spine of coxae IIj anterior spine of coxae II
may be well developed, greatly enlarged and hook-like
(Echinonyssus), or absent (Patrinyssus). All ambulacra
with pair of well developed claws.
Genus Hirstionyssus Fonseca, 19^8
(Figs. 7-11)
Ichoronyssus of authors, in part
Llponyssus of authors, in part
Hirstionyssus Fonseca, 19h8, Proc, Zool, Soc. London 118(2):266, 29^.
Type by original designation, Dermanyssus arcuatus, Fonseca, 19^8, sonsu Oudemans, 1913 (not Koch, 1839 ) t
£= Hirstionyssus talpae Zemskaya, 1955» (In Bregetova
et a l., Pored. Faune SSSR 59s360)J, synonymy by Evans
and T ill, 1966, Bull. Brit. Kus. (Natur. Hist.) Zool.
l^(5):277-278.
Neoichoronyssus Fonseca, 19^1, Cienc. 2:263. Type by original
designation and monotypy, Liponissus wernockl Fonseca,
1935* (Jameson, 1950, Proc. Ent. Soc. Washington
52(^):l62, makes Hirstionyssus Fonseca, 19^-8, a sub
genus of Neoichoronyssus Fonseca, 19^1, but this arrange
ment has not been recognized by other authors).
Echinonyssus H irst, 1925r Proc. Zool. Soc, London 1925:51* Type,
by original designation and monotypy, Echinonyssus nasutus
H irst, 1925. (Domrow, 1963, 1965» Proc. Linn, Soc. New
South Wales 88(2):200, and 90(2):l 6 ^, suggests that
Hirstionyssus Fonseca, 19^-8, is a synonym of Echinonyssus
Hirst, 1925, but this is not generally accepted as valid
at the presont time).
Diagnosis based on Adults
Medium sclorotizod nites ranging from iJ-00 to 950 microns
i n le n g th .
GNATHOSOMA. Chelicera with edentate chelae. In females
both choliceral digits dorso-ventrally flattened and
grooved along cutting facesj fixed digit with membranous
sheath. Moveable digit of males completely fused with deeply grooved spermadactylo. Pilus dentilis absent* dorsal seta reduced to microseta or absent. Internal taalao of hypo stoma simple, variable in length* com iculi membraneous| tectum capituli elongated, tongue-shaped, with anterior margin smooth or fim briated. Palpal trochanter without ventral process* palpal apotele two- tlned, Deutosternal denticles in single file or alternating double file rarely regular double file. Venter of gnathosoma with normal chaetotaxy,
VENTER. Tritostemum normal, bipartite, laciniae pilose, tritosternal base never armed. Sternal shield in female with two pairs of pores and usually with throe pairs of
setae, st. 3 rarely off shield* shield usually with antero lateral and postero-lateral projections* metastemal setae
free. Genital shield broad with broadly rounded tip, with
out sculpturing, with more heavily sclerotized medial strip
extending to tip of plate and lightly sclerotized lateral
regions* normally with one pair of genital setae on shield,
rarely with Jvl setae on postero-lateral edge. Anal shield
roughly poar-shapad to oblong-oval, with normal chaetotaxy*
position of paranal setae relative to anal field variable,
Opisthogastric area with variable number of setae, Holo-
vontral shield of male normally bearing eight pairs of setae,
excluding anal setae, rarely with nine or 10 p a i r s ,
DORSUM. Dorsal shield entire, bearing 23 to 2? pairs of usually simple setae in female, sometimes setae on posterior margin slightly barbed| setae zl never present) centrally located setae usually shorter than more laterally located setae, except for jl setae which are always small.
LEGS. Chaetotaxy of legs normal or with deficiencies in
chaetotaxy of genua and tibiae III and IV. Certain coxal
setae and ventral setae of femur, germ, and tibia sometimes
hypertrophied to form spinose setae, peg-like setae, or
claw-like setae. One or more coxae with distinct, ventral,
non-setigerous spur or spurs, usually posteriorly directed,
besides tho anterior spine of coxa II) anterior spine of
coxa II well developed.
Key to Females of the Genus Hirstionyssus
Sternal shield small and without postero-lateral projections
bearing st. 3 setae, i.e., st. 3 setae off shield on soft
integument. Paranal setae at anterior end of anal field.
Peritreme extending only to antorior margin of coxa III.
Posterior end of dorsal shield forming nari'ow, caudal pro
jection) dorsal setae z 6 and J2 absent, Coxal spur formula
1—3-2—1j with inner spur on coxae T, I I , I I I , and IV u s u a ll y
bifid and blunt. Seta pdg on genu IV absent, (Figs, 81-83)»
(Subgenus ”X" us, new subgenus). .... Jl. staffordl (p. l 6 h )
Sternal shield large and with postero-lateral projections
bearing st. 3 setae; st, 3 setae never off shield on soft 82
integument. Position of paranal setae variable but not
at anterior end of anal field. Peritreme extending ■
beyond anterior margin of coxa III, Posterior end of
dorsal shield variable in shape but never forming narrow,
caudal projectiont dorsal setae z6 and J2 present, Coxal
spur formula variable but never 1- 3- 2- 1 ; i n n e r s p u r o f
coxae never bifid. Seta pd^ on genu IV present,
(Subgenus Hirstionyssus js. s t r . ) ...... 2
2(1). With postero-lateral spur on coxa II; without spur on
coxa IV. i • 3
- Without postero-lateral spur on coxa II ; with or without
spur on coxa IV...... 10
3(2). Posterior margin of sternal shield slightly convex to
slightly concave (to 2nd pore). Setae av^ and pv^ on
tarsus II stout and claw-like. (Geomyid host group). . . . *4-
- Posterior margin of sternal shield moderately to deeply
concave (2nd pore to anterior to setae st, 2). Setae
av^ and pv^ on tarsus II normal, long and slender.
(Heteromyid host group)...... 7
*K3). Postorior margin of sternal shield distinctly convex.
Innor spur on coxa II short and rounded; inner spur
on coxa III very large and rounded. Postorior margin
of dorsal shield in shape of broad, blunt wedge
(Figs. 22-2*0, ...... H. lo n g ic h o la e (p . 99 ) 83
- Posterior margin of sternal shield straight to slightly
concave (to 2nd pore). Inner spur on coxae I I and I I I
moderate in size and acute to blunt. Posterior margin
of dorsal shield in shape of narrow, sharp vodge, . . . . . 5
5(4). With the normal 26 pairs of dorsal shield setae plu 3 th e
five r series setae ; all dorsal setae large, subequal
in length with j 2 , j 3 , z2 , z 4 , s 3 » and s4 setae slightly
longer. (Figs. 19-21) ...... H, thomorays (p. 97)
- Without the normal 26 pairs of dorsal shield setae,
setae Si absent and with setae Zl moved closer to
margin of dorsal shield; dorsal setae not all large,
not subequal in length...... 6
6(5). 'With a blunt apophysis on femur II; setae av^ and av-j on
tarsus II decidedly bulbous basaUy. Dorsal setae r2,
r3, and r4 normally developed; setae j4, j5, j 6 , z5» Jl ,
J2, and J3 very small; setae j2, z2, and si more than
twice as long. (Figs, 12-15) ...... H. femuralis (p. 92)
- Without blunt apophysis on femur II; setae av 2 an d av^
normal. Dorsal setae r2, r3> and r4 not normally
developed but with normal sotal bases; all dorsal shield
setae small, suboqual in length except setae jl, j 2 , j 3 *
z2, z4, and si which are somewhat longer. (Figs. 16-18)
...... H. geom ydis ( p . 95 )
7(3). Peritreme considerably wider anteriorly than posteriorly.
All dorsal sotao very robust and long, length oqual to 84
or greater than distance between adjacent setae.
(Figs. 31-32). .•■«• ...... H. triacanthus (p. 112)
- Peritreme definitely narrower anteriorly than posteriorly.
Dorsal setae not unusually robust and long ...... 8
8(7). Posterior margin of sternal shield deeply concave
(distinctly anterior to st. 2 setae). Peritreme
extends to middle or anterior margin of coxa I.
(Figs. 29-30)* ...... H* incomptjs (p. IO7 )
- Postorior margin of sternal shield modorately concave
(2nd pore to level of st. 2 setae). Peritreme extends
only to anterior margin of coxa I I ...... 9
9(8). Peritreme considerably wider posteriorly than antoriorlyi
posterior half distinctly divided and appears broken up.
(Figs. 25-26) ...... n, hilli (p. 102)
- Peritreme subequal in width throughout entire lengthj
posterior half normal, not divided or appearing to be
broken up. (Figs, 27-28) ...... H. "C" n . s p . (p . 105)
10(2), Dorsal setae r4, SI, and S3 absent. Coxal spur formula
0- 2- 2- 1 j with all spurs moderate in size and acute.
Setae av^ and pv^ on tarsus II stout and claw-like.
(Insectivore host group). 11
- Dorsal setae r4, SI and S3 present (except Si in some
individuals of H, obsoletus). Coxal spur formula
variable, as well as size and shape of spurs. Setae 85
av^ and pv^ on tarsus II variable. (Kajor Sciurid-
Muroid host group)...... 12
11(10). Dorsal setae r2 absent, Antero-ventral seta on coxa II
normal, long and slender. Line formed by end of genital
flap ribs straight or nearly so. (Figs. 33**35)
...... H. "BH n. sp. (p. 117)
- Dorsal setae r2 present. Antero-ventral seta on coxa II
stout, blunt, and peg-like. Line formed by end of
genital flap ribs moderately arched. (Figs. 38-38)
...... H. talpae (p. 120)
12(10). Setae av^ and pv^ on tarsus II stout and claw-like.
(Sciurid host group)...... 13
- Setae av^ and pv^ on tarsus II normal, long and slender.
(Kuroid host group) ...... 19
13(12). Inner spur on coxae II and III moderate in size and
rounded to almost truncate, ...... l*f
- Inner spur on coxae II and III small to large in size
and acute to slightly blunt ...... 16
1^(13). Peritreme short, oxtonding only to middle of coxa II.
V.ith small, sometimes acutely forked, spur on Coxa IV,
(Figs. 39-^2) ...... H. affinis (p. 12^)
- Feritreme extending beyond middle of coxa II but no
further than postorior third of coxa I. With or
without spur on coxa IV, ...... 15 86
15(1*0. ’Without spur on coxa IV. Seta av^ on tarsus IV acutely
spiniform. Line formed by end of genital flap ribs
straight or nearly so. (Figs. 43-** 6 ). .H, "DM n. sp. (p. 125)
- With moderate size, acute spur on coxa IV. Seta av^ on
tarsus IV bluntly spiniform. Line formed by end of
genital flap ribs slightly but distinctly arched.
(Figs. 47-50) ...... ••••••* H, occidantalis (p. 128)
16(13). Inner spur on coxa II small and usually blunt, Postorior
margin of sternal shield moderately concave (never beyond
setae st, 2). Line formed by end of genital flap ribs
slightly but distinctly arched 17
- Inner spur on coxa II large and acute. Posterior margin
of sternal shield moderately to deeply concave (usually
to or beyond setae st. 2). Lino formed by end of genital
flap rib3 slightly to strongly arched...... *• 18
17(16), Peritreme extending only to anterior margin of coxa II.
Inner spur on coxae II and III small and blunt; spur
on coxa IV very small and acute. Seta av-^ on tarsus IV
acutely spiniform. Distance between setae Zl and Si
slightly greater than length of setae Zl. (Figs, 51“54)
H, eutamiae (p. 13l )
- Peritreme extending to or beyond middle of coxa I. Inner
spur on coxa III larger and more acute than spur on
coxa II; spur on coxa IV short, broad, and acute. Seta
av^ on tarsus IV slightly enlarged basally but not 87
acutely spiniform. Distance between setae Zl and SI
much greater than length of* setae Zl. (Figs, 59-62)
...... H, utahensls (p, 136)
1 8 (1 6 ), Spur on coxa IV short, broad and acute. Stout claw-like
setae av-^ and pv^ on tarsus II long and curved. Seta
av^ on tarsus IV bluntly spiniform. Line formed by end
of genital flap ribs slightly but distinctly arched.
Posterior end of dorsal shield in shape of narrow, sharp
wedge. (Figs. 55-58)...... H. cynornys (p. 1 3 3 )
- Spur on coxa IV long, slender, and acute. Stout claw-like
setae av-^ and pv-^ on tarsus II short and straight. Seta
av^ on tarsus IV normal, long and slonder, line formed
by end of genital flap ribs strongly arched. Posterior
end of dorsal shield in shape of broad, blunt wedgo.
(F ig s . 63 - 65 ) ...... H.latiscutatus (p. 1^2)
19(12). Inner spur on coxae II and III absent) with indistinct,
broad, slightly raised apophysis in place of spur) outer
spur on coxa III and spur on coxa IV usually blunt.
Posterior margin of sternal shield deeply concave (beyond
st. 2 setae). All body setae (dorsally and ventrally)
unusually small, (Figs. 66 - 67 ). . . . H. "A" n. sp. (p. 11±6)
- Inner spur on coxae II and III prosent and well developed
(except spur on coxa II of H. brevisota)) outer spur on
coxa III and spur on coxa IV (when prcsont) acute.
Posterior margin of sternal shield slightly t o 88
moderately concave. All body setae not unusually
small...... 20
20(19). Inner spur on coxa II very broad and roundedj with spur
on coxa IV. Seta av^ on tarsus IV acutely spiniform.
Central dorsal setae and setae of Z and S series
(except Z5 and S5) minute, (Figs, 68-70). . H. torus (p. 1^7)
- Inner spur on coxa II variable but not very broad and
rounded; with or without spur on coxa IV. Seta av^ on
tarsus IV normal, long and slender. Central dorsal
setae and setae of Z and S series not a ll minute, .... 21
21(20). Inner spur on coxae II and III moderate in size and
rounded; without spur on coxa IV. Anal shield very
broad relative to length; with pair of lateral pores.
Posterior end of dorsal shield broadly rounded; setae
SI often absent or off lateral margin of dorsal shield.
(Figs. 71-72) H. obsoletus (p. Ih9)
- Inner spur on coxa II variable; inner spur on coxa III
always acute; with or without spur on coxa IV. Anal
shield not particularly broad; usually without lateral
pores. Posterior end of dorsal shield usually in shape
of broad, blunt or narrow, sharp wedge; setae SI always
p r e s e n t ...... 22
22(21), Inner spur on coxae II and III large and acute; without
spur on coxa IV, Genital setae never on postero-latoral
margin of genital shield, line formed by end of genital 89
flap ribs strongly arched* Figs. ( 7 3 - 7 * 0
...... H. isaballlnus (p. 15l)
- Inner spur on coxae II and III not large, usually small
to moderate in size and acute to blunt (except spur on
coxa II of H. breviseta); with or without spur on coxa IV.
Genital setae usually on postero-lateral margin of genital
shield. Line formed by end of genital flap ribs not-
strongly arched. 23
23(22). Inner spur on coxa II absent; with Indistinct, slightly
raised apophysis in place of spur; innor spur on coxa III
small and very acutely pointed; with spur on coxa IV.
Seta pl£ on tibia IV always present. Central dorsal
setae smaller than marginal setae, (Figs. 75-76)
H. broviseta (p. 157)
- Inner spur on coxae II and III small to moderate in
size and acute to blunt; with or without spur on
coxa IV. Seta pl^ on tibia IV absent. All dorsal
setae subequal in length...... 2*4
2*4(23). *l'*ith spur on coxa IV. Line formed by end of genital
flap ribs straight or nearly so. (Figs. 77~?8)
...... H. neotom ao ( p . 1 5 8 )
- VJithout spur on coxa IV. Line formed by end of gonital
flap ribs distinctly curved down. (Figs. 79-80)
H, bisetosus (p. 162) 90
Recognized Taxospecies of the Genus Hlrstionyssus
As a result of the proceeding numerical taxonomic analysis 25 taxospecies in two subgenora are formally recognized. In the follow ing section of this paper the female of each taxospecies is given a complete taxonomic treatment. A synonymy is given first, followed by the taxospecies diagnosis. Reference is made to figures which illus trate the diagnostic characters given for each taxospecies. Following the diagnosis are collection records which include the Noarctic records
of all specimens examined in this study plus a ll Nearctic records
reported in the literature and hero accepted as valid. For each taxo-
spacies the type record is indicated by an asterisk before the host and
locality. References are cited for all literature records, and the
source of the specimens examined is indicated in parenthesis following
the number of females examined. Following is a lis t of the abbrevia
tions used in referring to the source of specimens*
BMTIH B ritish Museum (Natural H istory), Cromwell Road, London, S.W. ?, England
CDR C. D, Radford, "Lampagoos", 85, Grosvenor Avenue, Torquay, Devon, England
DMA D, M. Allred, Department of Zoology and Entomology, Brigham Young University, Provo, Utah 84601
EWJ E. W. Jameson, Jr., Department of Zoology, University of California, Davis, California 95616
FJR F. J. Radovsky, George W. Hooper Foundation, Univorsity of California Medical Center, San Francisco, California 94122
GWK G. W, Krantz, Department of Entomology, Oregon State University, Corvallis, Oregon 9?33* OSU Ohio Stato University, Acarology Laboratory, College of Biological Sciences, Columbus, Ohio ^+3210
RWS R, W, Strandtmann, Bernice P. Bishop Museum, Honolulu, Hawaii 96819
USNM United States National Museum, Washington, D, C. 20250
The subgenora, major host groups, and taxospecies are arranged in the same order in which they were presented in the "Groups Formally
Recognised" section of the "Discussion*1*
Subgenus Hlrstionyssus s_. str.
D ia g n o sis
Deutosternal denticles 12 to 30 in numbers arranged basically
in alternating double file. Sternal shiold large and/or with
postero-lateral projections bearing st. 3 setae. Faranal
setae positioned from slightly anterior to slightly posterior
to level of middle of anal field, Feritrome not extroraely
shortj extending beyond posterior margin of coxa I I . Shape
of dorsal shield variable; posterior ond broadly rounded to
narrow, sharp, wedge shaped but never forming narrow, caudal
projection, Dorsal sotao z6 an d J2 presentj setae of R series
always on soft integument and never on dorsal shield margin.
Spur never prosent on coxa I | inner spur on coxae I - IV
variable in size, and shape, but never bluntly bifid; spur on
co x a IV, when present, always on latoro-ventral margin and never in same position as inner spur on coxae II and III.
Seta pd^ of genu IV present.
Hirstionyssus (H,) ferauralis Allred, 1957
(Figs. 12-15)
Hirstionyssus sp,, Allred and Roscoo, 1957» Trans, Amer, Kicroscop,
S o c, 7 6 (*O s396 (host, distr,)* Howell, Allred, and Beck, 1957»
Ecol. 38(1):83 (host, d istr.),
Hirstionyssus fem uralis Allred 1957» Amer, Midland Hatur. 57(2) **45**
(figs., host, d istr.) [Typoj Rockville, Washington Co., Utah*
Holotype, Dept, of Zool, and Entomol., Brigham Young Univ.,
Provo, Utah] * Allred and Beck, 1966, Brigham Young Univ. Sci.
Bull,, Biol. Ser, 8(1):25 (key, figs., host, distr.).
D ia g n o sis
Deutostemal denticles 12 - 13 in number* arranged in alternating
double file. Anterior margin of sternal shield slightly concave*
posterior margin slightly concave, usually no further than level
of setae st, 3» Line formed by end of genital flap ribs irregu
larly straight to slightly arched. Setae Jvl off genital shield
on soft integumont, but sometimes touching margin of shield.
Paranal setae positioned at lovol near middle of or slightly
anterior to middle of anal field. Poritreme subequal in width,
usually slightly wider posteriorly* extending to near middle of
coxa I* normal in appearance throughout. Greatest width of
dorsal shield always at level of s^ setae* lateral sides straight or nearly so, and converging toward posterior end|
posterior end in shape of narrow, sharp wedge. Setae SI
absent, with setae Zl moved close to margin of dorsal shield.
All other dorsal setae present and normally developed.
Setae Z5» S5» and sometimes Sk slightly barbed. Setae
j5 i z6 , J l , J2, and J3 very small, with setae j 2 , z 2 ,
and si more than twice as long; all other setae intermediate
in length. Coxal spur formula 0 -3 -2 -0 ; inner spur on coxa II
large and usually blunt; postero-lateral spur on coxa II
blunt to rounded; inner spur on coxa I I I large and acute to
blunt; outer spur on coxa I I I small and acute. Blunt
apophysis postero-ventrally on femur I I , S e ta e av-^ an d pv-^
on t a r s u s I I stout and claw -like; sotae avg and av^ on tarsu 3
II very long and slender but decidedly bulbous basally. All
other leg setae presont and normal.
Collection Records
Cltollus armatus (Kcnnicott)
UTAH: Salt Lake Co., Ft, Douglas; Coll: V. Bullough; 7 April
19^9; 3 fem ales (DMA), ( A llre d and Beck, 1 9 6 6 ),
Thomomys bottae (Eydoux and Gervais)
CALIFORNIA: C o lo rad o D e s e rt; C o ll: H, S , G e n try ; 1? F e b ru a ry
193^1 1 female (RWS)j Alameda Co., Berkley; Coll: J, B, Boxer;
June, 15 May 19^7J 5 females (RWS); San Mateo Co., M illbrao;
Coll: R. P. Maynard; 15 May 1953; 1 female (RWS). 94
TEXAS i Presidio Co.; Colli J. L. Regan; 14, 27 June 1948;
13 females (RWS),
UTAHt Juab Co.; Kane Co., Kanab; Colli D £, Beck; 22 April
1951I 7 fe m a le s (DMA), (A llr e d and B eck, I 966 ); Millard Co,,
Lyndyll; Colli D, K, Allrod and J, LaKunyonj 1 November 19^9 1
2 females (RWS); San Juan Co., Bluff; Colli D, K. Allred;
4 Kay 1951; 1 female (DKA); ’Washington Co., Rockville; Colli
D, M. Allred; 18 December 1950; 1 female (DM), (Allred and
B eck, I 966 ),
Thomomys talpoides (Richardson)
UTAH; Juab Co., Mt. Nebo; Colli D, N, Allred; 24 September 1952;
1 fe m ale (DMA); San P e te C o ,; (A llre d and B eck, 1 9 6 6 ),
Thomomys sp ,
CALIFORNIA: San Mateo Co,, M llbrao; Coll: R. P, Maynard;
29 January, 15, 19 February 1949; 4 females (RWS).
NEW MEXICO: Lincoln Co.; Coll: W. F. Blair; 19 July 1947;
2 females (RWS),
UTAH: Washington Co,, Rockville; Colli D. K. Allred; 18 December
1950; 2 females (DMA), (Allred and Bock, 1966).
Cratogeomys castanops (Baird)
TEXAS: Presidio Co.; Coll: J, L, Regan; 6 , 12, 13 Juno, 1 July
1948; 11 females (RWS); same locality; Coll: S. Await; 14 June
1948, 6 females (RWS); same locality; Coll: W. F, Blair; 24 July
1947; 4 females (RWS); Terrell Co.; Colli B, G, Hightower;
10 June 1949; 2 females (RWS), 95
Cratogeomys pynmurus im parllis (Goldman)
MEXICO: Michoacan, II./ Patzcuaro; Coll: D. P. Furman; 7 November
1953! 3 females (RWS),
*Peromyscus eremicus (Baird)
*UTAH: ‘Washington Co., Rockville; Coll: D, M, Allred; 18 December
1950; 1 female (Holotype) (DMA), (Allred, 1957b),
Neotoma le p id a le p ld a Thomas
UTAH: Juab Co.j (Howell, Allred, and Beck, 1957); Tooele Co,;
(Allred and Roscoe, 1957)»
Hirstionyssus (H.) goomydis (Keegan, 19^6)
(Figs. 16-18)
Liponyssus goomydis Keegan, 19^6, Trans. Amer. Microscop. Soc. 65(l)t
69-71 [Type: Fort Leavenworth, Leavenworth Co., Kan,; Holotype,
U. S. N, M,] ,
Hirstionyssus geomydis, Fonseca, 19^8, Proc. Zool, Soc. London
118(2):297 (synon.); Jamoson, 1950, Froc, Biol. Soc. Washington
63:31-32 (key); Strandtmann and Worlan, 1953» Toxasjtop. Biol,
Med. 11 (*0:629 (key, fig s.); Strandtmann, 1956, J. Kansas
Sntomol. Soc. 29(h0):138 (descr, note); Strandtmann and Wharton,
1958, Inst. Acarology Contrib. *l-:103 (synon,, host, d istr.),
Neoichoronyssus geomydis, Jameson, 1950, Proc. Entomol. Soc. V/ashington
52(iv): 162-163 (key). D ia g n o sis
Doutostemal denticles 15 - 16 in number* arranged in
alternating double file. Anterior margin of sternal shield
straight to slightly convex* posterior margin irregularly
straight to slightly concave, usually no further than setae
st. 3. Line formed by end of genital flap ribs irregularly
straight. Setae Jvl off genital shield on soft integument,
but sometimes touching margin of shield. Paranal setae
positioned at level near middle of anal field. Feritrema
suboqual in width, usually slightly wider posteriorly*
extending to anterior half of coxa I* normal in appoarance
throughout. Greatest width of dorsal shield usually at level
o f s e ta e s 6 - Zl* lateral sides straight to slightly concave
and nearly parallel* posterior end in shape of narrow, sharp
wedge. Dorsal setae r2, r3» and r4 not normally developed*
with normal setal bases but with very small hyalin club-
shaped setae. Setae SI absent, with setae Zl positioned
somewhat closer to margin of dorsal shield than normal.
All other dorsal setae present and normally developed. Setae
Z5, S5, and SA- slightly barbed. All dorsal shield setae small
subequal in length, except setae jl, j 2 , j 3 > s 2 , zh, and si
which are somewhat longer. Coxal spur formula 0-3-2-0* inner
spur on coxae II and III large and usually acute; postero
lateral spur on coxa II acute to blunt* outer spur on coxa III 97
moderate in size and acute. Setae av^ and pv^ on tarsus II
stout and claw-like; all other leg setae present and normal.
Collection Records
Citellus tridecemlineatus (M itehill)
NE3RASKA: Clay Co., Saronville; Colli E. V. Konarek; 3 September
1936| 1 female (RWS),
Thomomys bottae (Eydoux and Gervals)
COLORADO: El Paso Co.; (K iller and Ward, i 960 ).
Thomomys talpoides (Richardson)
COLORADO: El Paso Co. j (K iller and V.'ard, i 960 ),
*Geomys bursarius (Shaw)
COLORADO! El Paso Co.; (M iller and Ward, i 960 ).
*KANSASi Leavenworth Co., Fort Leavenworth; Colli H. L . Keegan*
6 March 19^; 1 female (Holotype) (USNM), (Keegan, 19*^6)*
NEBRASKA: Clay Co,, Saronvillej Colli E. V. Komarok; 2, 3
September 1936; 5 females (RWS),
TEXAS: Leon Co.; Colli 0, E, Hunt; 23 April, 19 November I9^9l
3 females (RWS),
Hirstionyssus (H,) thomomys Allred and Bock, I 966
(Figs. 19-21)
Hirstionyssus thomomys Allred and Beck, 1966, Brigham Young Univ. Soi,
Bull., Biol. Ser. 8(l)i26 [Typei E. Bald Ktn., Summit Co.,
Utah; Holotype, Dept, of Zool, and Entoraol., Brigham Young
Univ., Provo, U tah], D ia g n o sis
Deutosternal denticles about 15 in numberj arranged in
alternating double file. Anterior margin of sternal shield
slightly concavej posterior margin slightly concave, usually
no further than setae st. 3» Line formed by end of genital
flap ribs slightly but distinctly arched. Setae Jvl off
genital shield on soft integument, but usually touching
margin of shield. Paranal setae positioned at level near
middle of or slightly anterior to middle of anal field.
Peritrome subequal in width, usually slightly wider
posteriorly! extending to anterior half of coxa I; normal
in appearance throughout. Groatest width of dorsal shield
always at level of s^ setae; lateral sides straight to
slightly convex and converging toward posterior end; posterior
end in shape of narrow, sharp wedge. All dorsal setae present
and normally developed. Setae Zl moved very close to SI,
distance betweon setae Zl and SI less than length of Zl. Setae
Z5 and sometimes S3 slightly barbed. All dorsal sotae moderate
to large, subequal in length, but with J2, j 3 » z2, z^, s3 » an d
sb slightly longer. Coxal spur formula 0-3-2-0; inner spur on
coxa II moderate In size and usually blunt} postero-lateral
spur on coxa II blunt to roundod; inner spur on coxa III la r g e ,
and acute; outer spur on coxa III moderate in size and acuta
to blunt. Setae av^ and pv^ on tarsus II stout and claw-like.
All other leg setae present and normal. Collection Records
*Thomomys talpoldos (Richardson)
IDAHOt R u tte C o ., Id ah o F a l l s , AEC-NRTSj 13 June 1967j 1 fomal©
(DMA).
♦UTAH: Summit Co., E. Paid Ktn.j Colli D. M. Allrod and M. L.
K lllp a c k ; 8 August 1957J 1 female (Holotype) (DM)} Utah Co.,
Provo Canyonj July 1956} 1 female (DM), (Allrod and Beck, 19 66 ) .
Thomomys sp.
IDAHO 1 Butte Co., Idaho Falls, AKC-NRTS} 20 October 1966*
3 females (DM).
Hirstlonyssus (H.) longlchelae Allred and Reck, I 966
(Figs. 22-2*0
Hirstionyssus geomydls, Allred (not Keegan, 19*^6), I957f Amer.
Midland Natur, 57(2) ii+5**—i*'56 (fig s., host, d istr,),
Hirstionyssus longichelae Allred and Bock, 1966, Brigham Young Univ.
Sci, Bull., Biol. Sor. 8(l)i25 ^Typei Diamond Fork Canyon,
Utah Co., Utah} Holotype, Dept, of Zool. and Entomol.,
Brigham Young Univ., Provo, Utah] .
D ia g n o sis
Doutosternal denticles 1*V - 16 in number; arranged in
alternating double file. Anterior margin of sternal shield
straight to slightly convex; posterior margin slightly to
strongly convex. Lino formed by ond of genital flap ribs
usually slightly but distinctly arched. Setae Jvl off genital 100
shield on soft integument, but usually touching margin of
shield, Paranal setae positioned at level near middle of
or distinctly anterior to middle of anal field, Peritreme
subequal in width, usually slightly wider posteriorly*
extending to near middle of coxa I* normal in appearance
throughout. Greatest width of dorsal shield usually at
l e v e l o f sk setae* lateral sides straight or nearly so and
approximately parallel* posterior end in shape of broad,
blunt wedge. All dorsal setae present and normally developed.
Distance between setae Zl and S i greater than length of Zl,
Setae Z5» S5» S^, S3, and J3 usually slightly barbed. All
dorsal setae small, subequal in length, with setae j 2 » j3 ,
z2, si, and s2 slightly longer. Coxal spur formula 0-3-2-0*
inner spur on coxa II moderate in size and rounded* postero
lateral spur on coxa II blunt* inner spur on coxa III v e r y
large and rounded* outer spur on coxa III small and acute to
blunt. Setae av^ and pv^ on tarsus II stout and claw-like.
All other leg setae present and normal.
Collection Records
Ochotona princeps (Richardson)
UTAH: Summit Co., Bald Ktn.* Coll: D, M, Allred* 8 August 1957*
1 fem ale (DMA), (A llro d and Beck, 19 66 ).
Thomomys bottao (Eydoux and Gervais)
CALIFORNIA: Plumas C o ., Thompson Creek* C o ll: E . W, Jameson, Jr.*
16 October 19^8* 1 female (EWJ), 101
♦Thomomys talpoidos (Richardson)
IDAHO: 3utto Co., Idaho Falls, ASC-NRTS; 7 March 1967; 1 female
( DMA)•
OREGON: Hood River Co.| Coll: E. Kardos; 21 Octobor 1956;
4 fo m alo s (GWK).
♦UTAH: Duchssno Co., Timber Canyon; Coll: D. M. Allred;
20 August ‘1957; 2 females (DMA); Summit Co., Kt. Baldy; Coll:
C, D. Jorgensen et a l.; 25 August 1956; 1 female (DMA); Utah
Co., Diamond Fork Canyon; Coll: D. M. Allred; 2 6 Juno 1951;
1 female (Holotypo) (DM), (Allred and Beck, 1966),
Peromyscus maniculatus (Baird)
IDAHO: Butte Co., Idaho F alls, AEC-NRTS; 22 September 1966;
1 fem ale (DMA).
UTAH: Garfield Co., No. 3ouldorj Coll: R. Loo and S. Durrant;
15 July 1953* 2 females (DMA), (Allrod, 1957b, m isidentified as
H, go o m y d js) .
Onvchomys loucogastor (Niod-Kouwied)
IDAHO: Butte Co., Idaho Falls, AEC-NRTS; 30 July 1966; 1 female
(DMA),
Nustola fronata Lichtenstein
UTAH: Iron Co., Cedar Breaks Nat. Monuraont; Coll: D, M. Allred;
18 July 1953; 1 female (DMA), (Allred and Beck, 1966, m isidenti-
fiod as H» fomuralis). 102
Hirstionyssus (H. ) h llli (Jameson, 1950)
(Pigs, 25-26)
Noolchoronyssus h llli Jameson, 1950, Proc, Entoraol. Soc, Washington
52(4)s165-166 [Types No. Reno, Washoe Co., Nev. j Holotype,
U. S . N. M .] .
Hirstionyssus h llli, Strandtmann and Morlan, 1953» Texas Rep. 3iol.
Ked. 11(4):629 (key, figs.)} Strandtmann and Wharton, 1958t
Inst, Acarology Contrib, 4sl04 (synon,, host, d istr,)j
Allred and Beck, 1966 , Brigham Young Univ. Sci. Bull,, Biol.
Ser, 8(1):30 (key, figs., host, distr.).
D ia g n o sis
Deutosternal denticles 12 - 17 in number} arranged in
alternating double file. Anterior margin of sternal shield
irregularly straight to slightly convex; posterior margin
moderately concave (second pore to lovol of setae st, 2 ),
Line formed by end of genital flap ribs irregularly straight
to slightly arched. Setae Jvl off genital shield on soft
integument, and usually touching margin of shield. Paranal
setae positioned at level noar middle of or slightly
posterior to middle of anal field. Feritremo much wider
posteriorly than anteriorly, with posterior half distinctly
divided and appearing to be broken up; extending to anterior
half of coxa II, Place of greatest width of dorsal shield
variable from lovel of setae s4 to level of SI; lateral sides 103
straight and parallel* posterior end in shape of narrow, sharp
wedge. All dorsal setae present and normally developed. Dis
tance between setae Zl and Si equal to or less than length of
Zl, Setae Z5, S5, and S^ slightly barbed. All dorsal setae
moderate to large and subequal in length with anterior
podonotal setae somewhat longer. Coxal spur formula 0-3-2-0*
inner spur on coxae II and III moderate to large and acute to
b l u n t j postero-lateral spur on coxa II large and acute to
blunt( outer spur on coxa III small and acute to blunt. All
leg setae present and normal.
Collection Records
Eutamtas minimus (Bachman)
IDAHOt Butte Co., Idaho Falls, ASC-NRTS* 5 July 19671 1 female
(DMA).
Perognathus flavus Baird
UTAH: San Juan Co., Kontozuma Creekj Coll: D, M. Allrodj 7 June
1955; 5 females (DMA), (Allred and Beck, 1966),
Perognathus longimombri3 (Coues)
UTAHt Washington Co,, Beaver Dam Wash} Coll: D, M. Allred*
17 A p r il 1952* 1 fem ale (DMA), (A llre d and Beck, 1 9 6 6 ).
* Perognathus parvus (Poale)
CALIFORNIA: Lassen Co., E. Chilcoot; 16 October 19^9; (Jameson,
1 9 5 0 b ).
IDAHO: Butte Co., Idaho Falls, AEC-NRTS * 22 May, 15-16 July,
10 August* 11 fe m a le s (DMA). 104 -
♦NEVADA: Washoe Co., No, Reno* Coll: R. Traub and E. W.
Jameson, Jr.j 24 August 1949* 1 female (Holotype) (USNM),
(Jameson, 1950b)j Nye Co., Mercury, AEC-NTS; (Allred, 19^3).
UTAH: Carbon Co., Pricej Coll: D. M. Allred and D E, Beck*
8 June 1 9 5 1 I 4 fe m a le s (DMA); Ju ab C o ., Jo y ; C o ll: D E . B eckj
1 June 1951* 10 females (DMA)* Kano Co., Cottonwood Creek* Coll*
D. M. Allred et a l.* 10 Juno 1958* 4 females (DMA)* Utah Co.,
Cedar Valley* Coll: L. Woodbury* 3 October 1951* 1 female (DMA),
(Allred and Bock, 1966).
Ferognathus sp.
NEVADA: Nye Co., Mercury, AEC-NTS* 20 A pril, 21 June i960*
4 fe m a le s (DMA).
UTAH: V/ayne Co.* (Allred and Bock, 1966).
Dipodomys o rd ii VJoodhouse
UTAH* Tooele Co., Tooele* 15 August - 7 November 1951* (Keegan,
1953)* Utah Co.; (Elzinga, i 960 * Allred and Bock, 1966 ).
Poromyscus eromicus (Baird)
UTAH: Washington Co., Beaver Dam Wash* Coll: G. Grant et a l. 1
16 A p r il 1952 * 5 fe m a le s (DMA), ( A llr e d , 1 9 5 7 b ).
Poromyscus maniculatus (’Wagner)
IDAHO: B u tte C o ., Id a h o F a l l s , AEC-NRTS; 19 A ugust 1967* 1 fe m a le
(DMA).
UTAH: Kane Co.* Sevier Co.* (Allred, 1957b)* Tooole Co., Tooele;
15 November 1951* (Keegan, 1953)* Utah Co.; (Elzinga, i 960 * A llr e d
and Bock, I 966 ). 105
Onychomys leucogaster (Wied-Heuwied)
IDAHOt Butte Co., Idaho Falls, AEC-NRTS* 30 July 1966; 1 female
(DMA).
UTAHi Kane Co,, AdairviUe ; Colit C. Pritchett ; 21 June I960;
1 femalo (DMA), (Allrod and Beck, 1966); Tooolo Co.* (Ho, 1962*
Allrod and Beck, 1966); Utah Co.* (Elzinga, I960; Allred and
Beck, 1966),
Nootoma lopida lepida Thomas
UTAHt Tooele Co., 15 August 195i» (Keegan, 1953)*
Hirstionyssus (H.) "C" n. sp.
(Figs. 27-28)
Neoichoronyssus h illi, Jamoson, 1950, Proc. Entomol. Soc. Washington
52(4): 165-166 (key), (record from Ferognathus hispidus only),
Hirstionyssus h illi, Allred, 1957 * Anor. Midland Natur. 57 (2 ) i 456
(figs., host, d istr.), (record from Poromyscus eromicus only);
Allred, 1963, Proc. Entomol, Soc. Washington 65 (3)t232 (host,
d istr,), (rocord from Ferognathus formosns only).
D ia g n o sis
Deutostemal donticlos 14 - 16 in number* arranged in
alternating double file. Anterior margin of stom al shield
slightly convex; posterior margin moderately concave (second
poro to level of setae st. 2). Lino formed by end of genital
flap ribs straight to slightly arched. Setae Jvl off genital
shield on soft integument, but usually touching margin of 106
shield, Paranal setae positioned at level near middle of or
slightly posterior to middle of anal field. Peritrema sub
equal in width, usually slightly wider posteriorly; extending
to anterior half of coxa II; normal in appearance throughout.
Greatest width of dorsal shield at level of s3 setae; lateral
sides straight and converging slightly toward posterior end;
posterior end in shape of narrow, sharp wedge. All dorsal
setae present and normally developed. Distance between setae
Zl and SI equal to or less than length of Zl. Setae Z5 and
S5 slightly barbed. All dorsal setae moderate to large, sub
equal in length with anterior podonotal setae somewhat longer,
Coxal spur formula 0-3-2-0; inner spur on coxa II largo and
usually blunt; postero-lateral spur on coxa II moderate in
size and acute to blunt; inner spur on coxa III moderate in
size and usually blunt; outer spur on coxa III small and acute
to blunt. All log setae present and normal.
Collection Records
♦Ferognathus formosus Korriam
♦NEVADA: Nye Co., Mercury, AEC-NTSj 19 August I960; 1 female (DMA)j
13 Juno 1961; 1 female (Holotype) (DMA), (Allred, 1963, m isidentl-
f ie d a s H. h i l l i ) .
Porognathus hispidus Baird
KANSAS: Moade C o ., SW Meade; Coll* J. E, K ill and P. E. Crowe;
July 19^0; 1 female (E'.vJ), (Jameson, 1950b, m isidentifiod as
H. h i l l i ) . 107
Perorayscus eremicus (Baird)
UTAHt Washington Co., Rockville; Colli D E. Reck et a l.|
13 July 1953; 2 females (DM), (Allrod, 1957b, m isidontified
as H, h llli).
"unknown host”
NEVADA: Nye Co., Mercury, AEC-NTS; 28 August I959l 1 female (DMA).
Type M aterial
Holotype female from Porognathus formosus, collected at the
Nevada Test Site, Mercury, Nye Co., Nevada, 13 June 19&1, collection code 6FB, field no. 13; one paratype female from samo host and locality, collocted 19 August 19^0, collection code 6CL, field no. ^+51 to be deposited in the acarology collection. Department of Zoology and
Entomology, Brigham Young University, Provo, Utah, One paratype female
from Ferognathus hispidus, collocted 13 miles southwest of Koade,
Koade Co., Kansas, July 19^0, by J. E, H ill and P, E. Crowo; in the
collection of E. '■/, Jameson, Jr., Department of Zoology, University of
California, Davis, California.
Hirstionyssus (H.) incomptis (Eads and Hightower, 1952)
(Figs. 29-30)
Neoichoronyssus incomptis Eads and Hightower, 1952, Proc, Entomol. Soc.
Washington 5^(1) :32-35 [Type: Terry Co., Texas: Holotype,
U. S. N. M.] .
Hirstionyssus incomptis, Strandtmann and Morlan, 1953* Texas Rep,
Biol. Mod. 11(4-) j629 (fig s., key); Strandtmann and Wharton, 108
1958, Inst, Acarology Contrib. ^jlO^ (synon., host, d istr.);
Allred and Beck, 1966, Brigham Young Univ. Sci. B ull., Biol,
Ser, 8(i)t30-31 (key, figs,, host, distr.).
D ia g n o sis
Deutostemal denticles usually about 19 in number ; arranged in
alternating double file* however, in the holotype female there
were 30 arranged in regular double file. Anterior margin of
sternal shield straight to slightly concave; posterior margin
deeply concave (distinctly anterior to setae st. 2). Lina
formed by end of genital flap ribs slightly to strongly arched*
Setae Jvl off genital shield on soft integument, but usually
touching margin of shield. Paranal setae positioned at level
near middle of anal field. Peritreme subequal in width,
usually slightly wider posteriorly; extending to near middle
of coxa I; normal in appearance throughout. Greatest width of
dorsal shield always at level of s3 setae; lateral sides
straight or nearly so and converging toward posterior ond;
posterior ond in shape of narrow, sharp wedge. All dorsal
setae present and normally developed. Setae Z5» S5» S^, and
sometimes S3 slightly barbed. All dorsal sotae large, sub
equal in length with anterior podonotal setae somewhat longer.
Coxal spur formula 0-3-2-0; inner spur on coxae II and III
moderate in size and acute to blunt; postero-lateral spur on
coxa II acute to blunt; outer spur on coxa III small and acute
to blunt. All leg setae present and normal. 109
Collection Records
Cite H us leucurus (Kerriam)
UTAH: Tooele Co,, Granite Peak; Coll: E. A, Shippa; 19 August
19531 1 female (DMA); Utah Co.; (Elzinga, I960; Allred and Beck, 1966 ).
Citellus varlegatus (Erxloben)
UTAH: Washington Co.; (Allred and Beck, I 966 ).
Eutamias sp,
IDAHO: B u tte C o ., Id ah o F a l l s , AEC-HRTS; 7 June 1966; 1 fem ale
(DMA).
Ferognathus longimembris (Coues)
UTAH: Kane Co., Hole-In-The-Rockj Coll: D E, Beck 16 Kay 1953;
1 female (DMA); same Co., E. Kanab; Coll; D. M, Allrod and C. L.
Hayward; 20 May 1956; 1 female (DMA), (Allred and Beck, 1966).
Dipodomys merrlami Kearns
UTAH: Washington Co,, Grafton; Coll: D E, Beck and D, M, Allred;
17 December 1950; 1 fom ale (DMA), ( A llre d and B eck, 1966 ).
* Dipodomys o rdli Voodhouse
IDAHO: B u tte C o ., Id a h o F a l l s , AEC-HRTS; 7 1 3, 15. 16, 30 June,
16 July, 12 August, 27 October 1966 , and 2 July 1967; 9 females
(DMA).
*TEXASi Terry Co.; Colli V. I. Kilos; 16 January 19^7, 1 female
(Holotype) (USHM), (Eads and Hightower, 1952); same locality;
Coll: W, H. Williams; 16 January 19^8; (Eads, Kenzies, and Milos,
1952)| Bailey Co.; Colli E, Garrett; 1^ March 1965; 1 female (RWS); Cochran Co.; Colli W. H. YJilliams and T, Crossland; 21 April
19^3; Dawson C o ,; C o ll: V. I . M ile s ; 5 M arch 19*^8; (Eads an d
Hightower, 1952); Dawson Co.; Coll: G. Carpenter; 3 March 19**8|
Gaines Co.; Coll: K« J. George; 11 March 1$&9» (Eads, Menzies, and Miles, 1952).
UTAH: Box Elder Co.; Carbon Co.; Daggett Co., Bridgeport; Coll*
D E. Beck; 11 June 1953* 2 fe m a le s (DMA); D uchesne C o ., R o o s e v e lt;
C o ll: D E. Bock; fe m a le s (DMA); same l o c a l i t y j C o ll: M. L .
K illp a c k ; 22 A ugust 1952; 1 fe m a le (DMA); Em ery C o .; Juab C o .,
Sand Dunes; CoU: D E. Beck et a l.; 3 April 1951; 6 females (DMA); same locality; Coll: H. J. Phillips; 18 June 1955; 1 female (DMA);
Kane Co., E, Kanab; Coll: M. D. Coffey; 9 May 1952; females
(DMA); San Ju an C o ., So. Koab; C o ll: D, K. A llr e d o t a l . ; 7 -8
May 1951; 10 fe m a le s (DMA); same C o ., B lu f f; C o ll: D E . Beck e t a l . ; 3 -5 May 1951; 13 fe m a le s (DMA); San F e te C o ,; S e v ie r C o .,
Richfied; Coll: D E. Beck; 30 April 1952; 7 females (DMA);
Tooele Co., Wig Mtn.; Coll: E. A. Shippo; lh May 1953; 1 female
(DMA); same locality; CoU: D, Parker and J, Smith; 23 Juno 19531
1 fem ale (DMA); same C o ., S k u ll V a lle y ; 27 May 195**; 1 fe m a le
(DMA), (Allred and Beck, 1966); Utah Co.; (Elzinga, I960; 196*0;
Utah Co., Lyndyll; Coll: D. M. Allred; 7 October 1950; 1 female
(DMA); same C o ., C edar V a lle y ; C o ll: D, M. A llre d e t a l . ;
25 March 1951; 1 female (DMA); same Co., Chimney Rock Pass; Coll:
M, D. Coffey; 23-28 April 1953* ** females (DMA); same locality; Ill
Colit D E. Back et a l,; 11 Juno 19531 1 female (DMA); Washington
Co.; Wayne Co.; (Allred and Beck, 1966 ).
Dipodomys microps (Kerriam)
UTAH: Juab Co.; Tooolo Co.; Utah Co.; Washington Co.; (Allred
and Beck, I 966 ).
Dipodomys sp,
UTAH; Grand Co.; San Juan Co., Arches Nat, Monument; Coll: V. J,
T ip to n ; 12 May 19^9; 2 fe m a le s (DMA), (.A llre d an d Beck, I 966 ) .
Reithrodontomys megalotjs (Baird)
UTAH: Tooele Co.; Utah Co.; (Ho, 1962); Washington Co.; (Allred
and Beck, I 966 ).
Poromyscus crinitus (Kerriam)
UTAHt San Juan Co., Red Mesa; Coll: D E. Beck et a l,; 27 August
1953; 1 fem ale (DMA), ( A llr e d , 1 9 5 7 b ).
Poromyscus orenicus (Baird)
UTAH: Washington Co., Leeds; Coll: D 5. Bock and K. L. Killpack;
1^ J u ly 1953; 2 fem a le s (DMA), ( A llre d and Bock, I 966 ).
Feromyscus maniculatus (Wagner)
IDAHO: Butte Co., Idaho Balls, AEC-NRTS; 16 November 1966, 6 J u ly
1953; 2 fe m a le s (DMA).
UTAH: Carbon Co.; (Allred, 1957b); Juab Co.; (Allred and Beck,
1966)1 San Juan Co., Bluff; Coll: D. K. Allrod et a l.; 3 May
1951l 1 female (DMA), (Allred, 195?b); Tooele Co.; Utah Co.,
Chimney Rock Pass; Coll: K. D. Coffey; 23 April 1953; 2 females 112
(DMA)| Wayne Co., Doad-Horse-Point; Colli D S. Beck; 10 July
1958; 1 female (DMA), (Allred and Beck, 1966),
Ne otoma lepida lopida Thomas
UTAH: Tooele Co.; (Allred and Roscoe, 1957); Utah Co.; (Allred
and Beck, 1966).
Klcrotus longicaudus (torriam )
UTAH: Garfield Co.; (Allred and Beck, 1966),
Rattus norvogicus (Berkenhout)
OTAHi Utah Co.; (Allred and Beck, 1966 ).
Hirstionyssus (H.) triacanthus (Jameson, 1950)
(Figs. 31-32)
Keolchoronyssus triacanthus Jameson, 1950, Proc. Entomol. Soc.
’Washington 52(4) :166 ^ Typo 1 Pyramid Lake, Washoe Co.,
Nevada; Holotype, U, S. N. M.} .
Hirstionyssus triacanthus, Strandtmann and I'orlan, 1956, Texas Rep.
Biol, tod. 11(4):629 (key, figs*); Strandtmann and Wharton,
1958, Inst. Acarology Contrib. 4:108 (synon., host, d istr.);
Allrod and Bock, 1966, Brigham Young Univ. Sci. B ull., Biol.
Ser. 8(1):30 (koy, figs., host, d istr.).
D ia g n o sis
Deutosternal denticles usually about 21 In number; arranged
in alternating double file; however, in the holotype female
there wore 26 arranged in regular doublo file. Anterior
margin of sternal shield slightly concave to slightly convex; posterior margin moderately to deeply concave (second pore
to beyond setae st. 2). Line formed, by end of genital flap
ribs slightly but distinctly arched. Setae Jvl off genital
shield on soft integument, but usually touching margin of
shield. Paranal setae positioned at level near middle of
anal field. Feritreme considerably wider anteriorly than
posteriorlyi extending to anterior half of coxa Ij normal
in appearance throughout. Greatest width of dorsal shield
always at level of s3 - sh setae; lateral sides straight or
nearly so and converging toward posterior end; posterior end
in shape of narrow, sharp wedge. All dorsal setae present
and normally developed. Setae Z5* S5» and S4 usually barbod
Distance between setae Z1 and SI much less than length of Zl
All dorsal setae very robust and long, length usually equal
to or greater than distance between adjacent setae. Coxal
spur formula 0-3~2-0; inner spur on coxae II and III large
and acute | postero-lateral spur on coxa II largo and acuta
to blunt; outer spur on coxa III moderate in size and acute.
All log setae present and normal.
Collection Records
Lopus californicus Gray
UTAH: Juab Co., Goshen Meadow; Coll: D. Parker and J. Smith;
July 1953} 1 female (DMA); Tooele Co.; (Allred and Beck, 1966 ). UA
C itellus tovmsenclll (Bachman)
IDAHOj B u tto C o.i Id ah o F a l l s , AEC-NRTS; 1 6 -1 8 June 1 9 6 ?I
2 fe m a le s (DMA),
Citollus loucurus (Merriara)
UTAH: Tooele Co.| Utah Co.; (Elzinga, I960; Allred and Beck, 1966 ).
Perognathus longlmombris (Coues)
UTAH: Kane Co,; (Allred and Beck, 1966),
Perognathus parvus (Pealo)
IDAHO: B u tte C o ., Id ah o F a l l s , AEC-NRTS; 8 June 19 6 ?; 1 female
(DMA).
Ferognathus formosus Ferriara
UTAH: T ooele Co . 5 (Allred and Beck, I 966 ).
* Dipod omys merriami Koams
ARIZONA: Pima Co., Tucson; Coll: C. E, Yunkor; 16 December 1958|
1 fem ale (OSU),
* NEVADA: 'Washoe Co., Pyramid Lake j Coll: E. W. Jamoson, Jr.;
9 October 19^9; 1 female (Holotype) (USNM)* (Jameson, 1950b);
Nye Co., Mercury, AEC-NTS; Coll: M. A. Goates; 1? March, 7 Juno,
10 Ju n e 1961; 7 fe m a le s (DMA), (G o a te s, 1963 ),
UTAH: Washington Co., Grafton; Coll: D. 11. Allred; 17 December
19501 2 females (DMA); same Co., Beaver Dam Wash; 21 December
1950* 3 fem ales (DMA), (A llre d an d B eck, 1966 ).
Pipodomys ordil Woodhouso
IDAHO: Butto Co., Idaho Falls, AEC-NRTSj 27 January, 21, 2b, 26 Juno, 31 July, 10, 11, 16 August 1966, 19 May, 8 Juno 19671
10 fe m a le s (DMA),
NEVADAt Nye Co., Mercury, ASC-NTS; h A ugust 1 9 6 li 2 fe m a le s (DMA).
TEXASi Dawson Co. 1 Colli M, J. George; 5 March 19*4-8; (Eads,
Nenzies, and Miles, 1952).
UTAHi Beaver Co,| Box Elder Co., Lucin; Colli D E. Beck and M, L.
Killpack; 18 June 1952j ^ females (DMA); Daggett Co.; Duchesne Co. 1
Emery Co.} Juab Co., Joyi Coll: D. I-!. Allred et al.} 1 June 1951*
1 female (DMA)} same Co., Sand Dunes} Coll: D E. Beck et al. |
8 A p r il 19511 6 fe m a le s (DMA); same l o c a l i t y ; C o ll: H. J . P h illip s *
19-26 June 1955* 9 females (DMA); same locality; Coll: D. M.
Allred; 7 October 1950* 1 female (DMA); Kane Co.; M illard Co.;
San Juan Co., Bluff; Coll: D S. Beck and D. K. Allrod; 3“5 Ma7
1951* 5 females (DMA); same Co., So, Hoab; Coll: D. M. Allred
e t a l . ; 7 May 1951* 3 fo m a le s (DMA); San P e te C o .; C o ll: D E . Beck;
22 July 1951* 2 females (DMA), (Allred and Beck, 1966); Tooele Co.,
Tooele; 25 August - 23 October 1951* (Keegan, 1953)* Utah Co.;
(Elzinga, i960; 196 **-)* Utah Co., Cedar Valley; Coll: D. M. Allred;
1*4- A ugust 1957* 1 fem ale (DMA); 25 M arch 1961; 3 fe m a le s (DMA); same
Co., Chimney Rock Pass; Coll: M. D. Coffoy; 23 April 1953* 1 fornalo
(DMA); U in ta h C o .; V /ashington C o .; (A llro d and Bock, I 966 ).
Dipodomys microps (Kerriam)
NEVADA: Nye Co., Morcury, AEC-NTS; 18, 25 Kay, 31 October,
November, 6 , 21 December i 960 , 5 March, 7, H , 22 April, 8,
10 Juno 1961; 16 fe m a le s (DMA), (G o a te s, I 963 ). 116
UTAH: Box Elrior Co., Lucin; Coll: D E, Bock} 19 Juno 1952;
1 female (DMA); Juab Co.; Millard Co.; San Pete Co.j (Allred
and Beck, 1966); Tooele Co.; (Keegan, 1953)I Utah Co.; (Elzinga,
I960; 1964); Washington Co.; (Allred and Beck, 1966).
Dipodomys deserti Stephens
UTAH: Washington Co., Beaver Dam Wash; Coll: D E. Beck et a l.;
21 December 1950? 1 female (DMA),
Dipodomys sp.
UTAH: Grand Co.; San Juan Co., Arches Nat, Monument; Coll: V. J.
T ip to n j 12 May 19^9; 1 fem ale (DMA), ( A llre d and Beck, 1966 ).
Reithrodontomys megalotis (Baird)
UTAH: Tooele Co.; (Allrod and Beck, 1966); Utah Co.; (Elzinga,
I 960 ; 196 *0 .
Peromyscus crinitus (Korrlara)
UTAH: Emory Co,j (Allred and Beck, I 966 ).
Feromyscus orcmlcus (Baird)
UTAH: Washington Co., Leeds; Coll: D E. Beck et_ al.; 14 July
1953; 4 fe m a le s (DMA), ( A llr e d and B eck, 1966 ),
Peromyscus manlculatus (Wagner)
IDAHO: B u tte C o ., Id a h o F a l l s , AEC-NRTS; 23 Juno I 966 ; 1 fem ale
(DMA).
UTAH: Box Elder Co.; Juab Co.; Tooele Co.; (Allrod and Bock, 1966);
Utah Co.; (Elzinga, I 96 O; 1 9 6 4 ), 117
Onychomys torridus (Coues)
NEVADA: Nye Co., Mercury, AEC-NTS; 10, 27 June 1961 1 2 fe m a le s
(DMA), ( A llr e d , 1 9 6 2 ).
Neotoma lepida lepida Thomas
UTAH: San Pete Co.j (Allred and Beck, 19 66 ).
Neotoma cinerea (Ord)
IDAHO: B u tte C o ., Id a h o F a l l s , AEC-NRTSj 22 S ep tem b er 1 9 6 6 1
1 fem ale (DMA).
Rattus norvoglcus (Berkenhout)
UTAH: Utah Co.j (Allred and Beck, I 966 ),
Chordoiles minor Grinnell
IDAHO: B u tte C o ., Id a h o F a l l s , AEC-NRTS) 3 A ugust 1967*
1 fom ale (DMA).
Sceloporus graclosus Baird and Girard
IDAHO: B u tte C o ., Id a h o F a l l s , AEC-NRTS; 7 S ep tem b er 1966)
1 fem ale (DMA).
Hirstionyssus (H.) "B" n, sp,
(Figs. 33-35)
Hirstionyssus sp., Jamoson, 1950» Mammal, 31:140 (host, d istr.).
Hirstionyssus arcuatus, Strandtmann and Morlan (in part) (not Koch,
1839)» 1953» Texas Rep. Biol, Mod. 11(4):632 (key, fig s., host,
d istr.); Strandtmann and Wharton (in part), 1958, Inst.
Acarology Contrib, 4:100. 118
D ia g n o sis
Doutostornal denticles 15 - 18 in number* arranged in
alternating double file. Anterior margin of sternal shield
slightly to strongly convex* posterior margin moderately
concave (second pore to level of setae st. 2). Line formed
by end of genital flap ribs straight or nearly so. Setae
Jvl off genital shield on soft integument, but touching margin
of shield. Genital shield broad throughout with lateral sides
straight and nearly parallel. Anal shield unusually broad
relative to length* with pair of lateral pores, Faranal setae
positioned at level near middle of or slightly posterior to
middle of anal field. Feritreme subequal in width, usually
slightly wider posteriorly; extending to anterior half of
coxa I* normal in appearance throughout. Greatest vridth of
dorsal shield at level of setae Zlj lateral sides usually
slightly concave and converging slightly toward anterior endj
posterior end broadly rounded. Setae r2, r*J-» SI, and S3 absent*
setae S2 sot in from margin of shield a distance equal to or
greater than length of S2, All other dorsal setae present and
normally developed. Sotao Z5 and usually S5 slightly barbed.
Central dorsal shiold setae small, about half the length of
most anterior, lateral, and posterior marginal setae, Coxal
spur formula 0-2-2-lj inner spur on coxao XI and III largo and
acutej outer spur on coxa III moderate in size and acuto* spur
on coxa IV moderately long, broad, and acute. Setae av-^ and pv^ 119
on tarsus II stout and claw-like. All other leg setae present
and normal.
Collection Records
Sorex fumous M iller
PENNSYLVANIA: Venango Co., S. Oil City; C oll: N. D. Richmond
and J. J, Christiani 4 Kay 1947; 1 female (RWS).
*Blarina brevicauda (Say)
* MARYLAND: Prince Georges Co., Patuxent Refuge; Coll: R, 0.
Drummond and L. Stickol; 21 April 195^* 2 females (OSU)j same
locality and collectors; 29 April 195^t 1 female (Holotype) (OSU),
(Drummond, 1957» m isidentified as H, arcuatus); same Co., Greonbelt;
Coll: C. E. Yunker; 8 August 1953! 1 female (OSU); same Co.,
Riverdale; Coll: F. C. Bishopp; 20 June 1934; ^ females (RWS).
NEW JERSEY: Burlington Co., Koorestown; Coll: G. R. Lunz;
16 April 1933i 1 female (RWS),
NEW YORK: Tompkins Co., 5. Ithaca; Coll: E. W. Jameson, Jr.;
15 March, 24 May 1947; 2 females (RWS); same Co., SW E e seu ch ;
Coll: E, W, Jameson, Jr.; 25 Juno 1948; 2 females (EWJ),
(Jameson, 1950c).
OHIO: Erie Co., Castalia; Coll: R. J. Stoll; 20 June, 14, 28
July, 4, 11 August 1965; 7 females (OSU).
PENNSYLVANIA: Beaver Co., E, Industry; Coll: N. D. Richmond and
J. J, Christian; 20 May 1947; 8 females (RWS); Mercer Co., Mercer;
Coll: N. D, Richmond and J, J. Christian; 16 July 1947; 1 female
(RWS); Pike Co.; Coll: F. Harper; 7 October 19^5; 1 female (RWS); 120
Venango Co., Oil City; Colli N. D. Richmond and J. J. Christian;
2 May 19^7 J i female (RV73) i Wayne Co.; Colli F. Harper j 10 August
19*+6J 1 female (RWS).
Scalopus aquaticus (Linnaeus)
OHIO: Franklin Co., Gahanna; Colli D. E. Johnston; 3 March
1969; 2 females (OSU).
Parascolops breweri (Bachman)
NEW YORK 1 Tompkins Co., E. Kewfield; Coll: E, V7. Jameson, Jr.;
8 July 19^3; 1 female (RWS),
Type M aterial
Holotype female from Blarina brevicauda, collected at the
Patuxent W ildlife Refuge, Prince Georges Co., Maryland, by R, 0.
Drummond and L. Stickel, 29 April 195^» Army collection no. 296 ; t o be deposited in the U, S, National Museum. One paratypo female from
Blarina brevicauda, collected near Castalia, Erie Co., Ohio, h August
1965l in the acarology collection of the Acarology Laboratory, The Ohio
Stato University, Columbus, Ohio. One paratype female from Blarina brevicauda, collected 2 miles east of Industry, Beaver Co,, Pennsylvania,
20 May 19^7» by N. D, Richmond and J. J, Christian; in the collection of
R. W, Strandtmann, Bernice P. Bishop Museum, Honolulu, Hawaii,
Hirstionyssus (H.) talpae Zemskaya, 1955
(Figs. 36-33)
Liponyssus arcuatus, Oudeinans (not Koch, 1839), 1913» Arch. Naturgesch.
Abt. A 79(9)t68. '* .121
Liponyssus occidentalism Jameson (not Ewing, 1923)» 19^7, Univ. Kansas
Publ. Mus. Natur. Hist, 1(7) *1^2 (host, d istr.).
Hirstionyssus arcuatus, Fonseca (sensu Oudemans, 1913» not Koch,' 1839),
19^8, Proc. Zool, Soc. London 118(2) :298 (synon.); Strandtmann
and Korlan (in part), 1953, Texas Rep. Biol. Med. 11(h) :632
(key, figs., host, d istr.)j Strandtmann and Wharton (in part),
1938, Inst. Acarology Contrib. h:103 (synon., host, d istr.).
Hirstionyssus talpae Zemskaya, 1955, in Brogotova ot a l., Oprod. Faune
SS3R 59*380 [Type: U.S.S.R.j Holotype, Zool. Inst. Acad. Sci.
U.S.S.R., Leningrad] ; Bregetova, 1956, Opred. Faune SSSR
61:165-197 (key, figs., host, d istr.); Evans and T ill, 19&6,
Bull. B rit. Mus. (Natur, H ist.) Zool. lh(5) *277-285 (synon.,
key, figs., host, distr,).
Hirstionyssus palustris Allred and Beck, 1966, Brigham Young Univ.
Sci. Bull., Biol, Ser. 8(l):19-20 [Type: Schofield, Carbon
Co,, Utah; Holotype, Dept, of Zool. and Sntomol., Brigham Young
Univ., Provo, Utah] . HEW SYNONYMY,
D ia g n o sis
Deutosternal donticlos 12 - 16 in numbor; arranged basically
in alternating double file. Anterior margin of sternal shield
slightly to strongly convex; posterior margin modoratoly to
sometimes deeply concave (second pore to slightly boyond setae
st, 2), Lino formed by end of genital flap ribs slightly but
distinctly arched. Setae Jvl off genital shield on soft
integument, but touching margin of shield. Genital shield 122
broad throughout, with lateral sides straight and nearly
parallel. Anal shield unusually broad relative to length*
with pair of lateral pores. Paranal setae positioned at
level near middle of or slightly posterior to middle of anal
field. Peritreme subequal in width, usually slightly wider
posteriorly* extending to anterior half of coxa IIj normal
in appearance throughout. Greatest width of dorsal shield
usually at level of Zl setae{ lateral sides slightly concave
to straight and parallel to slightly converging toward
anterior end; posterior end forming very broad, blunt wedge.
Setae r^, SI, and S3 absent. All other dorsal setae present
and normally developed. Setae Z5 and sometimes S5 slightly
barbed. Central dorsal shield setae small, usually half as
long as anterior, lateral, and posterior marginal setae.
Coxal spur formula 0-2-2-1; inner spur on coxae II and III
large and acute; outer spur on coxa III moderate in size and
acute; spur on coxa IV short, broad, and acute to blunt.
Antero-ventral setae on coxa I I spinose or peg-liko, and acute
to blunt. Setae av^ and pv-^ on tarsus I I stout and claw-like.
All other leg setae present and normal.
Collection Records
Sorox cinoreus Kerr
WYOMINGi Albany Co., Brooklyn Lake; Collj E. W. Jameson, Jr.|
29 August 19^31 ^ females (EV.'J and RWS), Sorex furoeus K iller
HEW YORK: Tompkins Co., E, Ithaca; Colls E. W. Jameson, Jr.;
20 August 19^7* 1 female (RWS).
Sorex palustris Richardson
UTAH: Carbon Co., Schofield; Coll: D E. Beck; 29 July i960;
2 females (DllA), (Allred and Beck, 19&6, described as lb palustris).
Blarina brevicauda (Say)
MARYLAND: Prince Georges Co., Riverdale; Coll: R, 0. Drummond;
17 May 195^1 1 female (OSU).
Cryptotis parva (Say)
KANSAS: Douglas Co., Lawrence; Coll: E. W. Jameson, J r.;
30 September 19^5} 3 females (EWJ and RWS), (Jameson, 19^7,
misidontifiod as H. occidentalis).
MARYLAND: Prince Georges Co., Patuxent Refuge; Colit R, 0.
Drummond and L. Stickel; 6 May 195^; 2 females (OSU), (Drummond,
1957 t identified as H, arcuatus).
Slgmodon hispidus Say and Ord
NORTH CAROLINA: Make Co., Raleigh| Coll: R. 0. Drummond;
21 June 1953; 1 female (OSU).
Mote
In addition to the Nearctlc distribution given abovo, H. talpae is recorded by various authors from a number of insectivoros and some rodents throughout the Palearctic region and in parts of the Oriental region. The typo host and locality are Talpa ouropaea in the
U, S, S, R, Typo specimens could not be obtained for this study, but 124 several specimens from the typo host and locality, identified by
N, G. Bregetova, were made available by the B ritish Museum.
Hirstionyssus (H.) affinis (Jameson, 1950)
(Figs. 39-42)
Neoichoronyssus affinis Jameson, 1950* Proc. Entomol. Soc. Washington
52(4)i 163-165 [Typei No. Quincy, Plumas Co., Calif. 1 Holo
type, U. S. N. M. ] .
Hirstionyssus affin is, Strandtmann and Morlan, Texas Rep. Biol. Med.
Il(4)s630 (key, fig s.); Strandtmann and Wharton, 1958* Inst,
Acarology Contrib. 4ii00-101 (synon., host).
D ia g n o sis
Deutostomal denticles 15 in number; arranged in alternating
double file. Anterior margin of sternal shield moderately con
vex; posterior margin moderately concave (second pore to setae
st. 2), Lino formed by end of genital flap ribs slightly but
distinctly arched, Sotao Jvl off gonital shield on soft
integument but touching margin of shield. Paranal setae
positioned at level near middle of anal field. Poritreme sub
equal in width, usually slightly wider posteriorly; extending
only to posterior half of coxa II; normal in appearance through
out, Greatest width of dorsal shield at level of setae Zl - Si;
lateral sides straight or nearly so and almost parallel;
posterior end in shape of broad, blunt wedge. All dorsal setae
present and normally developed. Distance between setae Zl and SI equal to or less than length of Zl. Setae Z$, S5, and S4
usually slightly barbod. Central dorsal shield setae somewhat
smaller than anterior, lateral, and posterior marginal setae.
Coxal spur formula 0-2-2-1; inner spur on coxae II and III
moderate in size and rounded to almost truncate; outer spur
on coxa III small and acute; spur on coxa IV short, broad, and
acuto. Setae av^ and pv^ on tarsus II stout and claw-likej
seta av^ on tarsus IV acutely spiniform. All other leg setae
present and normal.
Collection Records
♦Tamiasciurus douglasii (Bachman)
♦CALIFORNIA: Plumas Co., No, Quincy; Coll: E. W, Jameson, Jr.
I5 Kay 1549; 1 female (Holotype) (U3NM), (Jameson, 1950b).
Hirstionyssus (H.) "D" n, sp.
(Figs. 43-46)
Hirstionyssus affinis, Strandtmann and 1'orlan (in part) (not Jameson,
1950b), 1953. Texas Rep. Biol. Ked, 11(4):630 (key, figs.,
host, d istr,); Strandtmann and Wharton (in part), 195^, Inst.
Acarology Contrib, 4:100 (synon., host, d istr,); Allred and
Bock, 1966, Brigham Young Univ. Sci. B ull,, Biol. Ser. 8(1):19
(key, figs., host, distr,). 126
D ia g n o sis
Deutostemal denticles 15 - 18 in number; arranged in
alternating double file. Anterior margin of sternal shield
slightly to moderately convex; posterior margin moderately
concave (second pore to setae st. 2), Line formed by end of
genital flap ribs usually irregularly straight. Setae Jvl
off genital shield on soft integument, but sometimes touching
shield. Paranal setae positioned at level near middle of or
slightly posterior to middle of anal field, Peritreme sub
equal in width, usually slightly wider posteriorly; extending
to anterior margin of coxa II or posterior margin of coxa I;
normal in appoarance throughout. Greatest width of dorsal
shield usually at level of Zl setae, although may be approxi
mately same at level of setae s3 - s*J*; lateral sidos straight
or nearly so and approximately parallel; posterior end forming
broad, blunt to narrow, sharp wedge. All dorsal setae present
and normally developed. Distance between setae Zl and SI
slightly greator than length of Zl. Setae Z5» S5» and S^
usually slightly barbed. Central dorsal shield setae somewhat
smaller than anterior, lateral, and posterior marginal setae.
Coxal spur formula 0-2-2-0j inner spur on coxae II and III
moderate in size and rounded to almost truncate; outer spur on
. coxa III moderate to small in size and acute. Setae av-^ and pv^
on tarsus II stout and claw-like; seta av^ on tarsus IV acutoly
spiniform. All other leg setae present and normal. Collection P.ocordg
*Butamias mini mus (Bachman)
CALIFORNIA! Modoc C o., A ltu r a s ; C o lls Kirkwood j 22 Juno 19^9;
1 fem ale (RWS), (Strandtmann and Morlan, 1953)*
’•'UTAH: Box Elder Co., Yost; Colls D E. Becks11 J u ly 1957*
6 females (Holotype) (DMA); same Co., George Creek; Colls D E,
Beck et a l.; 7 December 1957J 1 female (DMA), (Allred and Bock,
1966 ).
Foromyscus boylli (Baird)
CALIFORNIA: Plumas Co., E. Quincy; Coll: E. W, Jameson, Jr.;
*4- October 19501 1 female (RWS), (Strandtmann and Morlan, 1953)*
Type M aterial
Holotype female from Butamias minimus, collected at Yost, Box
Elder Co., Utah, by D E, Bock, 11 July 1957, field no. ^200; to bo doposited in the acarology colloction, Department of Zoology and
Entomology, Brigham Young University, Provo, Utah. One paratypo
female from Eutamias minimus, collected at Alturas, Modoc Co.,
California, 22 June 19^9, by Kirkwoodj and one paratypo femalo from
Peromyscus boylli, collected 5 miles oast of Quincy, Plumas Co.,
C alifornia, *4- October 1950» by S. vl, Jameson, Jr.; in ‘the colloction
of R, W. Strandtmann, Bernice P. Bishop Museum, Honolulu, Hawaii, 128
Hlrstionyssus (H.) occidentalis (Ewing, 1923)
(Figs. ^7-50)
Liponyssus occidentalis Ewing, (1923), Proc, U. S, Nat, Hus. 62(13)*20
^Typet Lolo, Missoula Co., Mont.; Lectotype, U. S. N. M,J 1
Radford, 1943, Parasitol. 35:64-65 (catalogue).
Hirstionyssus occidentalis, Jameson, 1950, Proc. Biol, Soc. Washington
63:31-32 (key)| Strandtmann and Morlan, 1953, Texas Rep. Biol.
Med, ll(4):628-632, 636 (key, figs., descr. note); Strandtmann,
1956, J. Kansas Entomol. Soc. 29(4);138 (descr. note);
Strandtmann and Wharton, 1958, Inst. Acarology Contrib. 4il06
(synon., host, d istr.).
Neolchoronyssus occidentalis, Jameson, 1950, Proc. Entomol. Soc.
'Washington 52(4) : 162-163 (key).
Hirstionyssus punctatus Allred and Reck, 1966, Brigham Young Univ. Sci.
Bull., Biol. Ser. 8(1):20 [Type: Bald Ktn., Summit Co., Utah;
Holotype, Dept, of Zool. and Entomol., Brigham Young Univ.,
P ro v o , U t a h ] , NEW SYNONYMY.
D ia g n o sis
Doutosternal denticles 15 - 18 in number; arranged in alternating
double file. Anterior margin of sternal shield slightly convex;
posterior margin moderatoly concave (second pore to setae st. 2).
Line formod by end of genital flap rib3 usually slightly but
distinctly arched. Setae Jvl off genital shield on soft integu
ment, but sometimes touching margin of shield, Paranal setae 129
positioned at level near middle of or slightly posterior to
middle of anal field, Peritremo subequal in width, usually
slightly wider posteriorly! extending to anterior half of
coxa XIj normal in appearance throughout. Greatest width of
dorsal shield at level of Zl setae j lateral sides straight
or nearly so and converging slightly toward anterior end|
posterior end forming broad, blunt to narrow, sharp wedge.
All dorsal setae present and normally developed. Distance
between setae Zl and 31 slightly greater than length of Zl.
Setae Z5, S5» and sometimes slightly barbed. Dorsal
shield setae subequal in length with central dorsal setae
slightly smaller. Coxal spur formula 0-2-2-lj inner spur on
coxae II and III moderate in size and rounded to almost
truncate; outer spur on coxa III smaller and acutej spur on
coxa IV moderate in size and acute. Setae av^ and pv^ on
tarsus II stout and claw-like j seta av-^ on tarsus IV bluntly
spiniform. All othor log setae present and normal,
Colloction Records
Eutamlas minimus (Bachman)
UTAHi Box Elder Co., Lynnj July 1957J (Allrod and Beck, 19&6,
described as H. punctatus).
Eutamias quadrlvlttatus (Say)
UTAH: Summit Co., Bald Ktn.j Collt D E, Beckj 25 August 195^1
2 females (DI'A)i same locality; Coll: D. K. Allred and M. L. 130
KiUpackj 8 August 1957J 1 female (DMA), (Allred and Beck, 1966,
described as H. punctatus).
♦Tamjasclurus hudsonicus richardsoni (Bachman)
*MONTANAi Missoula Co,, Loloj Colli W, V, King} 22 June 1910}
2 females (Lectotype and cotype) (US]JM)» (Ewing, 1923).
Glaucomys sabrinus (Shaw)
UTAH: D a g g e tt C o ., Deep Creek} 29 June 1958} 1 fem ale (DMA),
(Allred and Beck, 1966, described as H. punctatus),
Peromyscus maniculatus (Wagner)
UTAHi Kano Co., Cottonwood Creek} June 1953} (Allred and Beck,
1966, described as H. punctatus).
Type M aterial
The typo of H. occidentalis was originally based on "...the one of two contained females, which is mounted dorsal side u p ,.,.", and collected from Tamiasciurus hudsonicus richardsoni at Lolo, Missoula
Co., Montana, 22 June 1910 by W. V, King, Subsequently those two
femalo specimens were remounted on separate slides with no designation
of the primary typo, thus they have since been considered cotypes. The
one cotype which is in best condition was remounted in 1967, and is
herein designated the lectotype and this is indicated on the slide
l a b e l . 131
Hirstionyssus (H.) eutamiae Allred and Bock, 1966
(Figs. 51-5*0
Hirstionyssus eutamiae Allred and Beck, 1966, Brigham Young Univ. Sci.
Bull,, Biol. Ser. 8(l):21-22 [Typej Aspen Grove, Utah Co.,
Utah; Holotype, Dept of Zool. and Entomol., Brigham Young Univ.,
Provo, Utah] .
D ia g n o sis
Deutosternal denticles 15 - 16 in number; arranged in alternating
double file. Anterior margin of sternal shield slightly to mod
erately convex; posterior margin moderately concave (second pore
to setae st. 2). Line formed by end of gonital flap ribs
slightly but distinctly arched. Setae Jvl off genital shield
on soft integument and usually removed from margin of shield by
distance greater than setal base diameter. Paranal setae
positioned at level near middle of anal field. Perltreme sub-
oqual in width, usually slightly wider posteriorly; oxtonding
to anterior half of coxa II; normal in appoaranco throughout.
Greatest width of dorsal shield at level of s6 or Zl setae;
lateral sides straight to slightly convex and converging toward
anterior ond; posterior end forming broad, blunt to narrow, sharp
wedge. All dorsal setae present and normally devolopod. Dis
tance between setae Zl and SI slightly greator than length of Zl.
S e ta e Z5 and S$ slightly barbed. Dorsal shiold setae subequal in
length with central dorsal setae slightly smallor, Coxal spur formula 0-2-2-1; inner spur on coxae II and III small and
usually bluntj outer spur on coxa III moderate in size,
slender, and acute; spur on coxa IV very small and acute*
Setae av-^ and pv^ on tarsus II stout and claw-like; seta
av^ on tarsus IV acutely spiniform. All other leg setae
present and normal.
Collection Records
Eutamias quadrivittatus (Say)
UTAH: Utah Co., Provo, Rock Canyon; Coll: R. K. Utley; 19 October
1956; 1 female (DMA); same Co., Schofield Resevoir Canyon; Colli
D E, Reck; 7 Juno 1957f 1 female (DMA), (Allred and Reck, 1966 ) .
Eutamias tovmsendii (Bachman)
CALIFORNIA: El Dorado Co., Echo Summit; Coll: E. '•[, Jameson, Jr.;
June 1962; 2 females (EMJ),
*Eutamias sp,
*UTAH: Utah Co., Aspen Grove; Coll: L. Tseu; 16 October 195&I
1 female (Holotype) (DMA), (Allrod and Beck, 1966).
Roithrodontomys mogalotis (Baird)
UTAH: Utah Co., Cedar Valley; November 1952; (Allrod and Beck, 1966 ). 133
Hirstionyssus (H.) cynomys (Radford, 1941)
(Figs. 55-58)
Liponyssus cynomys Radford, 1941, Parasitol. 3^(3) :313~314 [Typei
Antonito, Conejos Co., Colo.j Holotype, B rit. Mus. (Natur.
H i s t , ) ] \ Radford, 1943, Parasitol. 35*64-65 (catalogue).
Hirstionyssus cynomys, Fonseca, 1948, Proc. Zool. Soc. London
118(2) :297 (synon.); Strandtmann, 1958, J. Kansas Entomol.
Soc, 29(4)i138 (descr. note); Strandtmann and Wharton, 1958,
Inst. Acarology Contrib. 4:103 (synon., host, d istr,),
Neoichoronyssus cynomys, Jameson, 1950, Proc. Entomol, Soc. Washington
52(4)t162-163 (key).
Hirstionyssus occidentalis (Ewing, 1923), Strandtmann and Morlan, 1953,
Texas Rep. Diol. Med. 11(4) 1632 , 636 - 63 ? (key, figs., host,
d istr.), synonymy subsequently retracted by Strandtmann, 1956,
J. Kansas Entomol. Soc. 29(4):138, and Strandtmann and Wharton,
1958, Inst. Acarology Contrib. 4 j 1 0 3 .
Hirstionyssus invagjnatus Allred and Beck, 1966, Brigham Young Univ.
Sci. Bull,, Biol. Ser. 8(l):2l-22 [Type: Provo Canyon, Utah
Co., Utah; Holotype, Dept, of Zool. and Entomol., Brigham Young
Univ., Provo, Utah] . NEW SYNONYMY.
Hirstionyssus angustus Allred and Beck, 1966, Brigham Young Univ. Sci,
Bull., Biol, Ser. 8(l);24-25 [Type: Colton, Utah Co., Utah;
Holotype, Dept, of Zool. and Entomol., Brigham Young Univ.,
P ro v o , U ta h ] . NEW SYNONYMY. 13^
D ia g n o sis
Deutosternal denticles 1^ - 16 in number; arranged in alternating
double file. Anterior margin of sternal shield slightly convex;
posterior margin usually deeply concave (to or beyond setae
st. 2), Line formed by end of genital flap ribs slightly but
distinctly arched. Setae Jvl off genital shield on soft integu
ment, but sometimes touching margin of shield, Paranal setae
positioned at level near middlo of or distinctly posterior to
middle of anal field, Feritremo subaqual in width, usually
slightly wider posteriorly; extending to posterior half of
coxa I; normal in appearance throughout. Greatest width of
dorsal shield variable from setae s3 — s4- to Zl - Si; lateral
sides straight or nearly so and parallel; postorior end in shape
of broad, blunt to narrow, sharp wedge, AH dorsal setae present
and normally developed. Distance between setae Zl and SI much
greater than length of Zl, Setae Z5 and sometimos S5 slightly
barbed. Central dorsal shield setae somewhat smaller than
anterior, lateral, and posterior marginal sotae, Coxal spur
formula 0-2-2-1; innor spur on coxae II and III largo and acute;
outer spur on coxa III moderate in size and acute; spur on
coxa IV short, broad, and acute. Sotae av^ and pv^_ on tarsus II
stout and claw-like; sota av^ on tarsus IV bluntly spiniform.
All other leg setae present and normal. 135
Collection Records
*Cynomys sp,
*COLORADO: Conejos Co., Antonitoj Coll: G, M. Kohls; 7 November
1932; 1 female (Faratype) (CDR), (Radford, 19^1).
Cltellus armatus (Konnicott)
UTAH: Box Elder Co., Goose Creek Ktns.; Coll: R, K. Hansen;
9 Kay 1952; 1 female (DMA); Cache Co., Blacksmith Fork Junction;
Juno 1953; Rich Co., Laketcwn; June 19531 same Co., Randolph;
17 July 1951; 2 females (DMA); San Fete Co., E. Mt. Pleasant;
Coll: D E. Beck; 1 August 1951; Summit Co., Blacks Fork; Coll:
R. K, H ansen; 29 J u ly 1952; 1 fem ale (DMA); U tah C o ., Provo
Canyon; Coll: D B, Back; 2 6 June 1951» duly 1957; ^ females (DMA);
same Co., Spanish Fork Canyon; Coll: D E. Beck; 7 July I960;
1 female (DMA); same Co., Soldiers Summit; Coll: D E, Beck;
7 June 1951» 1 female (DMA); Wasatch Co., Current Creek; Coll:
D B. Beck; 30 June 1953; 2 females (DMA); came locality; Coll:
R. M. Hansen; 2 August 1952; ^ females (DMA); same Co., W, Lake
Creek Summit; Coll: D E. Beck; 7 August 1953; 3 females (DMA);
same Co., W, Wolf Creek Summit; Coll: M. D. Coffey and M. L.
K illp a c k ; 6 August 1953; 1 female (DMA); same Co., Strawberry
Resevoir; June 1951; (Allred and Beck, 1966, described as
H. invaginatus and H. angustus).
"Wyoming ground squirrel"
COLORADO: Grand Co., Kremling; Coll: N. B. Kiqutoff; 13 April
1939; 1 female (RWS). Hirstionyssus (H,) utahensis Allred and Bock, 1966
(Figs. 59-62)
Hirstionyssus obsoletus, Jameson (in part), 1950» Proc* Biol, Soc.
Washington 63:32 (records from Peromyscus maniculatus and one
record from Neutrotrichus gibbsii only)} Strandtmann and Morlan,
(in part), 1953» Texas Rop. Biol. Med. li(4):630 (key, figs.,
host, distr.).
Hirstionyssus carnifex, Strandtmann and Korlan (in part), (not Koch,
1839), 1953* Texas Rep, Biol, Med, 11(4):632 (key, figs,, host,
d istr.)j Strandtmann and Wharton (in part), 1958* Inst.
Acarology Contrib. h t101-102 (synon., host, d istr.).
Hirstionyssus occidentalis, Strandtmann and Morlan (in part), (not
Ewing, 1923)f 1953f Texas Rep. Biol. Mod. ll(*J-):632 (key, fig s.,
host, distr.),
Hirstionyssus utahensis Allred and Beck, 1966 , Brigham Young Univ. Sci.
Bull,, Biol, Ser, 8(l):22-2^ ^Typo: E. Bald Mtn., Summit Co.,
Utahj Holotype, Dept, of Zool. and Entomol,, Brigham Young Univ.,
Provo, Utah J .
Hirstionyssus tarsalls Allred and Beck, 1966, Brigham Young Univ. Sci.
Bull., Biol. Ser. 8(1):18-19 [Type: Deep Creek Camp Ground,
Daggett Co., Utah} Holotype, Dept, of Zool. and Entomol,,
Brigham Young Univ., Provo, Utah J . NEW SYNONYMY. 137
D ia g n o sis
Doutosternal denticles It - 22 in number ; arranged in alternating
double file. Anterior margin of sternal shield slightly convex;
posterior margin moderately concave (second pore to setae st. 2),
Line formed by end of genital flap ribs irregularly straight to
slightly arched. Setae Jvl off genital shield on soft integu
ment, but touching margin of shield, Paranal setae positioned
at level near middle of or slightly posterior to middle of anal
field, Peritreme suboqual in width, usually slightly wider
posteriorly} extending to anterior half of coxa I; normal in
appearance throughout. Greatest width of dorsal shield usually
at level of Zl - SI setae; lateral sides straight to slightly
convex and approximately parallel; posterior end in shape of
broad, blunt wedge. All dorsal setae present and normally
developed. Distance between setae 21 and Si much greater than
length of Zl. Setae Z5 and sometimes S5 slightly barbed. Kost
dorsal shield setae subequal in length but with j2, j3» z2, and
si distinctly longer, Coxal spur formula 0-2-2-1; inner spur
on coxa IT small and acute to blunt; inner spur on coxa III
moderate in size and acute; outer spur on coxa III short, broad,
and acute; spur on coxa IV moderato in size, short, broad, and
acute. Setae av^ and pv^ on tarsus II stout and claw-like; seta
av-^ on tarsus IV slightly enlarged basally but not distinctly
spiniform. All other leg setae present and normal. Collection Records
Sorex trowbridgii Baird
CALIFORNIA: Plumas C o ., Q uincy; C o lli E . W. Jam eson, J r . |
20 October 19^91 1 female (RW3), (Jameson and Brennan, 1957)»
Neutrotrichus gibbsii (Baird)
CALIFORNIA! Plumas C o ., Q uincy; C o llt E . 17. Jam eson, J r . |
20 April 19^9; 1 female (RWS), (Jameson and Brennan, 1957).
Cite11ns armatus (Konnicott)
UTAHi San Pete Co.; Summit Co.; Utah Co.; Wasatch Co.; (Allrod
and Bock, 1966),
Citellus tridecenllnoatus (H itchill)
UTAH: Duchesne Co., Roosevelt; Coll: M. L. Killpack; 19 April
1953l 1 fem ale (DMA), (A llr e d and R eck, 1 9 6 6 ).
Citellus lateralis (Say)
UTAH: Beaver Co., Dolano RaJigor Station; Colli D E. Beck|
25 June 1957; 1 female (DMA); Carbon Co.; Summit Co.; Utah Co.;
(Allred and Bock, 1966).
Eutamias minimus (Bachman)
UTAH: San Juan Co,; Sevier Co.; Summit Co., Bald Ktn; Colli
D E. Deck; 2U- A ugust 1956; 1 fem ale (DMA); D uchesne C o ., R o o s e v e lt;
Coll: D E, Beck; 23 August 1952; 1 female (DMA), (Allrod and Beck, 1966 ),
*Eutamias quadrivittatus (Say)
*UTAH: Daggett Co.; Duchesne Co.; San Juan Co.; San Pete Co.;
Sevier Co., W. Koosharem; Coll: D E, Beck; 20 July 1953; 1 female 139
(DM)j Summit Co., E. Bald Mtn. ) Collr D. M. Allred and M, L.
K illp a c k j 8 August 1957J 3 females (Holotype and paratypes) (KiA)f
Utah Co.} (Allred and Beck, 1966),
Eutamias sp.
UTAH: Wayne C o ., P le a s a n t C reek) C o lit D. H, A llre d ) 7 June
I960) 1 female (DKA), (Allred and Beck, 1966).
Thomomys talpoidos (Richardson)
UTAH: Utah Co.) (Allred and Beck, I 966 ).
Porognathus longimembrls (Couos)
UTAH: Kane Co.) (Allred and Bock, I 966 ),
Dipodomys ordii Woodhouse
UTAH: Box Elder Co., Lucin; 23 Hay 1953) 1 female (DM)) Uintah
Co., Jonsen; Coll: D E, Beck) 9 June 1953 i 2 females (DMA),
(Allred and Beck, 1966)} San Juan Co.) Tooele Co.) Utah Co.)
(Elzinga, 196h, m isidontified as H. Occident a lls) Allred and
Beck, 1966),
Pipodomys mlcrops (Kerriam)
UTAH: Tooele Co.; Utah Co.) (Elzinga, 196h, m isidontified as
H. occidentalis) Allred and Bock, 1966 ),
Reithrodontomys mogalotis (Baird)
UTAH: Utah Co, | November 1958 ~ March 1959) (Elzinga, 19&J-)
Elzinga and Roes, 196,+ , m isidentified as H. occidontalls))
Carbon Co., Price) Coll: D E. Beck) 8 June 1951) 1 female (DMA),
(Allred and Beck, 1966), Peromyscus crlnltus (Kerriam)
NEVADA! Hyo C o ., M ercury, AEC-NTSj 10 June 1961; 1 fem ale (DMA),
(Allred and Goates, 1964b, misidontified as H. carnifex).
UTAH: Kane Co., A dairville; 11 June 1958* 2 females (DKA)t
Tooele Co., Granite Mtn.| Colli E, A. Shippej 1, 7 April 1953*
2 females (DM), (Allred and Beck, 1966); San Juan Co. | Wayne Co.|
( A llr e d , 1957t>» m is id o n tif ie d a s H. o c c i d e n t a l i s ) .
Peromyscus eremicus (Baird)
MEXICO: Baja, San Quinton| Colli F. J. Radovsky; 16 February
1965* lab reared female progeny (FJR).
UTAH: Washington Co., Beaver Dam Wash; 16 April 1952} 1 female
(DMA), ( A llr e d , 1957b, m i s i d o n t i f i e d a s H. o c c i d e n t a l i s j A llro d
and Beck, 1966).
Peromyscus maniculatus (Wagner)
CALIFORNIA: Plumas C o ., Q uincyj C o ll: E , W, Jam eson, J r . |
23 February, 5, 12 April, 8, 14, 17 Juno, 16 September 1949,
17 July 1951, 14 females (RW5 and EWJ), (Jameson and Brennan,
1 9 5 7 ).
COLORADO: Montezuma Co., Mesa Verde Hat. Monument; 2 females (DMA).
IDAHO: Butte Co., Idaho Falls, AEC-MRTSj 11, 12 June, 17, 21 July
1966, 29 June 1967s 17 females (DM).
MICHIGAN: Cheboygan Co., Douglas Lake areaj Coll: W. J. Wrenn*
30 June, 11 July 1965s 2 females (03U).
MEW MEXICO: Santa Fo Co., Santa Fej Coll: H, B, Korlan; 30 August
1951; 1 female (RWS). OREGON: Harney Co., Steen's Ktns.* Colli C, G. Hansen*
19 A ugust 195^! 1 fe m ale (USNK)» (H anson, 196h m i s i d e n t i f i e d
as H. o b s o le t u s ) .
UTAH: throughout statej September 19^8 - August 19531 (Allred,
1957b, misidontified as H, occidentalis); Box Elder Co., Lynn*
Coll: D. K, Allred* 11 July 1957* 1 female (DMA)» (Allrod and
Bock, 1966, described as H. tarsalis); Garfield Co., So. Torey*
Coll: D. M. Allred* 22 June 19^3* 1 female (DMA)* Juab Co., Joy*
Coll: D. M, Allred elt al, * 1 June 1951* 1 female (DKA)j Kano Co.,
Cottonwood Creek; 10 June 1953* 5 females (DMA)* Piute Co.,
Kingston* Coll: D E. Beck et a l.* 26 June 1952* 1 female (DMA),
(Allred and Beck, 1966)* Tooele Co., Tooele; 15 May 1952*
6 females (DMA)* same locality* 20 I -lay 1952* (Keegan, 1953*
m isidentified as H. carnifex); Utah Co.* November 1953 - March 1959*
(Elzinga, 196h5 Elzinga and Rees, 196h, m isidontified as H,
occidentalis); same Co., Chimney Rock Pass* Coll: M. D, Coffey
et^ a l.* 28 March 1953* 13 females (DMA); same Co., Pleasant Grove*
Coll: D. Ash* 12 August 1961* 1 female (DMA).
Foromyscus loucopus (Rafinesque)
MICHIGAN: Cheboygan Co., Douglas Lake area* Coll: ’-‘J. J, Wrenn *
20 July 1965* ^ females (OSU).
OHIO: Erie Co., Castalia .Voods; Coll: R. J, Stoll; 29 Juno -
11 A ugust 1965* 6 fe m a le s (OSU), 1*4-2
Peromyscus boylil (Baird)
CALIFORNIA: Plumas C o ., Q u in cy ; C o ll: E . W. Jam eson, J r . |
19 November 19*1-8, 17 September 19*4-9» 29 July 1951J 3 females
(RWS and EWJ).
Peromyscus truei (Shufeldt)
UTAH: Kane Co.; San Juan Co.) Wayne Co.; Tooele Co.; (Allred
and Eeck, I 966 ),
Feromyscus sp.
IOWA: Boone Co.j Colli F, E. French; 13 July I960; 2 females
(RWS).
Neotoma lopida lepida Thomas
UTAH: Juab Co., NE Jericho; April 195^; (Howell, Allred, and
Bock, 1957t m isidentified as H. occidentalis); Kane Co., NE
A dairville; Coll: D. M, Allrod; 11 June 1958* 1 female (DMA);
Tooele Co., Granite Ktn,; Coll: E. A. Shippe; 7 April 1952;
1 fem ale (DMA), (A llr e d and B eck, I 966 ).
Hirstionyssus (H.) latiscutatus (de Keillon and Lavoipierro, 19*44)
(Figs. 63-65)
Ichoronyssus caraifex, Ewing (not Koch, 1839)* (1923), Proc, U. S.
Hat. Mus. 62(13)-.1*4-15.
Liponyssus latiscutatus de Keillon and Lavoipierre, 19*4-4, J. Entomol.
Soc. South Africa 7*62 £ Type t Newtown, Johannesburg, Transvaal,
So. Africa; Holotypo,?} . 1^3
Ichoronyssus orcadensls Turk, 19^6, Ann. Kag. Natur. Hist. ll(12)i
796 £ Type 1 Zoological Gardens, Regent's Park, England^
Holotype, probably private collection of F. A. Turk, Camborne
synonymy by Evans and T ill, 1966, Bull
B rit. this. (Natur. H ist.) Zool. lh(5)*278-230, 291.
Hirstionyssus latiscutatus, Fonseca, 19^* Proc. Zool. Soc. London
118(2);298 (synon.)i Strandtmann and Korlan, 1953* Texas Rep.
Biol. Fed, ll(h-):632 (key, figs., host, d istr.)j Strandtmann
and Wharton, 1958, Inst, Acarology Contrlb. h-:105 (synon.,
host, d istr,)t Evans and T ill, 1966, Bull. Brit. Mas, (Hatur.
H ist.) Zool. 1^(5)*278-230, 291-295 (synon., key, descr.,
host, distr.).
Hirstjonyssus arcuatus, Strandtmann and Korlan (in part) (not Koch,
1339)» 1953* Texas Rep. Biol. Med. 11(4):632.
Hirstionyssus carnifex, Strandtmann and Korlan (in part) (not Koch,
1839)» 1953* Texas Rep, Biol, fed, 11 (4-) 1632 (host, distr. )|
Strandtmann and Wharton (in part), 1958, Inst. Acarology
Contrib, 4-: 101-102 (synon., host).
Hirstionyssus muscull, Bregetova (not Johnston, 18h9), 1956, Opred.
Fauno 3SSR. 6ltl35 (key, fig s.), synonymy by Evans and T ill,
1966, Bull. Brit. Kus. (Natur. H ist.) Zool. lh(5):273-280, 2911
Strandtmann and Wharton, 1958, Inst. Acarology Contrib, h:105
(synon., host, distr,). 144
Hirstionyssus invaginatus Allrod and Beck (in part), 1966, Brigham
Young Univ. Sci, Bull,, Biol, Ser. 8(1):21 (records from Kus
musculus nest only).
D ia g n o sis
Deutosternal denticles 14- - 22 in number; arranged in
alternating double file. Anterior margin of stom al shield
slightly concave to slightly convex; posterior margin
moderately to deeply concave, usually to level near setae
st, 2, Line formed by end of genital flap ribs strongly
arched. Setae Jvl off genital shield on soft integument,
but touching margin of shield, Paranal setae positioned at
level near middle of anal fiold, Peritromo subequal in width,
usually slightly wider posteriorly; extending to anterior half
of coxa X; normal in appearance throughout. Greatest width of
dorsal shield at level of setae Zl - SI; lateral sides straight
to slightly convex and converging toward anterior end; posterior
end forming broad, blunt to narrow, sharp wedge. All dorsal
setae present and normally developed. Distance between setae
Zl and SI much greater than length of Zl, Setae Z5 and S5
slightly barbed. Host dorsal shield setae suboqual in length,
but with J2, z2, z4, and several anterior s series setae some
what longer, Coxal spur formula 0-2-2-1; inner spur on coxae
II and III largo and acute; outer spur on coxa III moderate in
size and acute; spur on coxa IV long, slender, and acute. 145
Setae av-^ and pv^ on tarsus II stout, claw-like, but rather
short. All other leg setae present and normal.
Collection Records
Rattus norvegicus (Berkenhout)
GEORGIA: Decatur Co., Bninbridge; Colli H. B. Korlan; 14 February
1 9 4 6 j 1 fe m ale (USNM).
TEXAS: Yoakum Co., Plains; Coll: Conica; 14 Kay 1947; 1 female
(RWS).
Kus musculus Linnaeu3
MASSACHUSETTS: Middlesex Co., Cambridge; 4 females (OSU).
MISSISSIPPI: Leflore Co.; Marshall Co.; Montgomery Co.; 1 July
1949 ~ 30 June 1950* (Smith, 1955» m isidentified as H. cam ifex).
MISSOURI: Boone Co., Columbia; Coll: L. Baseman; 14 July 1930»
29 September 1937; 6 females (RWS),
OREGON: Benton Co., Corvallis; Coll: H. E, Ewing; 9 January
1912, 1 female (USHM), (Firing, 1923» m isidentified as H. cam ifex).
UTAH: San Juan Co., Kigalia Ranger Station; Coll: D E. Bock;
9 June 1955; 2 fe m a le s (DMA); U tah C o ., L o h i; C o lli D E . B eck;
1 January 19511 1 female (DM), (Allred and Beck, 1966, m isidonti
fied as H. invaginatus (= cynomys)).
N ote
In addition to the Nearctic distribution given above, H.
latiscutatus is recorded by many authors from a wide spectrum of
rodent hosts throughout the world. The type host and locality are Ih 6
Rattus rattus in Mewtown, Johannesburg, Transvaal, South Africa,
Type specimens wero unavailable for examination in this study. Speci mens from Mus muscuius in B ritain, determined by G, 0, Evans who has examined the typo m aterial, wore made available by tho B ritish Museum.
Hirstionyssus (H,) "A" n. sp.
( F ig s . 66 - 67 )
D ia g n o sis
Deutostornal denticles lh - 16 in number; arranged in
alternating double file, Anterior margin of sternal shield
slightly convex; posterior margin deeply concave (beyond
setae st. 2). Line formed by end of genital flap ribs
moderately arched. Setae Jvl off genital shield on soft
integument and usually removed from margin of shield by dis
tance greater than setal base diameter, Paranal setae
usually positioned at level slightly anterior to middle of
anal field, Peritreme subequal in width, usually slightly
wider posteriorly; extending to middle of coxa I ; n o rm al i n
appearance throughout. Greatest width of dorsal shield at
level of Zl - SI setae; lateral sides straight or nearly so
and converging slightly toward anterior end; posterior end
in shape of broad, blunt wedge. All dorsal setae present and
normally developed. Distance between setae Zl and SI greater
than length of Zl. Setae Z5, S5» Sty, S3, and J3 usually
slightly barbed. All dorsal setae, as well as ventral opisthogastric setae, subequal in length and unusually small.
Coxal spur formula 0—1—1—1; inner spar on coxae II and III
absent; although with indistinct, broad, slightly raised
apophyses in place of spurs; outer spur on coxa III moderate
in size and acute to blunt; spur on coxa IV moderate in size
and blunt. All leg setae present and normal.
Collection Records
* Cynomys ludoviclanus (Ord)
♦KANSAS: Rarber Co., SW Aetna; 16 September 1953J ^ females
(Holotypo and paratypes) (RWS).
Type M atorial
Eolotype fomale, vrith collection data as given above and collec tion number RL530916-3, deposited in tho U. S, National Kuseum. Para types in the collection of R. W. Strandtmann, Bernice P. Bishop IJusoum,
Honolulu, Hawaii, and in the collection of tho Acarology Laboratory,
The Ohio State University, Columbus, Ohio.
Hirstlonyssus (H.) torus Allred and Beck, 1966
(Figs. 68-70)
Hirstionyssus torus Allred and Bock, 1966, Brigham Young Univ. Sci.
Bull., Biol, Ser. 3(l);27-29 [Type: Handing, San Juan Co.,
Utah; Eolotype, Dept, of Zool, and Entomol,, Brigham Young
Univ., Provo, Utah] . 1^8
D ia g n o sis
Deutostemal dentlclos 11 - 13 in number; arranged in
alternating double file. Anterior margin of sternal shield
slightly convex; posterior margin moderately concave (second
pore to setae st, 2). Line formed by end of genital flap
ribs irregularly straight to slightly arched. Setae Jvl
off genital shield on soft integument, but often touching
margin of shield, Paranal setao positioned at level near
middle of or slightly anterior to middle of anal field.
Feritreme suboqual in width, usually slightly wider posteriorly;
extending to middle of coxa I; normal in appearance throughout.
Greatest width of dorsal shield usually at level of setao Zl -
SI; lateral sides straight to slightly convex and converging
toward anterior end; posterior end in shape of broad, blunt to
narrow, sharp wedge. All dorsal setae present and normally
developed. Distance between setao Zl and SI much greater than
length of Zl, Setao Z5» S5» and S^J- slightly barbed. Dorsal
s e ta e jk, j5, j6, z5, z6, Jl, J2, J3, Zl, SI, S2, 33, and
very small, with all others moderate and suboqual in length,
Coxal spur formula 0-2-2-1; inner spur on coxa II very broad
and rounded; inner, spur on coxa III large and acute; outer spur
on coxa III moderate in size and acute; spur on coxa IV moderate
in size, slender, and acuto, Setae av^ and pv^ on tarsus II
normal, long and slender; seta av-^ on tarsus IV acutely spiniform, although not extremely so. All other log setae
present and normal.
Collection Records
*Sclurus aborti Woodhouse
COLORADO* Larimer Co., Livermore; Coll: S. C, McCampboll;
3 O cto b er 1926; 3 f o n a le s (USMI')» ( m is id e n tif ie d b y H. E, Ewing
as R. occidentalis).
♦UTAH: San Juan Co., Mo. Blending; Coll: D. K. Allrod and D E.
Bock; 6 Kay 1951 1 ^ females (Holotype and paratypes) (DMA),
(Allred and Beck, 1966).
Hirstionyssus (H.) obsolotus Jamoson, 1950
(Figs. 71-72)
Hirstionyssus obsoletus Jameson, 1950, Froc. Biol, Soc, Washington
63:32-3^ [Typo: Plumas Co., C alif.; Holotypo, U. S. N. M.] ;
Strandtmann and korlan, 1953» Texas Rep. Biol. Med. 11 (U-) t630
(key, figs., host, d istr.); Strandtmann and Wharton, 195^,
Inst. Acarology Contrib. hjl06 (synon., host, d istr.).
Mooichoronyssus obsolotus, Jameson, 1950, Proc. Entomol, Soc. Washington
52(^)il52-153 (key).
D ia g n o sis
Doutosternal donticlos 15 in number; arranged in altornating
double filo. Anterior margin of stornal shield slightly con
vex; posterior margin usually moderately concave (second pore to setae st, 2), although sometimes more dooply concave.
Lino formed by end of genital flap ribs strongly arched.
Setae Jvl off genital shield on soft integument, but touch ing margin of shield. Genital shield quite broad throughout with lateral sides straight and nearly parallel. Anal shield very broad relative to length; with pair of lateral pores; paranal setae usually positioned at level slightly posterior to middle of anal field. Feritremo subequal in width, usually slightly wider posteriorly; extending to anterior half of coxa I; normal in appearance throughout. Greatest width of dorsal shield variable, from level of setae s3 - s4 t o Zl - SI; lateral sides straight to slightly convex and approximately parallel; posterior end broadly rounded. Dorsal
setae SI usually absent or removed from shield and located on
soft intogument, although sometimes may be present and in
normal position on dorsal shield margin. All other dorsal
setae present and normally developed. Setae Zl removed from
setae 51 or from margin of shield by distance greater than
length of Zl. Setae Z5 and S5 slightly barbed. Dorsal setao
subequal in length, except for setae j2, j j , z2, zh, and
which are decidodly longor; central dorsal setae slightly
shorter than marginal setao. Coxal spur formula 0-2-2-0;
inner spur on coxae II and III moderate in sizo and broadly
rounded; outor spur on coxa III moderate in size and acute to
blunt. All log setao present and normal. 151
Collection Records
♦Sorex trowbrldgii Baird
♦CALIFORNIA! Plumas Co., Quincy; Coll: E. W. Jameson, Jr. ;
10 March 19^91 8 females (Paratypos) (USHM), (Jamoson, 1950a)j
same locality and collector; 29 December 1948, 9, 11, 12 March
1949; 4 females (EWJ) (Jameson and Brennan, 1957)«
OREGON: Benton Co,, S17 Philomath; C olli J, Wernz; 1 March 19&7I
2 fe m a le s (GWK).
Scapanus townsondii (Bachman)
OREGON: Benton Co.; Coll: J. F. Comoll and D. L, Kaysj 19 May
1966; 1 fem ale (GWK).
Hirstionyssus (H. ) isabollinus (Oudemans, 1913)
(Figs. 8-11, 73-7*0
Liponyssus isabollinus Oudemans, 1913, Ent. Ber. Amst. 3:384, and
Arch. Naturgesch. Abt. A 79(9) :S0 £ Type: Holland; Holotypo,
Leiden Museum, Leiden, Holland J .
Ichoronyssus isabollinus, Ewing (1923), Proc. U, S. Hat, Mus. 62(13):
15-16; Radford, 1943* Parasitol. 35:66 (catalogue).
Hirstionyssus Isabollinus, Fonseca, 1943, Proc, Zool. Soc. London
113(2):297 (synon.); Strandtmann and Morlan, 1953* Texas Rop.
Biol, Med. 11(4):630 (key, figs., host, d istr.); Strandtmann
and Wharton, 1958, Inst. Acarology Contrib. 4:104-105 (synon.,
host, d istr.); Allrod and Bock, 1966, Brigham Young Univ. Sci,
Bull., Biol, Sor. 8(l):29 (key, figs., host, distr.); Evan3 and 152
T ill, 1966, Bun, Brit, Kus. (Natur. H ist.) Zool, 1^(5) 3^95“
296 (descr,, figs., synon,, host, distr,),
Hirstionyssus arvicolae Zemskaya, 1955» in Brogotova et a l,, Opred,
Faune SSSR. 59:366, synonymy by Brogotova, 1956, Opred,
Faune 5SSR. 61*181,
D ia g n o s is
Doutosternal denticles 12 - 20 in number; arranged in
alternating double file. Anterior margin of sternal shield
slightly concave to slightly convexj posterior margin
moderately concave (second pore to setae st, 2), Line
formed by end of genital flap ribs strongly arched. Setao
Jvl off genital shield on soft integument, but often touch
ing margin of shield. Paranal setae positioned at level
noar middle of or slightly posterior to middle of anal field.
Poritreme suboqual in width, usually slightly wider posteriorly \
extending to anterior half of coxa I; normal in appearance
throughout. Greatost width of dorsal shield usually at level
of Zl - SI setao| lateral sidos straight to slightly convex and
gradually converging toward anterior ondj posterior end in shape
of broad, blunt to narrow, sharp wedge. All dorsal setae present
and normally dovolopod. Distance betwcon sotae Zl and SI more or
less equal to length of Zl, Sotae Z5 and S5 slightly barbed.
Dorsal shield setao subequal in length; with anterior, lateral,
and posterior marginal sotae somewhat longer, Coxal spur formula
0-2-2-0; inner spur on coxae II and III largo and acuto; outer spur on coxa III moderate in size and acute. All leg setae
present and normal*
Collection Records
Ochotona princops (Richardson)
DTAH: Summit Co., E, Bald Mtn.j Colli D, K. Allredj 8 August
19571 ^ fe m a le s (DMA), ( A llre d and R eck, 1 9 6 6 ),
Citellus armatus (Konnicott)
UTAH: Rich Co.; (Allred and Reck, 1966).
Eutamias quadrivittatus (Say)
UTAH: Utah Co,; (Allred and Rock, 1966).
Thomomys talpoides (Richardson)
UTAH: Carbon Co., So. Schofield; Colls D E. Rock and C. Pritchett;
30 J u ly i9 6 0 ; 1 fem ale (DI1A); U tah C o .j (A llr e d and R ock, 1 9 6 6 ).
Reithrodontomys megalotis (Baird)
UTAH: Utah Co., Lehi; Coll: D, M. Allrod; 4 February 1951 t
2 fe m a le s (DMA), ( A llre d and Beck, 1966 ),
Peromyscus maniculatus (Wagner)
IDAHO: B u tte C o ., Id ah o F a l l s , AEC-URTS; 11 Juno 1966; 1 fe m a le
(DMA).
UTAH: Garfield Co.; (Allred, 1957b); Kano Co.; (Allred and Beck,
1966 ).
Onychomys torridus (Coues)
UTAH 1 Washington Co.; (Allred and Bock, 1966 ). Leinus trimucronatus (Richardson)
CANADA.: Northwest Territory, Prince Patrick Island, Mould Bayj
Coll: C. 0, Handley; June, i August, 5 October 19^9* 20 females
(USNM)* Hudson Bay, Southhampton Island; Coll: H, B. Collins;
30 June 195^1 3 females (USNM).
ALASKA: Pt, Barrow; Coll: Q. Tomick; 31 July 19531 1 female
(EWJ); Nunivak Island; Coll: B. Bahero; August 1951* (Keegan
and Heeden, 1952),
Clothrionomys rutilus (Fallas)
CANADA: Northwest Territory, NW Mackenzie, Reindeer Station;
Coll: R, Behin; August 1963* (M itchell and Behin, 1965).
Clothrionomys sp.
CANADA: Quebec, Lake Maryroac; Coll: F. Harper; 28 July 1953*
5 fe m ale s (USNM).
Phenacomys intormcdius Korriara
CANADA: Manitoba, Ft. Churchill; Coll* B. Foster; 1 Juno 1956*
8 fo m a le s (USNM),
UTAH: Summit Co.; (Allred and Deck, 1966).
Microtus ponnsylvanicus (Ord)
CANADA* Ontario, So, Bay, Manitoulin Island; Coll: J, Budd and
D, E. Johnston; 22 September 1955* 1 female (OSU).
MICHIGAN* Emmett Co., Fellston Road; Coll: W, J, V/renn* 23 July
1965 * 2 females (OSU); same Co., SW Alanson; 29 July 1965*
2 females (OSU). 155 NEV YORK i Albany Co., Rensselaerville | Collj F. Harper;
21 O cto b er 19^9; 1 fem ale (USNM).
OHIO: Erie Co., Venice; Coll: R, J. Stoll; 1 July - 23 August
19^5l 95 females (OSD).
UTAH: Utah Co., Provo; Coll: D, K, Allred and J. Murphy; 2h Kay
19511 1 fem ale (DMA), (A llr e d and Beck, 1 9 6 6 ).
Kicrotus montanus (Peale)
CALIFORNIA: Plumas C o ., Q uincy; C o ll: E. '•/, Jam eson, J r . ;
22 March 19^9; 1 female (RWS).
OREGON: Kalamath Co., Geory Ranch; Coll: E. C. Hanson; 17 June
I960; 1 female (OSU); Harney Co., Steen's Htns,; Coll: C, G.
Hansen; 29 June 195^1 1 female (GWK), (Hanson, 1965 ).
UTAH: Utah Co., Provo, Canyon; Coll: D. K. Allred; 22 July
1956; 1 fem ale (DMA), ( A llre d and B eck, 1966 ).
Kicrotus longicaudus (Korriam)
CALIFORNIA: Plum as C o ., No. S i e r r a Nevada K tn s .; O cto b er 19^3 -
August 1951? (Jameson and Bronr.an, 1957)•
OREGON: Harney Co., Steen's Ktns,; Coll: C. G. Hansen; 1*4- August
195^! 1 female (USNM), (Hanson, 196^),
UTAH: Duchesne Co., Pole Crook Mtn.; Coll: K, L. Killpacki
2h August 1957; 1 female (DMA); San Juan Co.; (Allred and Beck,
1966)( Summit Co., Bald Mtn.; Coll: D. M. Allred et al.;
25 August 1956; 1 female (DMA); Carbon Co., Schofield Rosevoir;
7 June 1957; 1 fem ale (DMA), (A llre d and Bock, 1966 ). . 156
Hicrotus sp.
ALASKA: Umita; Coll: L. A. Jachowski; 6-25 June 19^7I 13 females
(USNM)I Big D e lta j C o ll: C, 3 . P h i l i p s ; 19 June 1951* (Keegan and
Heeden, 1952).
PENNSYLVANIA: Beaver Co., Industry* Coll: N. D. Richmond and
J. J. Christianj 13 May 19^3* 1 female (RWS).
UTAH: Morgan Co., Morgan; Coll: D E. Beck and son; 12 July 1952;
1 fe m ale (DMA), (A llre d and Beck, 1966 ).
Mus m usculns L in n aeu s
MICHIGAN: Ingham Co., Lansing; Coll: T. Tibbetts; 20 October
19^9» 1 fem ale (USNM).
"unknown host"
UTAH: Tooele Co.; (Allred and Pack, 1966).
N ote
In addition to tho Nearctic distribution given above, H. isabellinus is recorded by many authors from a very wide spectrum of rodent hosts and several carnivore hosts throughout the Palearctic and
Oriental regions. The typo host and locality are Mustela sp. in
Holland, Type specimens were unavailable for examination in this study.
Representative specimens, identified as H. isabellinus by reliable workers, from several hosts and localities throughout its distribution were examined. 157
Hirstionyssus (H. ) brevisota Strandtmann and Korlan, 1953
(Figs, 75-76)
Hirstionyssus breviseta Strandtmann and Korlan, 1953* Texas Rep, Biol,
I'od, 11(4):629-635 [Typo: Mastin Ranch, Santa Fo Co., New
Mexico; Eolotype, U. S, N, M,} ; Strandtmann and VJharton, 1959,
Inst, Acarology Contrib, 4:101 (synon,, host, d istr.).
D ia g n o sis
Doutosternal denticles 12 - 14 in number; arranged in alternating
double file. Anterior margin of sternal shield straight to
slightly convex; posterior margin usually moderately concave
‘ (second pore to setae st. 2), although sometimes only slightly
concave (st, 3 setae to second pore). Line formed by end of
genital flap ribs Irregularly straight to slightly archod. Setae
Jvl usually on postero-lateral margin of genital shield. Anal
shield quito long and narrow; paranal setae positioned at level
near middle of or slightly posterior to middle of anal field.
Feritreme subequal in width, usually slightly wider posteriorly;
extending to middle of coxa I; normal in appearance throughout.
Greatest width of dorsal shield variable, from setae s3 to Si;
lateral sides straight to slightly convex and approximately
parallol; posterior end broadly rounded to shape of broad, blunt
wedge. All dorsal sotae present and normally developed. Dis
tance between setae Zl and SI much greater than longth of Zl.
Setae Z5 and S5 slightly barbed. Central dorsal shield setao 158
small! with anterior, lateral, and posterior marginal setao
subequal and moderate in length. Coxal spur formula 0-1-2-1;
inner spur on coxa II absent, although with indistinct,
slightly raised apophysis in place of spurj inner spur on
coxa III small and very acutely pointed* outer spur on
coxa III small and acutej spur on coxa XV small and acute#
AH leg setae present and normal.
Collection Records
*Ne otoma alblgula albigula Hartley
*MEW MEXICO: Santa Fe Co., Mastin Ranch; Coll: H. B. Korlan;
18 November 1952j 2 females (Holotypo and paratype) (USNM and
RWS), (Strandtmann and Morlan, 1953)*
TEXAS: Lubbock Co.; Coll: E, Garrett; 30 April 19^5J 3 females
(RWS).
Nootoma fusipes Baird
CALIFORNIA: Alameda C o., B e rk le y , S tra w b e rry Canyon; C o ll:
F. J. Radovslcy; 19 August I96h; lab reared female progeny (FJR).
Hirstionyssus (H. ) neotomae (Eads and Hightower, 1951-)
(Figs. 77-78)
Nooichoron.yssus nootomao Eads and Hightower, 1951 > Proc. Entomol. Soc.
Washington 53(5):295-298 [Typo: Zavala Co., Texas; Holotype,
U. S . N. M .] .
Hirstionyssus neotomao, Strandtmann and Morlan, 1953» Texas Rep, Biol#
Ked. 11(h):629-632 (key, fig s., host); Strandtmann and Wharton, 159
195S, Inst, Acarology Contrib. 4:106 (synon,, host, d istr,)|
Allred and ^eck, 1966, Brigham Young Univ. Sci, B ull,, Biol,
Ser, 8(1) :29 (key, figs,, host, d istr,).
D ia g n o s is
Deutostex'nal denticles 12 - 14 in numberi arranged in alternating
double file. Anterior margin of sternal shield slightly convex;
posterior margin usually moderately concave (second pore to setae
st. 2), although sometimes only slightly concave (setae st, 3 to
second pore). Lino formed by end of genital flap ribs irregu
larly straight. Setae Jvl usually on postoro-latoral margin of
genital shield, although sometimes slightly removed onto soft
integument but s till touching margin of shield, Paranal setae
positioned at level near middle of or slightly posterior to
middle of anal field, peritrome subequal in width, usually
slightly wider posteriorly; extending moro or loss to middle
of coxa Ij normal in appearance throughout. Greatest width of
dorsal shield at levol of setae Zl - 51 j latoral sides straight
or nearly so and slightly converging toward anterior end;
posterior end in shape of broad, blunt to narrow, sharp wedge.
All dorsal setae present and normally developed. Distance
between setao Zl and SI usually equal to or greater than length
of Zl. Setae Z5 and S5 slightly barbed. Dorsal shield sotae
subequal and quite long, with j2, j3» z2, and z4 longest. Coxal
spur formula 0-2-2-lj inner spur on coxa II small and acute to
blunt; inner spur on coxa III small and acute; outer spur on 160
coxa III small and acute; spur on coxa IV sm 'll, slender, and
acute. Seta p!L, on tibia IV absent. All other leg setae
present and normal.
Collection Records
Cynomys ludovicianus (Ord)
KANSAS: Barber Co., SW Aetna; 16 September 1953t 2 females (RWS),
Thomomys talpoides (Richardson)
UTAH: Carbon Co., So. Schofield; Coll: D S. Beck and C. Pritchett;
30 J u ly I9 6 0 ; 1 fem ale (DMA), ( A llre d and B eck, 19 6 6 ),
Dipod omys b rd ii V.oodhouse
UTAH: San Juan Co.; (Allred and Beck, 3966),
Ferognathus parvus (Feale)
UTAH: Juab Co.; ICane Co., Cottonwood Creek; Coll: D. M. Allred;
10 June 1953; 3 females (DMA), (Allred and Beck, 1966 ).
Liomys irroratus (Gray)
TEXAS: Cameron Co.; June i 960 - November 1962; (Eads, Trevino,
and Campos, I 965 )•
Feromyscus maniculatus (Wagner)
UTAH: Kane Co., Cottonwood Creek; 10 Juno 1953; 30 females (DMA),
(Allrod and Beck, 1966).
Peromyscus truoi (Shufeldt)
UTAH: Kane Co., Cottonwood Crook; 10 June 1953; 10 females (DMA);
San Juan Co., Dead-Korse-Foint; 7 May 1959; 1 female (DMA),
(Allred and Beck, 1966). 161
*Neotoma nlcropus Baird
*TEXAS: Zavala Co,; Colli B, G, Hightovier; 19 October 195^1
1 female (Holotype) (USNK)l Ector Co.j Colli V, T, Miles;
19 Kay 19^9J (Eads and Hightower, 1951).
Ne otoma albigula albigula Hartley
NSW MEXICO: Santa Fe Co., Santa Fe; Coll: H. B, Korlan; ZU- June
1952; 1 female (RWS),
Meotoma lepida lepida Thomas
NEVADA: Nye C o ,, M ercury, AEC-MTS; 13 - 15 June 1961j 3 females
(DMA), (A llr e d and G oates, 1 9 6 ^ a ),
UTAH: Kane Co., Cottonwood Creek; Coll: D, M. Allred; 10 June
1953; h-3 fe m a le s (DMA); same l o c a l i t y ; C o ll: C, P r i t c h e t t ;
17 June I960; 1 female (DMA), (Allred and Bock, I 966 ).
Meotoma c in e r e a (Ord)
CALIFORNIA: Plumas C o ,; C o ll: E. W. Jameson, J r.; 13 October
19^9; 1 female (R'NS).
IDAHO: B u tte C o ., Id ah o F a l l s , AEC-NRTS; 27 - 23 Septem ber 1966 j
15 fe m a le s (DMA),
UTAH: Daggett Co., Bridgeport; Coll: D E, Beck et a l, j 11 June
1953! 6 fe m a le s (DMA); Duchesne C o ., Tim ber Canyon; C o ll: D, M,
Allred; 20 August 1957! 6 fe m a le s (DMA), ( A llr e d and Beck, 1 9 6 6 ). 162
Kirstionyssus (H.) bjsetosus Allred, 1957
(Figs. 79-80)
Hirstionyssus bjsetosus Allred, 1957* Proc. Entomol, Soc. Washington
59(2):83-39 [Type: SS Cedar Ktns., Tooele Co., Utah} Holo-
type, U, S. N. M, ] j Allred and Beck, 1966, Brigham Young Univ.,
Sci, Bull., Biol. Ser, 8(1):29 (key, figs., host, distr.).
D ia g n o sis
Deutosternal denticles ih - 16 in numberj arranged in
alternating double file. Anterior margin of stornal shield
straight to slightly convex} posterior margin usually
moderately concave (second pore to setae st. 2 ), although
sometimes only slightly concave (setae st, 3 to second pore).
Lino formed by end of genital flap ribs never arched, but
distinctly curved down. Setae Jvl always on postoro-latoral
margin of genital shield. Paranal setae positioned at level
near middle of or slightly posterior to middle of anal field.
Peritreme subequal in width; usually slightly wider poste
riorly; extending to anterior half of coxa I; normal in
appearance throughout. Greatest width of dorsal shield at
l e v e l o f Z 1 - SI setae; lateral sides straight or nearly so
and gradually converging toward anterior end; posterior end
in shape of broad, blunt wedge. All dorsal setae present and
normally developed. Distance between setae Zl and SI slightly
greater than length of Zl. Setae Z5 usually slightly barbed. 163
Dorsal shield setae suboqual in length, with some anterior
marginal setae slightly longer. Coxal spur formula 0-2-2-0j
inner spur on coxae II and III small and usually acute; outer
spur on coxa III small and acute. Seta pl^ on tibia IV
absent. All other leg setae present and normal.
Collection Records
*Neotoma le p id a le p id a Thomas
•iSVADAs 'Jye C o., Mercury, AEC-NTS; 6 December 1961; 1 female
(DMA), (Allred and Goates, 196^a),
*UTAH 1 Tooele Co., SE White Rock, Cedar Ktns,j Coll; D. M.
Allred and S, Kulaik; 22 July 195^; sevei’al females (Holotype
and paratypes) (USHK and DMA), (Allred, 1957a )i same approximate
locality; April 195^ “ April 1955; (Allred and Roscoe, 1957)*
Subgenus "X"us, new subgenus
D ia g n o sis
Deutostomal denticles 10 - 12 in number; arranged basically
in single file. Stornal shield small, without postero-lateral
projections bearing setao st. 3 * •» se ta e s t . 3 off shield
on soft integument. Paranal setao at far antorior end of anal
fiold. Feritromo very short, extending only to antorior margin
of coxa III. Posterior end of dorsal shiold in shape of narrow,
caudal projection. Dorsal setae 26 and J2 absent. With bluntly
bifid spurs on some or a ll coxao (I - IV). Spur on coxa IV 1 & \
located in same position as inner spur on coxae II and III,
never on latero-ventral margin. Setae pd^ of genua IV absent.
Hirstionyssus ("X") staffordi Strandtmann and Hunt, 1951
(F ig s. 8 1 -3 3 )
Hirstionyssus staffordi Strandtmann and Hunt, 1951* J. Farasitol.
37(5)j^O-^62 [Typet Grady C o ., Georgia; Holotype,
I). S . N. M. ] ; Strandtmann and Morlan, 1953* Texas R ep.
Biol. Fed. 11(*4*):630 (key, fig s., host); Strandtmann and
Wharton, 1958, Inst. Acarology Contrib. ^ j 108 (synon., host,
d istr,); Allred and Beck, 1966, Brigham Young Univ. Sci. B ull.,
Biol. Ser. 8(1):18 (key, figs., host, distr.).
D ia g n o sis
Deutostornal denticles 10 - 12 in number; arranged in single
file. Anterior margin of sternal shield straight to slightly
convex; posterior margin irregular, without postero-lateral
projections bearing setae st. 3. Line formed by end of genital
flap ribs irregularly straight to slightly arched. Setae Jvl
off genital shield on soft intogument and removed from margin
of shield by distance greater than setal base diameter.
Faranal sotae positioned at antoi'ior end of anal field. F e r i-
treme suboqual in width, usually slightly widor posteriorly*
extending only to anterior half of coxa III; normal i n
appearance throughout. Greatest width of dorsal shield at
level of setae Zl - S2; lateral sides usually slightly concave 165
and converging toward anterior end; posterior end forming
narrow, caudal projection. Dorsal setae z 6 , J 2 , S l t an d S3
absentj with one pair of R series setae on postero-lateral
margin adjacent to usual position of setae S3. All other
dorsal setae present and normally developed. Distance
between setae Zl and margin of dorsal shield greater than
length of Zl. Setae Z5* S5» and the R series seta on margin
of shield slightly barbed. Dorsal shield setae j^, j5, j 6 ,
and z5 minute} setae jl, Jl, and J3 small but not minute} all
other setae moderate and suboqual in length. Coxal spur
formula 1-3-2-1} spur on coxa I moderate in size and blunt to
bluntly bifidj inner spur on coxae II and III large and
usually bluntly bifid} postero-latoral spur on coxa II moderate
in size and blunt to rounded} outer spur on coxa III small and
usually blunt 5 spur on coxa IV located in same position as
inner spur on coxae II and III, never on latoro-ventral margin,
usually small to moderate in size, and blunt to bluntly bifid.
Setao av-^ and pv^ on tarsus II stout and claw-like; seta pd^
of gonu IV absent. All other leg setae presont and normal.
Collection Records
♦Spilogale putorius (Linnaeus)
♦GEORGIAi G rady Co.} C o lli H. B, K o rlan f 12 O c to b e r - 23 November
19^8} 7 females (Paratypes) (USNK and RWS)f Decatur Co.} Colli
H. B. M orlan} 12-15 October 19^8} 9 females (Paratypes) 166 (USNM and RWS), (Strandtmann and Hunt, 1951)I Decatur Co.|
Grady Co,j Brooks Co.; Thomas Co,{ (Morlan, 1952).
Spilogale lnterrupta (Rafinesquo)
IOWA: Story Co., Amos; Colit S. W. Craig; 28 April 1960|
1 fomalo (RWS).
OKLAHOMA: Payno Co., Cushing; Coll: K, C. Emerson; 28 Octobor
193SI (Strandtmann and Hunt, 1951)*
Spilogale gracilis Kerrlam
UTAHt Utah Co., Provo Canyoni Coll: D E, Beck; 11 July 196l|
3 females (DKA), (Allred and Beck, 1966).
Spilogale gracilis (=leucoparia) gracilis Kerriara
TEXAS: P r e s id io C o .; C o ll: W, F . B l a i r ; 22 J u ly 19*^7; 1 fe m a le
(RWS), (Strandtmann and Hunt, 1951)*
Mephitis mephitis (Schrebner)
ILLINOIS: Carroll Co . 1 Coll: B. J. Verts; 8 July 1958*
1 female (OSU),
MICHIGAN: Oakland Co,, Royal Oak; Coll: A, Brown; 26 March 195*H
2 females (OSU).
Mephitis mephitis elongate Bangs
GEORGIA: Brooks Co,; Coll: H. B. Morlan; 15 March I 9U61
14- December 19^-8; 1 fomale (RWS); Decatur Co.; Coll: H, B, Morlan;
26 August 19^8; Grady Co,; Coll: H. B. Morlan; 21 January 19*1-91
Newton Co,; Coll: Travis and Komarek; 8 July 1936; 3 females (RWS),
(Strandtmann and Hunt, 195l)s Decatur Co,; Grady Co.; Brooks Co.;
Thomas Co.; (Morlan, 1952). Mephitis mephitis mesomelas Lechtenstein
TEXAS: Hayes Co.} Colli R. 3. Eads and G. C* Menzies; 1*1- July
19^7S Travis Co., Coll: M iller; 7 March 19^8; (Strandtmann and
Hunt, 1951).
"sk u n k "
MARYLAND: Coll: L, J. Stannard; 1957; 10 females (USKM)•
Hosts of Nearctic Hirstlonyssus Taxospecies
Class Mammalia
Subclass Theria
Infraclass Eutheria
Superorder Monodelphia
Order Insectivora
Suporfamily Soricoidea
Family Soricidae
Subfamily Soricinae
Genus Sorox Linnaeus
S. cinereus Kerr
H. talpae Zemskaya, 1955
S. fumeus M iller
H. "3" n. sp.
H. talpae Zemskaya, 1955
S. trowbridgii Haird
H, obsoletus Jameson, 1950
H. utahensls Allred and Peck, 1966 168
.S. palustris Richardson
Jl* talpae Zemskaya, 1955
Genus 31arina Gray
B. brevicauda (Say)
H. "B" n , s p ,
H. t a lp a e Zem skaya, 1955
Germs Cryptotls Pomel
C. parva (S ay)
H. talpae Zemskaya, 1955
Family Talpidae
Subfamily Scalopinae
Genus Keutrotrichus Gunther
N. gibbsii (Baird)
H. utahensis Allred and Beck, 1966
Genus Scapanus Pomel
S. townscndli (Bachman)
H. obsoletxts Jameson, 1950
Genus Scalopus E. Geoffroy-St. Hilaire
aquaticus (Linnaeus)
H. "B" n . s p .
Genus Parascalopus True
P. browori (Bachman)
H, "B" n . s p .
Family Loporidao
Subfamily Leporinae Genus Lapus Linnaeus
L. californicus Gray
H. triacanthus (Jameson, 1950)
Order Rodentia
Suborder Sciuromorpha
Superfamily Sciuroidea
Family Sciuridae
Subfamily Sciurinao
Genus Cynomys Rafinosque
H. cynomys (Radford, 19^1)
C. ludovicianus (Ord)
H. "A" n . sp .
H. nootomao (Eads and Hightov:or, 1951)
Genus C jtellus Oken
C, townsendii (Bachman)
H. triacanthus (Jameson, 1950)
C. armatus (Kennicott)
H. cynomys (Radford, 19^1)
H. femuralis Allrod, 1957
H, isabellinus (Oudomans, 1913)
H. utahensis Allred and Beck, I 966
tridecomlinoatus (Fitchill)
goomydis (Keogan, 19^6 )
H. utahensis Allred and Beck, I 966 C. variegatus (Erxleben)
H* incomptis (Eads and Hightower, 1952)
C_t leuourus (Eerriam)
H. incowptis (Eads and Hightower, 1952)
H. triacanthus (Jameson, 1950)
0* l a t e r a l i s (S ay )
H, utahensis Allred and Beck, 1966
"Wyoming ground squirrel"
H. cynomys (Radford, 19*4-1)
Genus Eutamjas Trouossart
H. eutamiao Allred and Beck, 1966
H. lncomptis (Eads and Hightower, 1952)
H, u ta h e n s is A llre d and Bock, 1966
E, minimus (Bachman)
H. h illi (Jameson, 1950)
H. "D" n . s p ,
H. occidentalis (Ewing, 1923)
H. utahensis Allred and Bock, 1966
E. quadrivittatus (Say)
ii* Qutamiae Allred and Beck, 19 66
H. isabellinus (Oudemans, 1913)
H. occidontalis (Ewing, 1923)
H. utahensis Allrod and Beck, 1966
E, tovmsendii (Bachman)
H, e u ta m ia e A llre d and Bock, I 966 171 Germs Sclurus Linnaeus
S. abort! Woodhouse
H. torus Allred and Beck, 1966
Genus Tamiasciurus Trouessart
T. hudsonicus richardsoni (Bachman)
H. occidentalis (Ewing, 1923)
T. douglasii (Bachman)
II* afflnis (Jameson, 1950)
Subfamily Pteromyinao
Genus Glaucomys Thomas
G. sabrinus (Shaw)
H. occidentalis (Ewing, 1923)
Superfamily Geomyoidea
Family Goomyidao
Subfamily Geomyinae
Genus Thomomys Wied-Kouwied
H. femurails Allred, 1957
5* thomomys Allred and Bock, 1966
T. bottae (Eydoux and Gervais)
Ii* fQTnuralis Allred, 1957
H. geom.yriis (Koegan, 19*4-6)
H. longlchelae Allred and Beck, 1966
T. talpoidos (Richardson)
H, femuralis Allred, 1957
H. geomydis (Koegan, 19^6) H. isabelllnus (Oudemans, 1913)
H. longlcholae Allrod and 3ockf 1966
H. nootomae (Eads and Hightower, 1951)
H. thomomys Allred and Beck, 1966
H. u ta h e n s is A m red and Bock, 1966
Genus Geomys Rafinesquo
G. bursarlus (Shaw)
H. geomydis (Keegan, 19^6)
Genus Cratogeomys Kerrlam
C, castanops (Baird)
H. femuralis Allred, 1957
gymnurus im parilis (Goldman)
H. f e m u ra lis A llre d , 1957
Family Hotoromyidae
Subfamily Ferognathinae
Genus Perognathus VJiod-Nouwied
H. h illi (Jameson, 1950)
P. flavus Baird
H. h i l l i (Jam eson, 1950)
P. longimombrls (Coues)
H. h illi (Jameson, 1950)
H. inoomptls (Eads and Hightower, 1952)
_H* triacanthus (Jameson, 1950)
_H. utahensis Allred and Beck, 1966 P. parvus (Peale)
H. h illi (Jam©son, 1950)
H, neotomae (Eads and Hightower, 1951)
H, triacanthus (Jameson, 1950)
P. formosus 1'orrlara
H. "C" n . s p .
triacanthus (Janason, 1950)
P. hjspldus Baird
H. "C" n . sp .
Subfamily Dipodomyinae
Genus Dipodomys Gray
H. incomptis (Eads and Hightowor, 1952)
H, triacanthus (Jameson, 1950)
D. m orriam i I'-oarns
inconptis (Eads and Hightowor, 1952)
H. triacanthus (Jamason, 1950)
D, oid .il Vfoodhousa
H. h i l l i (Jam oson, 1950)
H. incomptis (Eads and Hightowor, 1952)
H. nootomae (Eads and Hightowor, 1951)
H, triacanthus (Jameson, 1950)
H* utahensis Allred and Bock, I 966
D. microps (Korriam)
H. Incomptis (Eads and Hightowor, 1952) H. t r i a c a n t h u s (Jam sso n , 1950)
H. utahensis Allred and Pock, I 966
0. deserti Stephens
H, triacanthus (Jameson, 1950)
Subfamily Heteromyinae
Genus Liomys Kerriara
irroratus (Gray)
H, neotom ae (Eads and Hightower, 1951)
Subordor Myomorpha
Superfamily Kuroidea
Family Cricetidae
Subfamily Cricotinae
Gonus Roithrodontomys G igliola
R, mogalotis (Baird)
H. eutamiae Allred and Beck, 1966
H, incomptis (Eads and Hightower, 1952)
H. isabollinus (Oudemans, 1913)
triacanthus (Jamoson, 1950)
X, utahensis Allred and Beck, I 966
Gonus Peromyscus Glogor
H, u ta h e n sis A llro d and Beck, 1966
P. crinitus (Kerriam)
H. incomptis (Eads and Hightower, 1952)
II* triacanthus (Jameson, 1950)
utahensis Allred and Peck, I 966 175
P, eremicus (Baird)
H. femuralis Allred, 1957
H, h illi (Jameson, 1950)
H. incomptis (Eads and Hightower, 1952)
H. "C" n . s p ,
H. triacanthus (Jameson, 1950)
H. utahensis Allred and Beck, 1966
P. maniculatus (Wagner)
H, h illi (Jameson, 1950)
Ii* incomptis (Eads and Hightower, 1952)
H. isabellinus (Oudemans, 1913)
H. longichalaa Allred and Beck, 1966
H, neotomae (Eads and Hightower, 1951)
H. occidentalis (Ewing, 1923)
triacanthus (Jameson, 1950)
H. utahensis Allred and Beck, 1966
P.* lsucopus (Rafinesque)
Ii* utahensis Allred and Beck, I 966
P. boylii (Baird)
H. ”D" n , sp .
ii* utahensis Allred and Beck, I 966
P. truoj (Shufoldt)
H, neotomae (Eads and Hightowor, 1951)
ii* utahonsis Allred and Beck, 1966 Genus Onychomys Baird
0 . leucogaster (Wied-Kouwied)
H. h illi (Jameson, 1950)
H, longichelae Allred and Beck* 1966
0, torridus (Coues)
isabellinus (Oudomans, 1913)
.H. triacanthus (Jameson, 1950)
Genus Sigmodon Say and Ord
S. hispidus Say and Ord
J5* talpae Zemskaya, 1955
Genus ITeotoma Say and Ord
N. micropus Baird
H. neotomao (Eads and Hightower, 1951)
K. albigula albigula Hartley
H, breviseta Strandtmann and Morlan, 1953
il* uootomae (Eads and Hightower, 1951)
N, lepida lepida Thomas
H. bjsetosus Allrod, 1957
femuralis Allred, 1957
H. h illi (Jameson, 1950)
H, incomptis (Eads and Hightower, 1952)
H, neotomao (Eads and Hightower, 1951)
H. triacanthus (Jameson, 1950)
H. utahensis Allrod and Beck, 1966 N, fuclpos Baird
breviseta Strandtmann and 1'orlan, 1953
N. c in e r a a (O rd)
H. neotomao (Eads and Hightower, 1951)
H. triacanthus (Jameson, 1950)
Subfamily Kicrotinae
Genus Lernmus lin k
L. trimucronatus (Richardson)
H. isabellinus (Oudomans, 1913)
Gonus Clethrionomys Tilesius
H* isabellinus (Oudemans, 1913)
C. rutilus (Pallas)
H. isabellinus (Oudemans, 1913)
Gonus Phonaoomys I'orriam
P. intermedius Kerriam
H. isabellinus (Oudemans, 1913)
Genus Hicrotus Schrank
H. isabellinus (Oudemans, 1913)
M, pennsylvanicus (Ord)
H. isabellinus (Oudemans, 1913)
II* rcontanus (Pealo )
H. isabellinus (Oudemans, 1913)
M. longicaudus (Kerriam)
11* incomptis (Eads and Hightower, 1952)
H. isabellinus (Oudomans, 1913) Family Furidae
Subfamily Ihrinao
Genus Rattus G, Fischer
R, norvegicus (Berkenhout)
H, incomptis (Eads and Hightovrer, 1952)
_H. latiscutatus (de Keillon and Lavoipierre, 19^)
H. triacanthus (Jameson, 1950)
Genus I--us Linnaeus
M, musculus Linnaeus
H. isabellinus (Oudemans, 1913)
H# latiscutatus (de Keillon and Lavoipierre, 19^0
Order Carnivora
Suborder Fissipedia
Superfamily Canoidoa
Family Kustolidae
Subfamily Kustelinae
Genus I?ustela Linnaeus
M, frenata Lichtenstein
ii* louglcholae Allred and Beck, 1966
Subfamily Kophitinae
Genus Spilogale Gray
S. putorius (Linnaeus)
H. staffordi Strandtmann and Hunt, 1951
S_, interrupta (Rafinesque)
H, staffordi Strandtmann and Hunt, 1951 S. gracilis Kerriara
Jh staffordi Strandtmann and Hunt, 1951
S.* (= leucoparia) gracilis Kerri am
Jn staffordi Strandtmann and Hunt, 1951
Genus Kophitis S, Geoffroy-St. Hilaire and G. C u vier
£1* niophitis (Schrebnor)
H. staffordi Strandtmann and Hunt, 1951
Kt mephitis elongata Bangs
H, staffordi Strandtmann and Hunt, 1951
M, mephitis mosomelas Lechtonstein
H. staffordi Strandtmann and Hunt, 1951
"skunk"
H. staffordi Strandtmann and Hunt, 1951 unknown hosts
H, "C" n , s p ,
H. isabellinus (Oudomans, 1913)
C la ss Avos
Superorder Koognathae
Order Caprimulgiformes
Family Caprinulgidae
Genus Chordoilos
C, minor Grinnoll
H, triacanthus (Jamoson, 1950) Class Raptilia
Subclass Diapsida
Order Squamata
Suborder Lacertilia
Family Iguanidae
Subfamily Iguaninae
Genus S celop oru s Wiegmann
S_, graciosus Baird and Girard
H, triacanthus (Jameson , 1950) SUMMARY AND CONCLUSIONS
Relationships among the Nearctic members of the gonus
Hirstionyssus were examined using the techniques of numerical taxonomy.
The study comprised 105 individual specimens (OTU's) including females of a ll previously described and several undescribed Nearctic taxo- species. Each taxospecies was represented by from one to 20 replicates, one of which was a type specimen in a ll except three cases. Replicates from different hosts and localities were included to study the effect of intrataxon variation, and to provide an estimate of the effect of differ ent hosts and localities, A total of 152 characters (2h"£ qualitative and 76^ quantitative) was recorded from a ll areas of the body. Throe methods of transforming the basic data matrix were used and compared.
Coefficients of Pearson’s product-monent correlation (r) and Sokal's taxonomic distance (d) were computed, and analyzed by UFGKA (Unweighted
Pair-Group Method with Average linkage) cluster analysis, yielding a total of six phenograms. Subsequently, a revised Condonsed Ratio Corre lation matrix was computed from a reduced basic data matrix of 115 characters. This correlation matrix was analyzed by graphic analysis and illustrated in two-dimensional graphic diagrams.
Phonogram groupings obtained from six sim ilarity matricos wore compared with respect to the placement of 0TU*s and CTU clusters. The two phenograms based on standardized basic data and the distance 181 182 phonogram based on condensed basic data were quite different, and yielded low cophenetic correlation coefficients. The distance pheno grams based on standardized basic and condensed basic data were sim ilar to each other, but gave very unsatisfactory results. The standardized basic correlation phonogram was very different from a ll others and was considered to give the poorest illustration of relationships. Three of the phenograms based on condensed data were very sim ilar to each other and gave the most useful results. The correlation and distance pheno grams based on condensed ratio data yielded the highest cophenetic correlation coefficients, with the correlation phonogram based on con densed basic data the third highest. In the case of correlation, con densation of basic data and computing ratios of quantitative characters
prior to condensation yielded almost identical phenograms.
The relationships among OTU's and OTU clusters in the graphic
analysis diagrams based on correlation were analyzed and compared with
the phenograms produced by cluster analysis. These two-dimensional
graphic diagrams were found to illustrate phonetic relationships some what hotter than the phenograms. The relationships observed in the
numerical taxonomic analyses wore then compared with those expressed in
a reduced matrix of character state values of 30 readily recognizable
qualitative characters, and with host and geographic distribution.
As a rosult of this study 25 taxospecies, including four previ
ously undoscribed, were formally recognized. Five previously described
taxospecies were relegatod to synonymy. The congruence of phenetic
sim ilarity among these taxospecies with host and geographic distribution 183 was quite good. On the basis of the numerical taxonomic analyses and the distribution of diagnostic qualitative characters, these 25 taxo species were placed in six rather well defined host groups* A formal taxonomic change proposed was the recognition of two subgenora of
Nearctic Hirstionyssus; "X"us, new subgenus, to include the single taxospecies of the carnivore host group, H, staffordi Strandtmann and
Hunt; and Hirstionyssus s_. s tr., to include the other 2h taxospecies.
The following suggestions and conclusions are made as a conse quence of this investigation*
1. It is suggested that when most characters are measurements
and the OTU's differ in overall size, either correlation
coefficients based on condensed data should bo used, or
the quantitative characters should bo converted to ratios
of some sort prior to the condensation transformation;
good results may then be obtained from both correlation
and distance.
2. The inclusion of many highly correlated quantitative
characters in a study tends to cause excessive weighting
by those groups of characters, resulting in distorted
sim ilarity values.
3. Two-dimensional graphic analysis diagrams illustrate
phonetic relationships somewhat more accurately than
the one-dimensional phenograms. The blind acceptance of any one combination of sim ilarity coefficient and clustering procedure without reference to the distribution of diagnostic qualitative characters is unwarranted from a u tili tarian viewpoint. APPENDIX I
LIST OF FEMALE SPECIMENS (OTU-S) STUDIED FROM THE GENUS HIRSTIONYSSUS# HOLOTYPES ARE DESIGNATED by AN ASTERISK.
OTU HOST SPECIES COLLECTION DATE COLLECTION LOCALITY MUSEUM
GEOMY1D HOST GROUP
*FEMU 1 PEROMYSCUS EREMICUS IS DECEMBER 1950 UTAH- WASHINGTON CO.* ROCKVILLE DMA
FEMU 2 THOMOMYS BOTTAE 22 APRIL 1951 UTAH- KANE CO.* KANAB DMA
FEMU 3 THOMOMYS SP. 18 DECEMBER 1950 UTAH- WASHINGTON CO.* ROCKVILLE DMA
FEMU 4 CRATOGEOMYS CASTANOPS 7 JUNE 1948 TEXAS- PRESIDIO CO. RWS
*GEOM 1 GEOMYS BURSAR I US 6 MARCH 1944 KANSAS- LEAVENWORTH CO.* FT* LEAVENWORTH USNM
GEOM 2 GEOMYS BURSAR I US 23 APRIL 1949 TEXAS- LEON CO. RWS
*LONG 1 THOMOMYS TALPOIDES 26 JUNE 1951 UTAH- UTAH CO.* DIAMOND FORK CANYON DMA
LONG 2 OCHOTONA PRINCEPS 8 AUGUST 1957 UTAH- SUMMIT CO.* BALD MTN. DMA
LONG 3 THOMOMYS TALPOIDES 7 MARCH 1967 IDAHO- BUTTE CO.* IDAHO FALLS* AEC-NRTS DMA
*THOM 1 THOMOMYS TALPOIDES 0 AUGUST 1957 UTAH- SUMMIT CO.* BALD MTN. DMA
THOM 2 THOMOMYS SP. 20 OCTOBER 1966 IDAHO- BUTTE CO.* IDAHO FALLS* AEC-NRTS DMA
135 APPENDIX I CONTINUED
OTU HOST SPECIES COLLECTION DATE COLLECTION LOCALITY MUSEUM
HETEROMY’IO HOST GROUP
*HILL 1 PEROGNATHUS PARVUS 24 AUGUST 1949 NEVADA- WASHOE CO.* NORTH OF RENO USNM
HILL 2 PEROGNATHUS PARVUS 1 JUNE 1951 UTAH- JUAB CO.* JOY DMA
HILL 3 PEROGNATHUS PARVUS 15 JULY 1967 IDAHO- BUTTE COoi IDAHO FALLS* AEC-NRTS DMA
HILL 4 PEROGNATHUS FORMOSUS 13 JUNE 1961 NEVADA- NYE CO.* MERCURY* AEC-NTS DMA
HILL 5 PEROGNATHUS HISPIDUS JULY 1940 KANSAS- MEADE CO.* SW MEADE EWJ
* INCO 1 DIPODOMYS ORD I I 16 JANUARY 1947 TEXAS- TERRY CO. USNM
INCO 2 DIPODOMYS ORDII 30 APRIL 1957 UTAH- SEVIER CO.* RICHFIELD DMA
INCO 3 PEROMYSCUS MANICULATUS 16 JULY 1967 IDAHO- BUTTE CO.* IDAHO FALLS* AEC-NRTS DMA
*TRIA 1 DIPODOMYS MERRIAMI 9 OCTOBER 1949 NEVADA- WASHOE CO.* PYRAMID LAKE USNM
TRIA 2 DIPODOMYS ORDII 12 JULY 1958 UTAH- SAN JUAN CO.* MONUMENT VALLEY DMA
TRIA 3 DIPODOMYS MERRIAMI 16 DECEMBER 1958 ARIZONA- PIMA CO.« TUCSON OSU
SCI URID HOST GROUP
#AFFI 1 TAMIASCIURUS DOUGLASII 13 MAY 1949 CALIFORNIA- PLUMAS CO.* QUINCY USNM
AFFI 2 EUTAMIAS MINIMUS li JULY 1957 UTAH- BOX ELDER CO.* YOST DMA
AFFI 3 EUTAMIAS MINIMUS 22 JUNE 1949 CALIFORNIA- MODOC CO.* ALTURAS RWS 187 APPENDIX I - CONTINUED
OTU HOST SPECIES COLLECTION DATE COLLECTION LOCALITY MUSEUM
AFFI 4 PEROMYSCUS BOYLII 4 OCTOBER 1950 CALIFORNIA- PLUMAS CO** QUINCY RWS
CYNO 1 CYNOMYS SP* 7 NOVEMBER 1932 COLORADO- CONEJOS CO** ANTONITO CDR
*INVA I CITELLUS ARMATUS 22 JULY 1956 UTAH- UTAH CO* * PROVO CANYON DMA
INVA 2 CITELLUS ARMATUS 7 AUGUST 1953 UTAH- WASATCH CO** LAKE CREEK SUMMIT DMA
*ANGU 1 CITELLUS ARMATUS 7 JULY i960 UTAH- UTAH CO** COLTON DMA
NSPA 1 CYNOMYS LUDOVICIANUS 16 SEPTEMBER 1953 KANSAS- BARBER CO** AETNA RWS
NSPA 2 CYNOMYS LUDOVICIANUS 16 SEPTEMBER 1933 KANSAS- BARBER CO** AETNA RWS
*EUTA 1 EUT AM I AS SP* 16 OCTOBER 1956 UTAH- UTAH CO** ASPEN GROVE DMA
EUTA 2 EUT AM IAS TOWNSEND I I JUNE 1962 CALIFORNIA- EL DORADO COo* ECHO SUMMIT EWJ
*OCCI 1 TAMI ASCIURUS HUDSONICUS 22 JUNE 1910 MONTANA- MISSOULA C0*« LOLO USNM
*PUNC 1 EUTAMI AS QUADRIVITTATUS 25 AUGUST 1956 UTAH- SUMMIT CO** BALD MTN* DMA
PUNC 2 EUTAMIAS QUADRIVITTATUS 8 AUGUST 1957 UTAH- SUMMIT CO*• eALD MTN* DMA
PUNC 3 EUTAMI AS QUADRIVITTATUS 25 AUGUST 1956 UTAH- SUMMIT C0*» BALD MTN* DMA
PUNC 4 GLAUCOMYS SABRINUS 29 JUNE 1958 UTAH- DAGGETT CO** DEEP CREEK DMA
*TORU I SCIURUS ABERT1 6 MAY 1951 UTAH- SAN JUAN COoj BLANDING DMA 188 APPENDIX I CONTINUEO
OTU HOST SPECIES COLLECTION DATE COLLECTION LOCALITY MUSEUM
TORU SCIURUS ABERTI 6 MAY 1951 UTAH- SAN JUAN CO.* BLANDING DMA
TORU SCIURUS ABERTI 3 OCTOBER 1926 COLORADO- LARIMER CO.* LIVERMORE USNM
MUROID HOST GROUP
*BtSE NEOTOMA LEPIDA NEST 22 JULY 1954 UTAH- TOOELE CO.* CEDAR MTNS♦ USNM
BISE NEOTOMA LEPIDA NEST 6 OCTOBER 1954 UTAH- TOOELE CO.* CEDAR MTNS. DMA
BISE NEOTOMA LEPIDA 6 DECEMBER 1961 NEVADA- NYE CO.* MERCURY* AEC—NTS DMA
*BREV NEOTOMA ALBIGULA NEST 18 NOVEMBER 1952 NEW MEXICO- SANTA FE CO.* MASTIN RANCH USNM
BREV NEOTOMA ALBIGULA NEST 30 APRIL 1965 TEXAS- LUBBOCK CO* RWS
BREV LAB REARED COLONY 14 SEPTEMBER 1967 ORIG. EX. NEOTOMA FUS1PES 19 AUGUST 1964 CALIFORNIA- ALAMEDA CO.* BERKLEY FJR
I SAB MICROTUS AGRESTIS BRITAIN- HEBRIDES* ISLE OF MUCK BMNH
I SAB -FIELD VOLE- 29 N0VEM3ER 1936 BRITAIN- CHESHIRE* MACCLESFIELD COR
I SAB MICROTUS ARVALIS NEST 4 AUGUST 1967 GERMANY- NIEDER SACHSEN* 0LDEN3ERG OSU
I SAB APODEMUS AGRARIUS 8 MARCH 1955 U.S.S »R« USNM
I SAB CLETHRIONOMYS SP. 20 APRIL 1952 JAPAN- HOKKAIDO* SAPPORO EW J
I SAB LEMMUS TRIMUCRONATUS 13 JULY 1953 ALASKA- PT. BARROW USNM APPENDIX I CONTINUED
OTU HOST SPECIES COLLECTION DATE COLLECTION LOCALITY MUSEUM
I SAB 7 MICROTUS SP* 6 JUNE 1947 ALASKA- UM1TA USNM
I SAB 0 LEMMUS TR1MUCR0NATUS 1 AUGUST 1949 CANADA- NW TERRITORY* PRINCE PATRICK ISL* USNM
I SAB 9 PHENACOMYS I NTERMEDIUS 1 JUNE 1956 CANADA- MANATOBA* FT* CHURCHILL USNM
I SAB10 MICROTUS PENNSYLVANICUS 22 SEPTEMBER 1955 CANADA- ONTARIO* MANITOULIN ISLAND OSU
I SABI 1 CLETHRIONOMYS SP* 28 JULY 1953 CANADA- QUEBEC* LAKE MARYMAC USNM
I SAB12 MICROTUS LONG ICAUCUS 14 AUGUST 1954 OREGON- HARNEY CO.* STEEN-S MTNS* USNM
I SAB13 MICROTUS MONTANUS 22 MARCH 1949 CALIFORNIA-* PLUMAS CO.* QUINCY RWS
1 SAB14 OCHOTONA PRINCEPS 0 AUGUST 1957 UTAH- SUMMIT CO.* BALD MTN. DMA
I SABI5 MICROTUS LONGICAUCUS 25 AUGUST 1956 UTAH- SUMMIT CO.* BALD MTN* DMA
I SAB 16 MICROTUS PENNSYLVANICUS £9 JULY 1965 MICHIGAN- EMMETT CO.* ALANSON OSU
I SAB17 MICROTUS PENNSYLVANICUS 9 AUGUST 1965 OHIO- ERIE CO. OSU
I SAB 1 8 MICROTUS PENNSYLVANICUS 27 JULY 1965 OHIO- ERIE CO* OSU
I SAB19 MICROTUS SP* 13 MAY 1943 PENNSYLVANIA“ BEAVER CO.. INDUSTRY RWS
ISAB20 MICROTUS PENNSYLVANICUS 21 OCTOBER 1949 NEV/ YORK- ALBANY CO.* RENSSELAERVILLE USNM
LATI 1 MUS MUSCULUS 3 DECEMBER 1963 BRITAIN- NEW CASTLE-UPON-TYNE BMNH 1 9 0 APPENDIX I - CONTINUED
OTU HOST SPECIES COLLECTION DATE COLLECTION LOCALITY MUSEUM
LATI 2 MUS MUSCULUS MASSACHUSETTS- MIDDLESEX CO** CAM3RIDGE OSU
LATI 3 RATTUS NORVEGICUS 14 FEBRUARY 1946 GEORGIA- DECATUR CO*» BA 1NBRIDGE USNM
LATI 4 MUS MUSCULUS 9 JANUARY 1912 OREGON- BENTON CO*« CORVALIS USNM
LATI 5 MUS MUSCULUS 1 JANUARY 1951 UTAH- UTAH CO*I LEHI DMA
*NEOT 1 NEOTOMA MICROPUS 19 OCTOBER 1950 TEXAS- ZAVALA CO* USNM
NEOT 2 CYNOMYS LUDOVICIANUS 16 SEPTEMBER 1953 KANSAS- BARBER CO** W* AETNA RWS
NEOT 3 PEROMYSCUS MANICULATUS 10 JUNE 1958 UTAH- KANE CO* * COTTONWOOD CREEK DMA
NEOT 4 NEOTOMA CINEREA 13 OCTOBER 1949 CALIFORNIA- PLUMAS CO*. QUINCY RWS
*TARS 1 PEROMYSCUS MANICULATUS 23 JUNE 1958 UTAH- DAGGETT CO** DEEP CREEK DMA
*UTAH 1 EUTAMIAS QUADRIVITTATUS 8 AUGUST 1957 UTAH- SUMMIT CO** BALD MTN* DMA
UTAH 2 EUTAMIAS MINIMUS 24 AUGUST 1956 UTAH- SUMMIT CO** BALD MTN* DMA
UTAH 3 PEROMYSCUS MANICULATUS 10 JUNE 1958 UTAH- KANE CO* * COTTONWOOD CREEK DMA
UTAH 4 PEROMYSCUS MANICULATUS 19 AUGUST 1954 OREGON- HARNEY CO** STEEN-S MTNS* USNM
UTAH 5 LAB REARED COLONY 14 SEPTEMBER 1967 ORIG. EX* PEROMYSCUS EREMICUS 16 FEBRUARY 1965 MEXICO- BAJA* SAN QUINTON FJR
UTAH 6 PEROMYSCUS SP* 13 JULY 1960 IOWA- BOONE CO® RWS 191 APPENDIX I - CONTINUED
OTU HOST SPECIES COLLECTION DATE COLLECTION LOCALITY MUSEUM
UTAH 7 PEROMYSCUS LEUCOPUS II AUGUST 1965 OHIO- ERIE CO* OSU
INSECTIVORE HOST GROUP
CARN 1 TALPA EUROPAEA" 21 OCTOBER 1953 U • S* S*R* BMNH
NSPB 1 BLARINA BREVICAUDA 4 AUGUST 1965 OHIO- ERIE CO* OSU
NSPB 2 BLARINA BREVICAUDA 29 APRIL 1954 MARYLAND- PRINCE GEORGES CO** PATUXENT REF* OSU
NSPB 3 BLARINA BREVI CAUDA 20 MAY 1947 PENNSYLVANIA- BEAVER CO*» INDUSTRY RWS
TALP 1 TALPA EUROPAEA 19 JUNE 1943 U*S«S«R* BMNH
TALP 2 TALPA EUROPAEA 29 DECEM3ER 1951 BRITAIN- SUFFOLK* LAKENHEATH BMNH
TALP 3 TALPA EUROPAEA NEST 13 DECEMBER 1942 EUROPE- BMNH
TALP 4 -SHREW- 4 FEBRUARY 1957 TAIWAN- PING-TUNG* CHAO CHOW DMA
TALP 5 BLARINA BREVICAUDA 17 MAY 1954 MARYLAND- PRINCE GEORGES CO«« RIVERDALE OSU
TALP 6 CRYPTOTIS PARVA 6 MAY 1954 MARYLAND- PRINCE GEORGES CO* » PATUXENT REF* OSU
.TALP 7 SIGMODON HISP I DUS 21 JULY 1953 NORTH CAROLINA- WAKE CO** RALEIGH OSU
TALP 8 CRYPTOTIS PARVA 30 SEPTEMBER 1945 KANSAS- DOUGLAS CO** LAWRENCE ' EWJ
TALP 9 SOREX CINEREUS 29 AUGUST 1948 WYOMING- ALBANY CO** BROOKLYN LAKE EWJ
*PALU 1 SOREX PALUSTRIS 29 JULY 1960 UTAH- CARBON CO#* SCHOFIELD DMA 192 APPEND t X I CONTINUED
OTU HOST SPECIES COLLECTION DATE COLLECTION LOCALITY MUSEUf4
OBSO 1 SOREX TROWBRIDGII 10 MARCH 1949 CALIFORNIA- PLUMAS CO.* QUINCY USNM
OBSO 2 SOREX TROWBRIDGII 10 MARCH 1949 CALIFORNIA- PLUMAS CO.* QUINCY USNM
CARNIVORE HOST GROUP
STAF 1 SPILOGALE PUTORIUS 12 OCTOBER 1948 GEORGIA- GRADY CO* USNM
STAF 2 SPILOGALE PUTORIUS 13 OCTOBER 1948 GEORGIA- DECATUR CO. USNM
STAF 3 MEPHITIS MEPHITIS 26 MARCH 1954 MICHIGAN- OAKLAND CO.* ROYAL OAK OSU
STAF 4 SPILOGALE GRACILIS 11 JULY 1961 UTAH- UTAH CO.* PROVO CANYON DMA APPENDIX II
LIST OF CHARACTERS UTILIZED FOR HIRSTIONYSSUS FEMALES* THE ADJECTIVE -LITTLE- MEANS LOWER CASE#
GNATHOSOMA - 17 CHARACTERS
1# WIDTH AT BASE.OF PALPS# 2* GREATEST LENGTH (BASE TO PALP TROCHANTER)* 3# LENGTH PALP GENU# 4* WIDTH PALP GENU# 5* LENGTH SECOND CHELICERAL SEGMENT (BASE TO TIP OF FIXED DIGIT)# 6# LENGTH MOVEABLE DIGIT OF CHELI CERA* 7* LENGTH CAPITULAR SETA (CS)• 8# LENGTH HYPOSTOMAL SETA 3 (HYP3)• 9# LENGTH HYPOSTOMAL SETA 1 (HYP1)# . 10# DISTANCE CS - CS• 11* DISTANCE HYP3 - 3# 12* DISTANCE HYP1 - 1# 13* DISTANCE HYP1 - 3# 14* DISTANCE HYP3 - CS* 15. DISTANCE CS - GNATHOSOMA BASE# 16* NUMBER OF DEUTOSTERNAL DENTICLES* 17* arrangement o f deutosternal d e n t i c l e s * (1) b a s i c a l l y s i n g l e f i l e (2) alternating d o u b l e f i l e (3) REGULARLY DOUBLE FILE
STERNAL AREA - 17 CHARACTERS
18* GREATEST WIDTH TRITOSTERNUM BASE# 19* LENGTH TRITOSTERNUM BASE* 20* LENGTH TRITOSTERNUM LACIN IAE* 21* LENGTH STERNAL SHIELD (IN MIDLINE)* 22* LEAST WIDTH STERNAL SHIELD (BETWEEN COXAE II)* 23* SHAPE OF ANTERIOR MARGIN OF STERNAL SHIELD# (1) SLIGHTLY CONCAVE (2) STRAIGHT (3) SLIGHTLY CONVEX (4) STRONGLY CONVEX 24# SHAPE OF POSTERIOR MARGIN OF STERNAL SHIELD# (1) STRONGLY CONVEX (2) SLIGHTLY CONVEX 193 19^ APPENDIX II - CONTINUED
(3) STRAIGHT 14J IRREGULAR (5) SLIGHTLY CONCAVE (TO SECOND PORE) (6) MODERATELY CONCAVE (SECOND PORE TO ANTERIOR LEVEL ST2) <7) DEEPLY CONCAVE (DISTINCTLY ANTERIOR TO ST2 > 25* LENGTH ST1 (STERNAL SETAE !)• 26* LENGTH ST2* 27* LENGTH ST3a 28 a LENGTH ST4* 29* DISTANCE ST1 - 1o 30a DISTANCE ST2 - 2a 31a DISTANCE ST3 - 3a 32a DISTANCE ST1 - 2a 33a DISTANCE ST2 - 3a 34a POSITION OF ST3 SETAEa (1> ON STERNAL SHIELD (2) OFF STERNAL SHIELD
GENITAL SHIELD - 10 CHARACTERS
35a LEAST WIDTH AT OR ANTERIOR TO GENITAL SETAE (GEN)a 36a GREATEST WIDTHa 37a PLACE OF GREATEST WIDTHa (1) AT GEN SETAE (2) BETWEEN GEN AND ZVI SETAE (3) AT 2V1 SETAE 38a DISTANCE FROM LINE FORMED BY END OF GENITAL FLAP RIBS TO POSTERIOR MARGIN OF GENITAL SHIELD (MEASURED ON MIDLINE)a 39a DISTANCE FROM LEVEL OF GEN SETAE TO POSTERIOR MARGIN OF GENITAL s h i e l d * 40a CURVATURE OF LINE FORMED BY END OF GENITAL FLAP RIBSa (1) CURVED DOWN (2) IRREGULAR (3) STRAIGHT OR NEARLY SO (4) SLIGHTLY BUT DISTINCTLY ARCHED (5) STRONGLY ARCHED 41o LENGTH GEN SETAEa 42a LENGTH JV1 SETAEa 43o POSITION OF JV1 SETAEa (1) ON GENITAL SHIELD (2) OFF BUT TOUCHING GENITAL SHIELD (3) OFF AND REMOVED FROM GENITAL SHIELD BY DISTANCE GREATER THAN SETAL BASE DIAMETER* 44a DISTANCE GEN - GEN* 195 APPENDIX II - CONTINUED
ANAL SHIELD AREA - 13 CHARACTERS
45 a LENGTH EXCLUSIVE OF CRIBUM# 46a GREATEST WIDTHa 47• WIDTH AT CRIBUM ATTACHMENTa 48a LENGTH PARANAL SETAEa 49* DISTANCE PARANAL - PARANAL* 50* POSITION OF PARANAL SETAE RELATIVE TO ANAL FIELD* <1) ANTERIOR END (2) DISTINCTLY ANTERIOR <3 J NEAR MIDDLE (4) DISTINCTLY POSTERIOR (5) POSTERIOR END 51 a LENGTH POSTANAL SETA* 52a DISTANCE PARANAL - POSTANAL* 53a DISTANCE ANTERIOR MARGIN OF ANAL SHIELD TO PARANAL SETAEa 54* DISTANCE ANTERIOR MARGIN OF ANAL SHIELD TO ANTERIOR MARGIN OF ANAL FIELD# 55* DISTANCE POSTERIOR MARGIN OF ANAL FIELD TO POSTERIOR MARGIN OF ANAL SHIELD (CRIBUM EXCLUDED)# 56* LENGTH JV3 SETAEa 57* LENGTH JV5 SETAEa
PER ITREME - 4 CHARACTERS
58a RELATIVE WIDTH (ANTERIOR VERSUS POSTERIOR)* (0) MUCH GREATER POSTERIORLY THAN ANTERIORLY (1) SUBEQUAL - USUALLY SLIGHTLY WIDER POSTER t ORLY (2) DISTINCTLY GREATER ANTERIORLY THAN POSTERIORLY 59* LENGTH MEASURED IN STRAIGHT LINE FROM POSTERIOR MARGIN OF STIGMA TO ANTERIOR TIP* 60. ANTERIOR EXTENT* (1> ANTERIOR HALF COXA I (2) POSTERIOR HALF COXA I (3) ANTERIOR HALF COXA W (4) POSTERIOR HALF COXA II (5) ANTERIOR HALF COXA III 61* SEGMENTATION OF POSTERIOR HALF* (1) NOT SEGMENTED - APPEARANCE SAME AS ANTERIOR HALF (2) DISTINCTLY BROKEN INTO SEGMENTS
DORSAL SHIELD - 45 CHARACTERS
62o LENGTH IN MIDLINE# 63a GREATEST WIDTH# 196 APPENDIX II - CONTINUED
64* PLACE Or GREATEST WIDTH OF DORSAL SHIELD® tl) LEVEL LITTLE S3-4 SETAE <2) LEVEL LITTLE S4-6 SETAE (3) LEVEL SI - Z1 SETAE 65* SHAPE OF LATERAL SIDES® (1) DISTINCTLY CONCAVE (2) STRAIGHT OR NEARLY SO C3> DISTINCTLY CONVEX 66* SHAPE OF POSTERIOR END* Cl> BROADLY ROUNDED (2) BROAD* BLUNT WEDGE (3) NARROW* SHARP WEDGE (4) NARROW« CAUDAL PROJECTION 67* SETAE LITTLE Z6* CO) ABSENT
77 LENGTHSETAE LITTLE J3o 78 LENGTH SETAE l i t t l e J6 • 79 LENGTH SETAE l i t t l e Z2 • 80 LENGTHSETAE l i t t l e S2 • 81 LENGTH SETAE l i t t l e S4 • 82 LENGTHSETAE J3* 83 LENGTH SETAE S2 • 84 LENGTH SETAE S5* 85 l e n g t h SETAE Z5* 86* DISTANCE LITTLE J3 - 3* 87• DISTANCE LITTLE J4 - 4* 88• DISTANCE LITTLE JS - 5* B9• DISTANCE LITTLE Z5 - 5# 90* DISTANCE LITTLE J6 - 6* 91* DISTANCE J1 - 1* 92* DISTANCE J3 - 3* 93* DISTANCE LITTLE J4 TO EDGE OF SHIELD (PARALLEL WITH MARGIN!• 94* DISTANCE LITTLE S4 TO EDGE OF SHIELD* 95* DISTANCE LITTLE J6 TO EDGE OF SHIELD* 96* DISTANCE J1 TO EDGE OF SHIELD* 97* DISTANCE Z1 TO EDGE OF SHIELD* 98* DISTANCE J3 TO EDGE OF SHIELD* 99* DISTANCE LITTLE Z2 - 4* 100* DISTANCE LITTLE J1 - 5* 101* DISTANCE LITTLE J3 - 4* 102* DISTANCE LITTLE J5 - LITTLE Z5* 103* DISTANCE LITTLE J5 - 6* 104* DISTANCE LITTLE J6 - JI • 105* DISTANCE Jl - 3* 106. DISTANCE J3 - Z5.
LEGS - 46 CHARACTERS
107* LENGTH PROXIMAL SETA OF COXA I* 108* LENGTH DISTAL SETA OF COXA I* 109* SHAPE SPUR ON COXA I* (0) ABSENT ( 1 > BLUNT (2) ROUNDED <3> BIFID AND BLUNT 110* LENGTH ANTERO-VENTRAL SETA ON COXA II* 111* FORM OF ANTERO-VENTRAL SETA ON COXA I I• <0) NORMAL* LONG AND SLENDER (1) STOUT* ACUTELY SPINOSE* AND STRAIGHT (2) STOUT* BLUNT* AND PEG-LlKE (3) STOUT* SPINOSE* AND HOOKED 112* LENGTH POSTERO-VENTRAL SETA ON COXA II* 113* LENGTH ANTERO-DORSAL SPINE ON COXA II* 193 APPENDIX II - CONTINUED
114* SHAPE INNER SPUR ON COXA II* 10) INDISTINCT! BROAD* SLIGHTLY RAISED APOPHYSIS £1) ACUTE £ 2 ) BLUNT (3) ROUNDED (4) BIFID AND BLUNT 115* LENGTH INNER SPUR ON COXA II* 116* BASAL WIDTH INNER SPUR ON COXA II* 117* SHAPE POSTERO-LATERAL SPUR ON COXA II* £0) ABSENT £1 ) ACUTE (2) BLUNT 13) ROUNDED 118* LENGTH ANTERO-VENTRAL SETA ON COXA III* 119* LENGTH POSTERO-VENTRAL SETA ON COXA III* 120* SHAPE INNER SPUR ON COXA III* (0) INDISTINCT* BROAD* SLIGHTLY RAISED APOPHYSIS £1 ) ACUTE (2) BLUNT (3) ROUNDED (4) BIFID AND BLUNT 121* LENGTH INNER SPUR ON COXA III* 122* BASAL WIDTH INNER SPUR ON COXA III* 123* SHAPE OUTER SPUR ON COXA III* £1 ) ACUTE £2) BLUNT 124* LENGTH OUTER SPUR ON COXA III* 125* BASAL WIDTH OUTER SPUR ON COXA III* 126o LENGTH SETA ON COXA IV* 127* SHAPE SPUR ON COXA IV* (0) ABSENT (1) LONG* SLENDER* AND ACUTE ' (2) SHORT* BROAD* AND ACUTE (3) BLUNT £4) BIFID AND BLUNT 128* BLUNT APOPHYSIS ON FEMUR II* £0) ABSENT £ 1 ) PRESENT 129* FORM OF SETAE AVI AND PV1 ON TARSUS II* £0) NORMAL* LONG AND SLENDER £1) STOUT AND CLAW-LIKE 130* FORM OF SETAE AV2 AND AV3 OF TARSUS II* (0) NORMAL* LONG AND SLENDER £ 1) LONG AND SLENDER BUT BULBOUS BASALLY 131* FORM OF SETA AVI ON TARSUS IV* £0) NORMAL* LONG AND SLENDER £1) ACUTELY SPINIFORM £2) BLUNTLY SPINIFORM 199 APPENDIX II - CONTINUED
132* FORM OF SETA PV1 ON TARSUS IV* (0) NORMAL* LONG AND SLENDER (1) BLUNTLY SPINIFORM 133* SETA PD2 ON GENU IV* <0> ABSENT (1> PRESENT 134 * SETA PL2 ON TIBIA IV* (0) ABSENT tl) PRESENT 135* GREATEST WIDTH COXA I* 136* LENGTH COXA I (M I D-VENTRALLY)• 137* LENGTH TIBIA I ( MID-DORSALLY)• 138* LENGTH TARSUS I* 139* GREATEST WIDTH COXA II* 140* LENGTH COXA II* 141* LENGTH TIBIA II* 142* LENGTH TARSUS II* 143* GREATEST WIDTH COXA III* 144* LENGTH COXA III* 145* LENGTH GENU III* 146* LENGTH TIBIA III# 147* LENGTH TARSUS III* 148* GREATEST WIDTH COXA IV* 149* LENGTH COXA IV* 150* LENGTH GENU IV* 151o LENGTH TIBIA IV* 152* LENGTH TARSUS IV* APPENDIX HI
ONE-HUNDRED AND FIVE OTU BY ONE-HUNDRED AND FIFTY-TWO CHARACTER HIRSTIONYSSUS BASIC DATA MATRIX# CONTINUOUS CHARACTER VALUES (MEASUREMENTS) ARE IN MICRONS#
OTU CHARACTERS 1 2 3 4 5 6 7 8 9 10 1 1 12 13 14 GEOMYID HOST GROUP
FEMU 1 78 68 21 20 1 09 58 19 15 19 44 17 13 24 32 FEMU 2 80 68 21 20 108 61 17 14 20 41 18 1 1 25 31 f e m u 3 76 71 27 21 107 60 21 15 18 49 21 17 25 33 FEMU 4 81 79 28 20 123 67 20 15 21 44 19 16 24 34 GEOM 1 62 61 24 20 100 50 18 13 16 29 10 1 1 20 23 GEOM 2 82 63 19 19 100 52 19 17 18 39 17 13 21 27 LONG 1 75 68 19 21 112 60 22 22 20 35 19 12 20 33 LONG 2 74 67 19 19 1 1 0 56 20 18 16 33 17 12 19 37 LONG 3 61 69 26 19 122 56 23 21 18 37 19 13 20 34 THOM 1 82 71 19 23 1 00 54 22 1 1 15 40 18 11 20 35 THOM 2 70 66 27 19 100 52 18 12 13 43 19 15 IS 35
HETEROMYIDHOST GROUP h i l l 1 66 58 20 17 84 45 16 12 13 37 16 10 17 29 HILL 2 72 60 22 19 106 48 13 1 0 10 39 17 1 1 19 32 h i l l 3 62 57 23 18 95 46 15 1 1 13 37 18 12 17 32 HILL 4 74 61 19 18 1 00 40 13 12 14 42 20 12 21 35 HILL 5 71 61 19 19 1 03 49 1 5 1 1 13 35 17 10 20 30 INCO 1 70 71 26 20 1 13 49 21 18 16 48 21 14 22 37 INCO 2 70 65 24 20 1 16 49 19 16 14 45 20 15 21 37 INCO 3 72 67 27 20 1 15 51 18 16 16 47 21 13 19 38 TRIA 1 75 70 25 20 1 1 1 49 21 20 18 46 21 16 21 36 TRIA 2 81 70 23 21 1 08 50 23 18 18 42 20 12 20 38 TR1 A 3 81 69 26 21 1 15 53 21 18 16 42 21 16 21 38
SC IURID HOST GROUP
AFFI 1 84 74 29 22 93 49 23 19 15 46 22 16 22 35 AFFI 2 84 77 29 22 1 02 55 27 18 13 49 21 17 23 38 AFFI 3 ‘ 84 77 30 22 1 00 56 26 22 15 50 22 19 24 37 AFFI 4 88 82 32 22 103 58 2S 19 16 46 21 17 23 38 CYNO 1 83 80 30 22 99 49 22 16 13 50 24 18 20 30 INVA 1 78 72 30 24 93 42 22 14 12 54 20 17 21 38 200 201 APPENDIX III - CONTINUED
OTU CHARACTERS 1 2 3 4 5 6 7 Q 9 10 1 1 12 13 14
INVA 2 83 73 29 22 90 46 20 15 12 52 21 17 19 34 ANGU 1 78 71 28 22 91 47 21 19 13 47 19 17 19 39 NSPA 1 71 56 13 19 92 40 12 12 14 34 16 12 15 27 NSPA 2 72 62 17 19 1 00 41 13 12 16 34 15 1 1 18 28 EUTA 1 78 72 30 24 I 02 52 22 14 12 45 23 18 22 33 EUTA 2 79 79 32 23 95 53 26 20 17 50 22 18 24 41 OCCI 1 80 74 29 22 81 40 23 16 16 51 24 20 22 35 PUNC 1 97 75 29 23 101 53 21 13 IS 50 25 17 23 37 PUNC 2 81 80 33 24 107 54 22 19 17 47 22 17 21 38 PUNC 3 95 75 31 24 1 07 54 21 19 16 49 26 18 23 37 PUNC 4 74 72 29 22 83 41 22 20 16 52 24 19 22 33 TORU 1 72 72 27 21 1 08 54 18 13 10 43 21 16 21 37 TORU 2 71 70 26 21 106 54 19 13 1 1 42 21 16 IS 37 TORU 3 88 79 28 20 122 54 18 IS 1 1 43 20 13 21 38
MUROID HOST GROUP
BISE I 83 78 17 21 95 42 24 21 12 40 22 14 19 34 BISE 2 77 70 17 20 92 45 25 21 12 39 20 14 17 34 BISE 3 78 63 17 19 90 40 23 20 12 36 20 13 19 34 BREV 1 67 57 20 18 84 41 20 17 1 0 32 18 10 21 31 BREV 2 59 64 25 17 89 52 19 17 13 37 23 17 21 31 BREV 3 50 59 24 18 90 44 18 14 13 36 19 14 17 32 I SAB 1 79 75 29 23 102 49 21 16 12 49 22 18 18 40 I SAB 2 97 74 23 23 I 06 53 21 18 16 43 20 17 24 40 I SAB 3 78 72 26 21 99 48 21 17 14 44 21 19 19 36 I SAB 4 74 73 27 22 98 48 23 17 14 47 22 18 20 41 I SAB 5 75 70 20 20 1 14 51 21 16 14 37 18 13 20 37 I SAB 6 90 85 33 26 109 53 21 19 16 53 24 20 24 49 I SAB 7 96 89 25 23 I 17 55 23 20 16 49 23 15 25 45 I SAB 8 88 94 33 24 1 15 51 23 20 16 52 23 19 24 47 I SAB 9 81 79 29 22 1 14 49 23 20 15 47 22 16 24 40 ISAB 10 96 81 31 25 1 20 54 21 18 16 46 25 18 21 43 I SABI I 86 SO 29 24 1 17 58 23 20 1 3 47 21 18 23 40 I SAB 12 81 78 29 21 1 1 1 52 22 20 13 46 22 16 21 43 I SAB 13 89 84 31 24 1 14 52 22 21 15 48 27 18 23 45 I SAB 14 94 87 25 20 1 16 56 23 23 17 52 24 17 26 48 I SAB 15 82 80 29 22 1 1 1 50 27 21 16 48 21 17 21 43 I SAB 16 83 81 27 21 n o 51 22 18 14 43 21 17 22 40 I SAB 17 81 78 20 22 1 12 55 1 5 20 15 48 24 18 20 40 ISA818 81 78 30 23 1 12 53 23 20 15 50 24 17 21 43 I SAB 19 77 69 27 22 I 09 53 23 21 1 1 50 23 17 18 38 ISAB20 79 75 28 22 1 12 58 19 16 15 49 21 16 23 39 LATI 1 89 83 31 22 1 02 47 25 20 16 52 26 17 28 43 LATI 2 88 87 3 1 23 103 47 27 21 15 53 25 18 27 45 LATI 3 90 80 3 1 26 1 03 45 25 20 16 40 21 10 31 43 202 APPENDIX I I I CONTINUED
OTU CHARACTERS 1 2 3 4 5 6 7 8 9 10 1 I 12 13 14
LATI 4 83 83 33 22 111 41 26 20 17 51 25 19 28 41 LATI 5 83 85 26 22 1 06 46 23 19 18 49 21 13 24 46 NEOT 1 65 61 21 17 91 39 19 18 1 I 35 20 12 18 .28 NEOT 2 79 61 17 20 95 42 21 18 15 34 18 13 20 29 NEOT 3 65 62 22 17 94 40 22 17 1 1 37 20 13 19 30 NEOT 4 72 62 22 18 93 39 19 16 12 33 18 14 19 29 TARS 1 82 71 22 22 93 51 22 18 12 43 20 13 21 37 UTAH t 87 72 28 21 1 02 50 24 18 15 42 21 13 22 39 UTAH 2 71 75 29 21 103 49 25 13 16 37 21 15 23 38 UTAH 3 76 70 27 21 98 50 25 20 15 43 22 13 19 40 UTAH 4 76 73 26 19 97 54 28 22 15 42 21 15 21 40 UTAH 5 63 70 24 19 1 06 48 22 16 9 41 22 14 20 36 UTAH 6 89 78 26 26 1 00 51 27 21 15 39 22 16 21 40 UTAH 7 71 72 27 20 95 47 21 19 15 39 23 16 19 37
INSECTIVORE HOST GROUP
CARN I 75 80 23 21 120 51 19 16 13 42 23 14 23 41 NSP8 1 76 67 26 21 93 44 13 14 1 1 43 23 17 25 32 NSPB 2 83 71 23 22 94 46 19 15 1 1 39 16 12 17 37 NSPB 3 77 63 21 24 93 44 18 12 10 39 18 1 1 17 33 TALP 1 72 65 24 22 93 45 22 15 12 47 20 13 19 34 TALP 2 79 72 26 22 89 41 19 13 11 49 22 16 20 36 TALP 3 83 72 25 22 94 46 23 17 12 43 20 12 21 39 TALP 4 75 62 24 22 91 48 17 13 11 42 22 16 19 32 TALP 5 77 61 23 20 88 43 14 13 9 3a 18 1 1 16 31 TALP 6 83 66 20 21 93 42 16 1 1 8 37 16 9 16 32 TALP 7 79 67 23 22 95 46 17 12 1 1 33 18 12 17 31 TALP 8 68 58 23 20 90 39 17 14 9 31 16 1 I 17 35 TALP 9 91 67 22 24 96 47 24 17 12 39 24 13 20 36 PALU 1 75 70 27 23 90 45 20 15 13 45 22 14 18 . 38 OBSO 1 86 70 23 20 1 1 I 49 21 IS 13 44 21 . 12 26 38 OBSO 2 87 67 24 22 1 10 59 20 15 13 46 22 14 25 38
CARN IVORE HOST GROUP
STAF 1 67 67 24 20 95 34 15 6 9 38 14 12 20 26 STAF 2 72 71 23 21 1 03 36 16 9 10 34 12 9 20 27 STAF 3 71 79 25 21 1 09 34 16 8 13 38 16 15 19 29 STAF 4 63 60 21 20 88 32 15 a 9 33 9 1 1 -18 25 203 APPENDIX III - CONTINUED
OTU CHARACTERS 15 16 17 18 19 20 21 22 23 24 25 26 27 28
GEOMYID HOST GROUP
FEMU 1 33 13 2 25 31 78 78 1 33 1 5 46 49 50 41 FEMU 2 34 13 2 20 27 67 76 127 1 5 47 47 42 39 FEMU 3 39 12 2 27 31 75 81 141 1 5 47 48 44 44 FEMU 4 44 13 2 28 27 80 67 152 1 5 42 43 40 38 GEOM 1 34 15 2 14 21 70 56 1 14 3 5 38 32 35 31 GEOM 2 34 16 2 23 25 63 59 1 14 2 4 34 33 34 33 LONG 1 34 14 2 21 31 70 83 132 3 2 32 33 33 33 LONG 2 32 16 2 21 29 71 79 127 2 2 31 23 27 29 LONG 3 33 14 2 21 30 73 76 125 3 2 42 29 29 30 THOM 1 35 IS 2 25 28 73 64 142 1 5 54 51 47 51 THOM 2 33 15 2 22 28 71 61 131 1 5 48 45 39 41 h e t e r o m y ID HOST GROUP h i l l I 27 14 2 21 23 63 34 98 2 6 31 27 21 26 h i l l 2 28 17 2 23 29 66 36 1 16 3 6 21 22 22 24 h i l l 3 23 12 2 21 29 69 36 104 3 6 30 27 20 26 HILL 4 23 14 2 22 29 68 32 106 3 6 27 24 21 .20 HILL 5 31 16 2 22 29 73 41 106 3 6 32 29 27 24 INCO 1 34 30 3 21 31 81 20 121 2 7 30 34 33 33 INCO 2 32 19 2 20 29 80 15 1 18 2 7 30 31 26 27 INCO 3 30 19 2 22 28 83 16 125 2 7 31 24 29 28 TRIA 1 33 26 3 21 32 78 26 1 16 I 6 47 47 44 44 TRIA 2 32 21 2 20 31 78 23 108 3 6 42 46 46 52 TRIA 3 33 21 2 22 30 75 20 110 3 7 46 44 44 44 SCIUR ID HOST GROUP
AFFI 1 33 15 2 22 32 83 38 1 17 4 6 42 42 42 36 AFFI 2 39 17 2 25 37 89 35 I 23 4 6 42 45 47 38 AFFI 3 39 15 2 27 34 90 36 127 3 6 40 42 44 36 AFFI 4 41 13 2 28 38 87 35 1 19 3 6 40 46 43 39 CYNO 1 33 15 2 23 30 84 23 126 3 7 39 39 44 40 INVA 1 33 16 2 25 31 83 25 132 3 7 35 37 37 36 INVA 2 37 16 2 22 32 83 24 128 3 7 36 35 36 34 ANGU 1 34 14 2 24 32 82 25 1 15 3 7 36 37 39 33 NSPA 1 23 16 2 24 26 53 1 7 1 14 3 7 25 28 27 19 NSPA 2 32 14 2 22 26 60 19 1 18 3 7 24 26 26 20 EUTA I 37 16 2 27 34 84 42 1 19 4 6 3a 40 44 39 EUTA 2 37 15 2 22 34 87 34 1 24 3 6 37 39 43 34 OCCI 1 38 IS 2 24 32 86 39 126 3 6 39 42 45 40 PUNC 1 41 17 2 27 33 82 41 128 3 6 43 45 53 46 PUNC *2 39 18 2 31 36 82 39 124 3 6 46 46 51 44 PUNC 3 40 16 2 21 32 87 41 128 3 6 43 43 50 45 APPENDIX III - CONTINUED
c h a r a c t e r s 15 16 17 18 19 20 21 22 23 24 25 26 28
33 15 2 21 27 78 38 121 3 6 45 47 45 34 11 2 21 36 91 34 110 3 6 29 29 35 35 12 2 23 34 89 32 1 09 3 6 29 29 32 35 13 2 27 32 84 32 1 17 3 6 29 30 33
«OST GROUP
42 15 2 24 32 78 50 1 17 2 5 40 40 34 36 16 2 25 31 80 42 1 14 3 6 39 41 36 33 14 2 23 32 75 51 115 2 6 42 39 37 29 12 2 21 30 72 27 93 2 5 27 27 28 31 14 2 21 28 67 28 94 2 6 30 30 31 30 14 2 19 28 66 33 98 3 6 28 30 29 37 IB 2 23 32 83 35 1 14 4 6 42 43 37 41 15 2 22 35 90 31 1 13 4 7 48 47 33 35 15 2 23 29 83 35 1 14 3 6 40 42 35 33 18 2 23 30 89 30 1 10 3 6 46 46 33 36 13 2 22 32 79 29 109 2 6 44 42 35 39 15 2 22 28 83 37 130 3 6 58 53 45 46 17 2 25 39 93 32 121 2 6 51 52 40 42 17 2 27 39 96 40 137 1 6 55 53 47 38 18 2 22 38 96 32 121 3 6 46 48 45 42 20 2 25 36 93 33 125 2 6 47 49 40 35 17 2 20 35 92 42 120 3 6 49 51 47 35 16 2 23 33 88 42 1 17 3 6 44 44 38 41 18 2 24 37 91 39 1 30 3 6 49 51 45 41 16 2 26 40 95 43 1 30 3 6 48 49 41 36 14 2 24 36 93 33 121 3 6 45 48 42 38 15 2 23 33 83 34 1 16 2 6 42 47 42 37 14 2 27 37 91 34 159 2 6 47 48 42 36 18 2 23 32 93 37 122 3 6 49 47 44 35 14 2 22 38 89 30 124 3 6 46 47 4 1 35 15 2 23 31 93 40 1 10 3 6 44 45 36 39 21 2 23 30 90 28 131 3 7 46 45 33 37 20 2 21 35 89 23 133 3 7 43 45 39 34 14 2 24 33 88 25 1 15 2 7 42 43 42 41 16 2 28 37 83 28 1 33 1 6 48 46 41 41 22 2 24 34 91 27 126 2 6 41 44 39 29 14 2 19 26 71 36 1 02 3 6 41 41 36 31 13 2 18 20 78 37 1 15 3 6 39 39 34 34 12 2 21 28 70 32 105 3 6 39 39 36 32 13 2 24 28 73 36 103 3 6 34 36 29 34 14 2 20 34 74 25 109 3 6 32 33 30 37 15 2 24 37 73 26 1 16 3 6 34 34 31 36 22 2 22 35 88 27 1 14 3 6 36 35 30 36 18 2 24 32 79 26 11 1 3 6 33 34 31 APPENDIX III - CONTINUED 20 5
OTU CHARACTERS IS 16 17 18 19 20 21 22 23 24 25 26 27 28
UTAH 4 34 17 2 22 32 83 28 120 3 6 37 37 33 33 UTAH 5 33 19 2 21 35 86 26 104 3 6 32 34 30 30 UTAH 6 39 14 2 24 37 89 30 124 3 6 41 41 37 38 UTAH 7 33 ia 2 23 34 81 27 1 12 3 6 38 38 34 36 INSECT IVORE HOST GROUP
CARN 1 34 8 1 23 34 84 72 122 4 5 41 47 45 38 NSPB 1 36 18 2 25 29 77 39 109 3 6 43 48 48 35 NSPB 2 31 15 2 24 31 78 42 122 4 6 48 52 49 39 NSPB 3 33 15 2 24 33 74 37 I 09 3 6 40 47 44 39 TALP 1 33 20 2 24 28 84 32 127 4 7 44 49 52 45 TALP 2 33 14 2 23 26 SO 25 1 14 4 7 44 45 44 34 TALP 3 34 19 2 25 33 84 33 121 3 7 53 51 51 42 TALP 4 33 12 2 21 30 79 40 120 4 7 38 42 41 31 TALP 5 30 14 2 21 25 73 34 1 10 4 6 39 38 42 29 TALP 6 31 16 2 20 28 72 3a 1 12 4 6 38 38 41 31 TALP 7 31 15 2 16 21 70 35 1 09 4 6 42 37 48 23 TALP 8 24 14 2 19 24 76 38 1 07 3 6 39 35 38 33 TALP 9 32 14 2 23 26 78 38 134 4 7 40 46 52 43 PALU 1 30 13 2 23 29 82 39 1 17 4 6' 49 49 56 44 OBSO 1 31 15 2 23 34 88 28 120 3 7 49 50 49 36 OBSO 2 33 15 2 23 33 86 37 125 3 6 51 52 49 37 CARN I VORE HOSTGROUP
STAF 1 31 10 1 21 21 65 33 1 24 3 4 42 36 34 19 STAF 2 35 10 1 22 24 70 37 127 3 4 45 43 37 22 STAF 3 34 12 1 18 27 67 38 125 2 4 33 36 34 22 STAF 4 27 12 1 17 28 50 33 1 15 3 4 36 33 39 17 206 APPENDIX III - CONTINUED
OTU CHARACTERS 29 30 31 32 33 34 35 36 37 38 39 40 41 42 GEOMYID HOST GROUP
FEMU 1 68 129 158 51 33 1 113 1 13 1 133 113 4 34 31 FEMU 2 61 121 149 45 32 1 108 107 2 1 17 1 04 3 33 30 FEMU 3 68 136 164 50 38 1 136 143 2 136 117 2 35 32 FEMU 4 72 148 178 53 31 1 118 123 2 149 109 4 33 29 GEOM 1 59 107 118 35 26 1 98 100 2 138 110 3 28 28 GEOM 2 60 109 125 38 30 1 91 95 2 94 96 2 28 23 LONG 1 61 119 143 46 24 1 92 94 2 1 19 94 4 21 21 LONG 2 59 115 144 46 24 1 98 98 1 1 29 96 4 17 16 LONG 3 57 118 143 43 26 1 1 08 1 09 2 124 100 3 17 15 THOM I 68 134 161 44 33 1 107 1 16 2 144 109 4 41 41 THOM 2 69 1 1 1 142 42 29 1 102 I 03 2 127 98 4 33 31
HETEROMYID HOST GROUP h i l l 1 58 98 112 30 18 1 83 94 2 124 90 4 24 21 HILL 2 67 115 143 35 29 1 88 105 2 121 91 4 28 23 HILL 3 65 103 118 30 19 1 83 91 2 127 89 2 27 23 h i l l 4 62 100 113 28 21 1 82 93 2 122 89 3 20 19 HILL 5 66 11 1 120 30 21 1 90 98 2 134 94 4 27 23 INCO 1 63 117 150 36 30 1 89 100 3 156 104 4 30 23 INCO 2 65 1 15 138 33 22 1 92 102 2 149 102 4 23 21 INCO 3 70 1 16 139 35 26 1 88 102 2 143 94 4 26 22 TRIA 1 63 1 10 134 33 23 1 68 86 3 144 103 3 45 39 TRIA 2 56 99 118 30 24 1 76 90 2 136 94 4 46 39 TRIA 3 61 103 126 28 26 1 78 93 2 154 104 4 48 39
SCIURID HOST GROUP
AFFI 1 61 1 08 133 39 30 1 94 1 03 3 166 99 4 30 26 AFFI 2 65 I 15 148 39 35 1 105 114 2 171 117 4 31 31 AFFI 3 69 122 157 38 31 1 110 1 16 2 164 1 18 3 27 27 AFFI 4 60 112 144 37 30 1 1 04 1 1 0 2 171 113 2 28 2a CYNO 1 63 120 156 37 35 1 112 1 17 2 181 126 4 27 24 INVA 1 70 133 170 41 33 1 108 1 1 0 2 175 1 16 4 27 27 INVA 2 69 125 156 40 33 1 112 1 13 2 171 112 4 23 24 ANGU 1 60 n o 139 37 30 1 99 100 2 167 110 4 26 24 NSPA 1 61 114 140 35 21 1 87 92 2 159 106 4 15 11 NSPA 2 64 117 144 35 24 1 90 94 2 151 102 4 17 1 1 EUTA 1 59 117 1 50 42 31 1 104 1 13 2 181 114 4 27 26 EUTA 2 61 117 156 42 33 1 114 121 2 182 122 4 24 23 occi 1 68 1 19 156 38 35 1 124 125 2 173 1 16 3 31 32 PUNC 1 63 118 147 39 31 1 115 1 19 2 173 119 4 35 31 PUNC 2 66 126 160 42 33 1 1 09 115 2 168 120 2 35 31 PUNC 3 61 123 155 44 35 1 113 122 2 178 128 4 34 32 APPENDIX 111 - CONTINUED
CHARACTERS 29 30 31 32 33 34 35 36 37 38 39 40
61 114 143 38 33 1 103 107 2 185 120 4 29 47 102 140 37 33 1 107 109 2 174 123 2 23 46 105 142 37 33 1 106 1 15 2 172 120 4 25 50 108 142 37 30 1 109 121 2 170 112 2 24 HOST GROUP
52 105 122 42 30 92 1 03 3 163 110 1 23 51 100 117 37 30 89 1 12 2 1 53 112 1 29 47 104 122 45 28 87 98 2 148 108 1 29 40 85 112 31 27 80 81 2 136 83 4 26 43 83 106 31 29 87 90 2 161 108 3 29 51 94 1 1 1 36 27 87 90 2 152 97 2 26 64 108 138 38 32 1 I 0 108 I 184 114 5 30 62 115 144 42 31 114 116 2 201 125 4 31 64 1 1 1 134 39 28 94 108 2 176 102 5 24 59 107 136 35 32 101 102 2 186 117 5 33 65 107 139 32 33 100 108 2 177 114 5 30 69 120 155 37 40 110 123 2 233 148 5 39 58 116 153 42 42 1 15 1 27 2 235 151 5 37 79 133 172 39 37 128 1 34 2 207 132 5 42 63 113 143 35 32 108 1 17 2 193 130 5 39 71 121 150 39 34 124 1 28 2 199 141 5 31 72 117 146 39 32 115 124 2 193 131 5 36 66 114 140 38 30 1 13 119 2 182 122 5 36 74 123 157 41 33 119 123 2 217 142 5 37 70 125 167 43 38 129 133 2 221 134 5 31 65 118 146 36 31 99 1 13 2 192 124 5 37 61 1 1 1 140 38 32 106 1 12 2 185 1 1 1 5 33 80. 153 205 45 38 124 133 2 191 128 5 39 67 1 15 145 37 34 115 120 2 197 133 5 37 66 119 148 40 31 117 1 23 2 193 129 5 38 66 107 138 35 30 1 1 0 1 18 2 179 1 14 5 33 67 121 151 42 33 1 18 123 2 190 122 5 34 68 123 162 37 38 1 1 7 126 2 186 130 5 33 61 119 150 40 30 1 1 0 121 2 192 131 5 31 67 122 156 41 35 125 134 2 188 127 5 37 64 117 147 37 36 114 1 29 2 1 85 125 5 32 39 97 125 39 27 74 80 2 147 105 3 29 47 1 1 1 140 42 27 78 84 2 145 105 3 32 43 99 129 36 25 90 95 2 137 101 2 30 43 104 130 38 27 87 88 2 143 104 3 26 49 103 127 36 27 97 98 2 144 92 3 24 53 100 142 34 30 100 1 15 2 157 107 3 28 52 1 1 1 145 37 30 101 1 05 2 164 118 4 28 52 105 138 37 30 100 97 1 168 117 4 25 208 APPENDIX III - CONTINUED
OTU CHARACTERS 29 30 31 32 33 34 35 36 37 33 39 40 41 42
UTAH 4 53 106 136 37 32 1 103 109 2 170 113 4 31 28 UTAH 5 51 101 126 34 29 1 98 103 2 157 1 02 4 30 24 UTAH 6 59 114 148 37 32 1 1 13 117 2 164 114 4 32 30 UTAH 7 50 108 136 39 27 1 101 107 2 155 107 2 27 27
INSECTIVORE HOST GROUP
CARN 1 72. 136 161 51 31 1 95 109 2 165 116 3 38 39 NSPB 1 63 112 132 40 36 1 100 I 13 3 164 112 3 35 29 NSPB 2 68 130 168 43 35 1 301 109 2 162 1 17 3 33 34 NSPB 3 58 108 131 42 31 1 97 n o 2 164 114 3 37 32 TALP 1 69 119 132 39 37 1 76 89 3 184 114 4 38 . 34 TALP 2 68 112 149 39 38 1 101 112 3 195 138 3 33 34 TALP 3 70 119 148 41 37 1 1 00 122 3 185 126 4 37 33 TALP 4 68 113 153 40 35 1 95 I 16 2 169 122 4 29 29 TALP 5 61 104 131 36 38 1 89 104 2 164 114 4 24 24 TALP 6 57 106 142 38 41 1 94 105 2 173 122 4 27 26 TALP 7 60 103 132 38 36 1 90 1 04 2 159 120 4 26 26 TALP a 60 103 126 38 34 1 96 106 2 167 116 4 27 25 TALP 9 74 127 177 41 41 1 107 1 12 2 1 80 122 4 37 32 PALU i 69 HO 14 1 34 38 1 1 00 1 10 3 186 128 5 32 32 OBSO l 70 116 143 37 35 1 101 100 1 190 122 5 33 29 OBSO 2 71 134 162 45 33 1 108 n o 2 188 125 5 32 28
CARNIVORE HOST GROUP
STAF 1 52 114 136- 40 29 2 83 85 2 179 146 4 22 23 STAF 2 46 119 144 43 28 2 88 95 3 171 152 4 25 22 STAF 3 48 112 141 42 29 2 89 88 3 170 141 4 20 21 STAF 4 43 103 119 39 27 2 86 90 3 161 136 4 22 23 209 APPENDIX I I I - CONTINUED
OTU CHARACTERS 43 44 45 46 47 48 49 50 51 52 53 54 55 56 GEOMYID HOST GROUP
FEMU 1 3 97 74 53 31 23 29 3 22 47 21 10 38 25 FEMU 2 3 83 70 53 43 21 27 2 19 45 19 1 1 34 24 FEMU 3 3 119 74 57 33 20 29 3 21 44 '4 1 1 37 28 FEMU 4 2 104 71 59 37 26 31 3 22 45 27 I 1 34 27 GEOM 1 2 80 78 60 35 22 29 3 22 50 24 12 41 30 GEOM 2 2 79 64 48 31 20 27 3 17 40 19 10 31 25 LONG 1 2 76 70 59 36 18 29 2 16 40 24 16 31 20 LONG 2 2 86 68 59 35 17 29 3 14 42 21 10 34 15 LONG 3 2 94 71 60 38 17 29 3 16 44 22 1 1 34 16 THOM 1 2 99 80 61 37 30 33 3 26 44 29 14 33 34 THOM 2 2 93 73 56 32 24 30 3 21 43 23 1 1 33 23 h e t e r o m y i d HOST GROUP
HILL 1 3 73 62 47 29 19 27 3 18 37 22 10 33 19 HILL 2 2 83 67 50 34 19 29 4 16 39 23 14 34 22 HILL 3 3 73 61 46 32 21 26 3 18 39 17 9 31 21 HILL 4 3 79 65 46 28 21 27 3 16 39 22 a 27 17 HILL 5 2 80 76 52 33 20 30 3 18 44 22 12 39 19 INCO 1 3 76 83 62 45 24 33 3 22 47 30 15 32 22 INCO 2 2 80 77 60 39 19 31 3 19 48 26 12 39 20 INCO 3 2 82 78 58 40 17 31 3 22 47 27 12 33 21 TRIA 1 2 62 82 61 36 30 35 3 33 47 29 12 43 41 TRIA 2 2 69 74 60 38 29 32 3 36 42 27 12 39 38 TRIA 3 2 69 78 60 35 29 36 3 35 48 27 1 1 37 44
SCIURID HO:ST GROUP
AFFI 1 2 92 83 71 43 30 39 3 30 46 23 13 41 30 AFFI 2 2 94 89 69 44 31 39 4 33 50 30 12 44 32 AFFI 3 3 99 90 69 36 30 41 4 33 50 33 13 46 29 AFF I 4 2 90 83 66 37 28 39 3 31 47 23 13 40 29 CYNO 1 3 96 85 72 41 33 39 5 34 43 32 1 1 47 33 INVAI 3 94 80 71 45 30 41 3 33 45 28 13 38 27 INVA 2 2 98 77 71 45 29 37 3 27 43 27 13 39 27 ANGU 1 3 81 78 68 41 30 38 4 28 46 26 1 1 41 28 NSPA 1 3 78 80 63 45 19 31 3 16 56 21 i l 50 12 NSPA 2 3 86 89 65 49 20 31 3 17 55 29 16 48 1 0 EUTA 1 3 95 86 69 41 30 41 3 30 50 30 11 42 27 EUTA 2 3 109 89 71 39 23 37 3 26 52 34 14 41 29 OCCI 1 3 101 88 71 41 29 38 3 32 50 35 15 42 34 PUNCI 3 103 89 72 42 33 41 4 33 SO 34 14 41 38 PUNC 2 3 102 95 72 40 32 38 4 33 50 37 1 1 52 36 PUNC 3 3 103 89 72 44 32 41 3 30 52 32 13 44 37 APPENDIX III - CONTINUED
CHARACTERS 43 44 45 46 47 48 49 50 51 52 53
3 91 89 72 43 32 40 4 30 48 33 3 94 85 66 43 25 33 3 23 52 26 23 2 93 81 68 49 25 35 3 25 48 26 23 3 83 84 65 38 21 35 3 24 48 30 21
HOST GROUP
1 87 90 59 32 24 32 3 28 49 33 2 8 1 79 88 58 32 26 32 3 28 47 31 30 1 78 86 59 31 26 32 4 28 47 29 28 1 72 69 48 24 20 25 4 21 43 20 24 1 77 83 50 29 22 28 3 21 48 24 26 1 79 77 53 33 22 29 3 20 43 28 24 2 95 83 71 40 28 37 3 30 53 26 31 2 101 87 78 49 29 39 3 31 52 28 31 2 89 88 61 41 24 31 4 24 48 30 26 2 86 80 68 39 29 38 3 33 43 28 35 2 83 80 70 41 21 36 3 23 50 23 31 2 96 101 84 48 33 44 3 39 62 32 37 2 98 100 79 48 33 42 3 35 59 33 35 2 114 94 82 51 30 42 3 41 59 30 41 2 93 95 75 44 35 42 3 41 58 32 42 2 106 86 71 42 30 41 3 33 52 28 34 3 107 90 71 44 32 41 3 36 61 27 37 2 101 92 70 43 33 37 3 39 56 31 40 2 99 99 82 50 34 45 3 41 58 36 44 3 1 1 1 102 81 49 30 43 3 33 65 33 33 3 90 90 75 43 35 42 3 38 53 28 40 2 97 89 70 42 29 39 3 33 57 30 40 3 112 90 70 43 32 39 3 38 54 34 42 2 100 94 69 49 30 40 3 39 57 31 42 2 100 92 72 47 33 40 4 39 55 31 41 2 101 83 70 47 30 38 3 30 51 27 31 3 102 100 74 41 29 41 3 34 58 37 34 2 106 95 76 44 31 40 3 34 61 28 34 2 99 101 65 33 28 36 4 30 58 33 34 2 11 1 104 72 41 28 37 4 33 59 33 39 2 105 97 71 39 28 39 4 31 59 31 32 2 63 76 50 26 23 29 4 25 40 28 26 1 65 81 54 31 23 30 3 28 46 27 28 1 76 80 54 27 23 31 4 27 47 26 28 1 72 76 53 26 23 30 3 26 43 27 26 2 79 73 59 30 22 33 3 23 46 24 22 2 98 77 65 33 24 36 3 25 46 27 25 2 89 82 65 38 27 37 3 28 49 29 23 2 85 78 59 34 23 30 4 26 44 28 24 211 APPENDIX III - CONTINUED
OTU CHARACTERS 43 44 45 46 47 48 49 50 51 52 53 54 55 56 UTAH 4 2 92 78 59 34 26 31 4 28 46 28 1 1 42 30 UTAH 5 2 89 74 56 31 23 30 4 24 43 26 9 40 23 UTAH 6 2 96 80 69 36 30 39 4 31 41 30 13 40 31 UTAH 7 2 87 73 56 31 27 30 3 31 42 26 1 1 37 29 INSECT I VORE HOST GROUP
CARN 1 3 82 101 79 40 28 38 2 38 71 25 16 56 37 NSPB I 2 90 89 77 41 27 38 4 28 53 30 13 50 27 NSPB 2 2 88 89 76 40 29 41 3 27 54 30 12 48 30 NSPB 3 2 84 85 78 41 23 39 3 29 52 27 13 44 29 TALP I 2 70 94 78 49 24 40 3 35 63 28 1 1 55 31 TALP 2 2 86 101 80 56 24 44 3 27 67 31 12 58 39 TALP 3 2 89 92 80 51 26 43 3 27 60 26 12 S3 32 TALP 4 2 88 98 77 49 30 43 4 31 65 32 1 1 57 27 TALP 5 2 84 73 69 39 22 34 4 26 44 27 1 0 40 20 TALP 6 2 E4 78 70 40 23 37 3 23 50 23 13 41 21 TALP 7 2 83 78 67 41 23 35 3 25 50 23 12 41 23 TALP 8 2 83 82 67 44 21 37 3 24 53 24 12 44 21 TALP 9 2 87 88 73 42 28 38 3 28 55 26 1 1 51 29 PALU I 2 88 84 78 48 26 42 3 26 56 23 12 43 25 OBSO 1 2 92 81 75 43 26 36 3 31 45 29 15 39 30 OBSO 2 2 97 80 74 45 28 36 3 30 50 28 15 30 28
CARN IVORE HOST GROUP
STAF 1 3 77 67 57 44 18 27 1 18 48 1 o 7 28 25 STAF 2 3 80 69 62 44 19 26 1 17 50 1 2 8 29 30 STAF 3 3 77 69 62 46 20 23 1 21 51 I 1 7 23 26 STAF 4 3 77 73 62 41 18 33 1 19 47 IS 5 27 30 212 APPENDIX III - CONTINUED
OTU CHARACTERS 57 58 59 60 61 62 63 64 65 66 67 68 69 70
GEOMYIO HOSTGROUP
FEMU 1 25 1 216 2 1 479 248 1 2 3 1 2 2 2 FEMU 2 23 1 203 1 1 445 230 1 2 3 1 2 2 2 FEMU 3 27 1 221 1 1 520 265 1 2 3 1 2 2 2 FEMU 4 30 1 223 1 1 487 267 1 2 3 1 2 2 2 GEOM 1 28 1 201 1 1 460 229 2 2 3 1 1 - I * 1 GEOM 2 22 1 192 1 1 418 216 3 1 3 1 1 1 1 LONG 1 16 1 199 2 1 443 252 2 2 2 1 _ 2 2 2 LONG 2 13 1 197 1 1 44 0 249 1 2 2 --1 2 2 2 LONG 3 15 1 195 1 1 444 277 3 2 2 1 2 2 2 THOM 1 38 1 239 1 1 492 272 1 3 3 1 2 2 2 THOM 2 25 1 220 1 1 478 259 1 3 3 1 2 2 2
HETEROMYID HOST GROUP
HILL 1 18 0 148 3 2 393 195 2 2 3 2 2 2 HILL 2 18 0 144 3 2 418 202 3 2 3 2 2 2 h i l l 3 21 0 142 3 2 399 190 1 2 3 2 2 2 HILL 4 19 1 143 3 1 430 21 0 1 2 3 2 2 2 h i l l 5 17 1 143 3 1 443 220 1 2 3 2 2 2 INCO 1 24 1 198 2 1 493 231 1 2 3 2 2 2 INCO 2 20 1 195 1 I 480 236 1 2 3 2 2 2 INCO 3 21 1 194 1 1 456 236 1 2 3 2 2 2 TRI A 1 40 2 195 1 1 478 209 1 2 3 2 2 2 TRIA 2 42 2 203 1 1 449 214 1 3 3 2 2 2 TRI A 3 42 2 206 1 1 455 218 1 3 3 2 2 2 SC I UR ID HOSTGROUP
AFFI 1 33 1 119 4 1 505 296 3 2 2 2 2 2 AFFI 2 38 1 184 3 1 572 327 3 2 3 2 2 2 AFFI 3 35 1 202 3 1 560 332 3 2 2 2 2 2 AFFI 4 34 1 208 2 1 560 340 1 2 2 2 2 2 CYNO 1 34 1 228 2 1 557 303 3 2 3 2 2 2 INVA 1 36 1 238 2 1 554 300 3 2 3 2 2 2 INVA 2 33 1 245 2 1 560 282 3 3 3 . 2 2 2 ANGU 1 32 1 225 2 1 540 288 1 3 2 2 2 2 NSPA 1 17 1 182 2 1 461 228 3 2 2 2 2 2 NSPA 2 16 1 195 2 1 472 208 3 2 3 2 2 2 EUTA 1 35 1 178 3 1 531 337 2 2 3 2 2 2 EUTA 2 30 1 182 3 1 568 318 3 3 3 2 2 2 OCCI 1 36 1 179 3 1 552 300 3 2 3 2 2 2 PUNC 1 41 1 194 3 1 549 331 3 2 3 2 2 2 PUNC 2 39 1 202 3 1 578 334 3 2 3 2 2 2 PUNC 3 40 1 195 3 1 575 327 3 2 3 2 2 2 APPENDIX I I I - CONTINUED
CHARACTERS 57 58 59 60 61 62 63 64 65 66 67
35 1 170 3 1 538 324 3 2 2 1 30 1 205 1 1 547 293 2 3 2 1 32 1 210 2 1 525 281 3 2 2 1 2 33 1 226 1 1 548 281 3 2 3 1 2 *OST GROUP
27 1 194 1 1 488 272 3 2 2 2 27 1 180 1 1 465 250 3 2 2 2 28 1 194 1 1 459 252 3 2 2 2 19 1 194 2 1 414 229 2 3 2 2 20 1 185 2 1 446 262 3 2 1 2 22 1 187 2 1 443 244 1 3 2 2 31 1 229 1 1 524 298 3 3 1 2 32 1 242 1 1 555 334 3 3 2 2 29 1 219 1 1 507 285 3 2 2 2 40 1 224 1 1 528 281 2 3 2 2 40 1 228 1 1 521 303 3 2 3 2 43 1 277 1 1 672 374 3 2 2 2 44 1 275 1 1 666 382 3 3 2 2 SO 1 265 1 1 646 378 3 2 3 2 50 1 236 1 1 568 318 3 2 2 2 40 1 247 1 1 620 342 3 2 3 2 41 1 24 4 1 584 345 3 2 2 2 49 1 222 1 1 547 306 3 2 2 2 51 1 254 1 1 628 340 3 2 2 2 37 1 273 1 1 651 384 3 2 3 2 47 1 244 1 1 564 312 2 3 2 2 42 I 24 1 1 1 552 306 3 2 2 2 43 1 255 1 1 565 306 3 2 3 2 45 1 234 1 1 560 301 2 3 2 2 46 1 246 1 1 570 316 3 2 3 2 34 1 244 1 1 550 303 3 2 2 2 41 1 234 1 1 604 332 3 3 3 2 34 1 242 1 1 571 309 3 2 3 2 41 1 236 1 1 586 326 3 2 2 2 43 1 240 1 1 601 340 3 2 2 2 41 1 246 1 1 578 326 3 2 2 2 25 1 170 1 427 21 1 3 2 2 2 27 1 183 1 1 420 217 3 2 2 2 28 1 177 1 1 416 225 3 2 2 2 32 1 179 I 1 425 21 0 3 2 3 2 26 1 205 1 1 473 285 3 3 2 2 27 1 214 1 1 506 295 3 2 2 2 28 1 203 1 1 515 300 3 3 2 2 26 1 201 1 1 488 286 3 2 2 2 APPENDIX III - CONTINUED
OTU CHARACTERS 57 58 59 60 61 62 63 64 65 66 67 68 69 70
UTAH 4 31 1 206 1 1 500 290 3 2 2 1 2 2 2 UTAH 5 26 1 197 1 1 458 264 3 2 2 1 2 2 2 UTAH 6 37 1 220 2 1 526 318 3 2 2 1 2 2 2 UTAH 7 32 I 213 1 I 478 284 3 2 2 1 2 2 2 INSECTIVORE HOST GROUP
CARN 1 47 1 276 1 1 576 326 3 1 1 1 2 2 2 NSPB 1 39 1 236 1 1 500 312 3 1 1 1 0 2 0 NSPB 2 40 1 249 1 1 524 316 3 1 1 1 0 2 0 NSPB 3 41 1 244 1 1 518 306 3 1 1 1 0 2 0 TALP 1 49 1 233 1 1 544 297 3 2 2 1 2 2 0 TALP 2 49 1 255 1 1 581 337 3 1 2 1 2 2 0 TALP 3 42 1 256 1 1 562 316 3 2 2 1 2 2 0 TALP 4 44 1 244 1 1 526 296 3 2 1 1 2 2 0 TALP 5 31 1 226 1 1 485 250 2 2 3 1 2 2 0 TALP 6 33 1 239 1 1 498 255 1 2 3 1 2 2 0 TALP 7 34 1 228 1 1 499 255 1 2 3 1 2 2 0 TALP 8 30 1 215 1 1 467 249 3 1 3 1 2 2 0 TALP 9 34 1 245 1 1 539 293 3 1 3 1 2 2 0 PALU 1 37 1 228 1 1 521 298 3 2 2 1 2 2 0 OBSO 1 32 1 232 1 1 520 299 3 3 1 1 2 2 2 OBSO 2 32 1 235 1 1 541 294 2 2 1 1 2 2 2
CARNIVORE HOST GROUP
STAF 1 42 1 81 5 i soa 266 3 1 4 0 2 2 2 STAF 2 45 1 78 5 1 517 276 3 1 4 0 2 2 2 STAF 3 42 I 87 5 1 51 0 262 3 1 4 0 2 2 2 STAF 4 43 1 78 5 1 459 260 3 1 4 0 2 2 2 215 APPENDIX III - CONTINUED
OTU CHARACTERS 71 72 73 74 75 76 77 78 79 80 81 82 83 84 GEOMYID HOST GROUP
FEMU 1 I I 0 1 0 2 29 1 1 35 IS 31 14 21 21 FEMU2 1 1 0 1 0 2 30 1 1 37 15 31 13 21 19 FEMU 3 1 1 0 1 0 2 28 1 1 34 16 22 1 1 21 21 FEMU 4 1 1 0 1 0 3 28 10 34 19 27 1 5 17 22 GEOM I I 1 0 1 0 3 13 10 15 14 15 I 1 15 20 GEOM 2 1 1 0 1 0 3 IS 8 20 1 1 15 1 0 13 16 LONG 1 1 1 1 1 0 5 19 1 12418 21 15 1620 LONG 2 1 1 1 1 0 5 16 10 20 16 17 13 12 16 LONG 3 1 1 1 1 0 2 17 1 1 20 12 19 13 1 3 15 THOM 1 1 1 1 1 0 1 36 33 42 27 42 32 33 30 THOM 2 1 1 1 1 0 2 29 24 33 18 33 23 21 22 HETEROMYID HOST GROUP
HILL 1 1 1 1 1 0 3 33 26 34 2529 22 22 20 HILL 2 1 1 1 1 0 3 29 23 33 25 29 22 20 21 h i l l 3 1 1 1 1 0 3 32 28 34 30 33 22 26 22 HILL 4 1 1 1 1 0 2 31 18 29 28 27 16 19 16 HILL S 1 1 1 1 0 2 33 26 34 27 32 21 24 21 INCO 1 1 1 1 I 0 4 35 28 34 30 31 24 25 23 INCO 2 1 1 1 1 0 3 29 23 31 23 26 19 21 21 INCO 3 1 1 1 1 0 3 33 24 35 30 33 23 27 22 TRI A 1 1 1 1 1 0 2 48 47 48 46 48 46 46 46 TRI A 2 1 1 1 1 0 2 49 49 49 42 49 45 45 45 TRI A 3 1 1 1 1 0 3 49 50 50 44 50 47 48 44 SCIURID HOST GROUP
AFFI 1 1 1 1 1 0 3 25 16 30 28 23 17 24 27 AFFI 2 1 1 1 1 0 3 28 18 33 31 32 24 25 31 AFFI 3 1 1 1 1 0 3 28 16 33 28 30 19 24 27 AFFI 4 1 1 1 1 0 3 24 14 28 29 29 16 23 24 CYNO I 1 1 1 1 0 2 20 1 1 27 22 25 14 20 23 INVA 1 1 1 1 1 0 2 21 1 1 29 23 24 13 20 24 INVA 2 1 I 1 1 0 4 19 12 28 23 22 1 3 20 22 ANGU 1 1 1 1 I 0 5 21 13 21 23 23 14 20 22 NSPA 1 1 1 1 1 0 5 13 10 13 14 1 2 14 12 19 NSPA 2 1 1 1 1 0 5 12 10 13 15 12 13 1 1 18 EUTA I 1 1 1 I 0 2 28 16 32 22 29 16 22 24 EUTA 2 1 1 1 I 0 2 22 12 24 16 23 14 19 21 OCCI 1 I 1 1 1 0 2 28 16 34 23 32 17 23 26 PUNC 1 1 1 1 1 0 3 28 17 34 31 3 1 20 28 31 PUNC 2 1 1 1 1 0 3 30 19 37 31 32 18 27 30 PUNC 3 1 1 1 1 0 3 30 17 30 31 32 18 27 32 APPENDIX III - CONTINUED
CHARACTERS 71 72 73 74 75 76 77 78 79 80 81 82 84
1 1 1 1 0 2 30 17 34 31 32 17 30 1 1 1 1 0 3 18 9 22 17 20 9 23 1 1 1 1 0 3 17 9 22 20 21 10 22 1 1 1 1 0 3 19 1 1 26 21 21 9 22
HOST GROUP
O 1 31 24 33 28 32 21 23 0 1 31 24 33 28 29 22 23 0 1 29 21 33 28 29 21 22 0 2 18 1 1 19 17 16. 1 1 18 0 2 17 1 0 20 18 18 1 2 17 0 2 14 10 20 18 21 15 18 0 2 27 19 36 24 30 17 24 0 2 30 18 38 25 33 16 27 0 2 24 16 28 21 27 15 23 0 2 17 18 20 19 22 17 28 O 2 29 19 35 33 28 17 29 0 2 35 18 42 28 39 18 36 0 2 33 19 43 31 39 18 33 O 2 33 18 42 29 39 17 38 0 2 39 15 45 33 45 27 42 0 2 31 20 39 27 33 17 31 O 2 29 19 37 24 35 21 30 O 2 37 25 44 31 40 27 41 0 2 38 26 47 33 42 26 42 O 2 31 17 38 28 35 18 23 0 2 36 26 43 31 39 26 38 o 2 34 23 44 31 39 24 39 0 2 39 27 44 34 39 26 38 0 2 40 26 45 33 41 24 38 0 2 39 23 44 34 40 23 38 0 2 29 15 34 21 31 18 26 0 2 26 13 34 23 26 13 20 0 2 27 13 36 28 28 14 22 o 2 26 15 33 26 28 15 16 o 2 29 17 37 27 32 17 23 0 2 26 14 32 27 29 12 22 0 2 32 26 34 28 30 26 21 0 2 33 26 33 28 31 24 24 0 2 31 27 34 27 31 26 24 0 2 26 18 31 24 26 19 23 0 2 26 15 31 27 27 13 17 0 2 28 15 33 28 29 1 6 19 o 2 26 16 31 23 27 14 20 0 2 28 17 33 25 21 16 23 217 APPENDIX III - CONTINUED
OTU CHARACTERS 71 72 73 74 75 76 77 70 79 80 81 82 83 84
UTAH 4 1 1 1 I 0 2 28 18 34 29 23 14 14 20 UTAH S 1 1 1 1 0 2 26 20 32 23 29 19 20 19 UTAH 6 1 1 1 1 0 2 30 20 40 30 24 19 23 27 UTAH 7 1 1 1 1 0 1 28 20 36 29 25 19 22 26 INSECTIVORE HOST GROUP
CARN 1 0 1 1 1 0 0 15 7 30 12 17 8 12 13 NSPB 1 1 1 0 0 0 1 21 10 32 15 24 12 22 19 NSPB 2 1 1 0 0 0 2 21 1 1 34 16 29 12 23 21 NSPB 3 1 1 0 0 0 2 20 9 30 15 27 1 1 21 20 TALP 1 1 1 0 0 0 2 16 1 1 28 12 28 1 1 18 22 TALP 2 1 1 0 0 0 2 15 9 34 13 31 9 23 22 TALP 3 1 1 0 0 0 2 19 12 35 17 34 12 27 23 TALP 4 I 1 0 0 0 2 26 13 39 15 30 8 30 26 TALP 5 I I 0 0 0 1 19 12 24 1 1 15 1 I 15 13 TALP 6 1 1 0 0 0 1 19 12 26 12 16 1 1 18 13 TALP 7 1 1 0 0 0 1 18 12 25 1 1 16 1 0 19 16 TALP B 1 1 0 0 0 2 17 1 2 25 10 24 1 0 15 13 TALP 9 1 1 0 0 0 2 20 13 27 12 28 12 20 IS PALU 1 1 1 0 0 0 1 20 1 1 33 13 28 1 1 17 16 OBSO 1 1 I 0 I 0 2 34 20 40 22 38 19 28 23 OBSO 2 1 1 0 1 0 2 34 21 40 22 37 21 28 24 CARNIVORE HOST GROUP
STAF 1 1 0 0 0 1 3 23 9 24 23 23 13 25 24 STAF 2 1 0 0 0 1 3 22 1 1 24 24 25 17 30 28 STAF 3 1 0 0 0 1 3 21 1 1 22 28 21 13 20 24 STAF 4 1 0 0 0 I 3 21 1 0 23 28 25 ! 3 20 22 2X8 APPENDIX III - CONTINUED
OTU CHARACTERS as 86 87 88 89 90 91 92 93 94 95 96 97 98 GEOMYID HOST GROUP
FEMU 1 23 29 43 38 97 63 39 50 67 16 84 81 12 41 FEMU 2 22 28 47 37 96 58 34 50 48 13 81 91 14 42 FEMU 3 23 30 53 50 122 84 41 52 63 17 83 91 15 42 FEMU 4 23 37 58 41 111 63 40 49 69 19 90 91 13 57 GEOM 1 23 33 41 33 93 51 46 53 61 20 95 98 26 52 GEOM 2 13 34 42 31 89 48 33 46 55 17 80 87 26 40 LONG 1 23 29 49 37 1 00 71 41 54 90 20 92 105 35 59 LONG 2 22 35 46 37 93 63 38 52 67 16 83 94 34 61 LONG 3 19 37 50 40 1 07 85 45 63 68 21 93 117 45 66 THOMI 29 33 53 41 1 03 49 33 49 89 21 108 109 22 50 THOM 2 21 31 54 39 112 57 34 39 69 16 98 96 17 42 HETEROMYID HOST GROUP
HILL I 24 23 43 33 81 37 35 36 51 16 79 80 20 42 HILL 2 26 23 48 42 91 38 39 39 49 15 80 80 18 49 h i l l 3 26 23 41 37 87 37 37 31 48 13 69 69 18 40 HILL 4 21 27 59 47 92 49 42 38 46 16 82 83 19 43 HILL 5 23 26 40 38 98 61 42 43 53 15 71 85 23 41 INCO 1 24 28 45 37 92 44 42 51 68 31 93 91 25 44 INCO 2 22 27 48 42 96 43 47 44 67 33 95 89 22 62 INCO 3 24 23 44 43 99 42 44 49 66 28 94 93 20 59 TRI A 1 47 29 43 41 91 38 38 32 58 26 81 80 26 44 TRI A 2 47 28 43 45 91 37 39 33 61 24 86 81 30 50 TRI A 3 44 30 47 40 95 49 39 31 63 30 76 78 26 45 SCIURID HOST GROUP
AFFI 1 29 38 52 34 1 05 87 35 71 83 26 100 131 26 53 AFFI 2 31 43 52 43 108 89 48 78 94 26 1 13 136 39 53 AFFI 3 29 32 63 42 123 96 45 81 89 30 1 09 146 36 61 AFFI 4 29 46 61 4 1 1 16 87 43 77 93 23 114 146 42 58 CYNO 1 32 40 61 46 1 05 118 31 84 109 28 96 135 50 64 INVA 1 30 43 55 49 1 1 1 99 43 84 112 22 91 124 53 56 INVA 2 30 37 52 46 1 03 99 41 73 104 20 87 124 41 59 ANGU 1 2G 43 57 50 107 1 1 0 43 67 100 24 93 1 19 48 56 NSPA 1 22 37 41 37 84 53 37 42 55 17 76 89 31 55 NSPA 2 24 38 42 31 89 56 38 45 52 14 71 85 23 44 EUTA 1 27 38 63 45 1 19 1 13 46 78 93 30 1 03 141 29 62 EUTA 2 27 47 65 41 1 25 96 52 86 83 31 106 152 46 63 OCCI 1 33 43 65 40 110 108 48 62 83 25 100 131 24 66 PUNC 1 33 45 52 41 117 91 43 66 113 25 114 138 36 63 PUNC 2 33 45 51 48 121 1 02 37 69 103 23 112 147 36 63 PUNC 3 32 47 52 42 1 14 87 41 59 98 21 117 146 35 60 APPENDIX - CONTINUED
CHARACTERS 8 5 8 6 8 7 8 8 8 9 9 0 9 1 9 2 9 3 9 4 9 5 9 6
3 0 4 3 5 0 4 6 1 2 1 8 6 3 7 5 5 8 9 2 2 1 1 0 1 4 2 2 4 3 8 6 8 3 2 9 8 7 6 5 0 6 3 9 8 2 0 1 0 1 1 1 2 6 3 2 6 4 4 6 1 3 9 1 0 0 6 7 4 3 7 1 7 8 1 8 1 0 4 1 1 6 4 8 2 6 5 0 6 6 3 6 9 9 . 6 2 4 4 7 6 7 3 1 7 9 3 1 2 0 5 9
HOSTGROUP
2 1 2 9 4 6 3 9 9 2 4 9 4 4 6 1 7 9 1 5 1 0 2 1 1 1 6 3 2 3 3 3 5 1 3 7 8 9 5 1 4 1 5 3 6 7 1 7 9 2 1 0 8 6 1 2 2 3 1 4 8 3 7 9 1 4 7 3 7 4 8 6 1 1 9 9 4 1 0 6 5 3 2 0 2 2 4 5 3 5 9 0 6 7 2 7 5 0 5 9 1 4 7 8 1 0 0 5 7 2 1 3 4 4 2 3 8 8 9 6 9 3 4 7 3 7 4 1 6 9 0 1 0 9 5 8 2 3 3 2 4 8 3 7 9 0 7 7 3 2 6 7 5 1 1 9 8 6 1 0 2 5 6 2 6 4 3 6 0 3 7 9 0 9 7 4 2 8 3 7 5 2 0 9 8 1 2 7 6 5 2 8 4 1 5 5 4 2 1 0 5 9 7 5 3 8 2 8 7 3 3 1 2 1 1 4 2 7 2 2 2 4 2 5 0 3 4 9 3 9 0 4 0 4 9 8 6 1 9 9 4 1 1 8 7 2 2 7 4 1 s a 4 1 1 0 3 8 6 3 7 7 1 7 7 2 0 9 6 1 2 2 7 1 2 9 3 9 5 6 4 3 1 0 2 7 0 4 2 6 3 7 5 2 4 1 1 4 1 2 5 7 2 3 6 4 8 6 3 4 0 1 0 6 8 8 5 5 9 4 9 0 3 2 1 3 4 1 5 7 7 3 3 3 4 3 5 3 3 9 1 1 0 9 2 6 2 8 9 1 0 9 3 9 1 3 2 1 5 9 7 9 3 7 5 2 7 4 4 6 1 0 9 8 5 6 8 6 9 9 0 3 1 1 3 4 1 5 6 7 9 4 2 4 1 5 5 3 8 1 0 3 8 5 3 9 6 1 8 3 2 0 1 0 3 1 3 2 7 0 2 8 3 9 7 3 3 5 1 0 8 8 8 6 5 8 9 8 1 3 3 1 2 3 1 4 1 6 3 3 6 4 8 6 1 3 9 1 0 9 1 0 3 6 6 7 9 9 8 3 0 1 1 5 1 2 7 7 7 4 1 3 7 5 2 3 7 1 0 0 7 9 4 3 6 9 8 0 2 6 1 1 5 1 4 1 6 9 4 1 3 9 4 8 3 8 1 1 0 7 7 4 4 8 6 9 1 2 8 1 2 6 1 4 3 7 6 3 2 3 8 7 4 5 5 1 0 7 1 0 7 5 5 9 0 9 0 3 6 1 2 9 1 5 7 7 8 4 1 3 8 51 3 5 9 8 7 2 4 5 6 8 9 0 2 3 1 1 9 1 3 0 6 7 3 9 4 3 5 8 4 0 1 0 4 7 1 3 7 8 4 7 1 1 9 1 2 1 1 3 7 6 4 4 3 4 3 55 3 4 1 0 4 8 1 4 2 8 2 7 9 2 8 1 1 2 1 3 1 6 2 4 4 3 7 5 3 3 2 1 0 4 8 5 4 3 6 9 7 6 2 3 1 1 0 1 3 3 8 1 4 0 33 5 3 4 2 1 0 5 7 8 4 6 6 7 8 1 2 9 1 2 0 1 3 6 6 4 2 8 3 9 5 0 4 0 9 8 7 5 6 0 . QO .78 2 1 1 0 8 I 1 9 7 6 2 0 3 2 6 6 4 0 1 1 4 9 2 5 5 7 9 8 2 1 6 1 1 7 1 3 2 68 2 2 3 4 5 7 4 7 I 0 7 9 3 4 3 8 3 8 7 1 8 9 9 1 3 6 7 7 3 1 3 3 5 9 4 0 1 1 3 9 4 4 3 7 5 8 1 2 2 1 1 4 1 4 1 7 4 2 4 3 4 6 3 3 5 1 1 1 9 S 4 4 9 0 7 8 2 0 1 0 6 1 4 3 6 5 2 2 3 1 6 4 4 1 1 1 3 9 3 4 9 8 3 8 1 1 9 1 1 0 1 3 6 66 2 3 3 2 4 8 3 5 8 2 4 7 3 0 4 3 5 3 1 4 7 9 8 6 60 2 3 3 2 4 6 3 2 8 4 4 2 2 8 5 4 6 0 1 6 8 6 9 4 5 5 2 5 3 2 4 7 3 6 8 8 3 9 3 3 4 9 5 7 1 5 8 5 9 3 5 7 2 7 3 1 4 9 3 8 8 4 4 4 3 1 4 3 5 2 1 1 7 8 9 0 5 2 2 0 32 5 6 4 0 1 0 2 8 3 4 1 66 7 8 1 1 9 5 1 2 4 5 7 2 3 3 2 5 8 4 2 1 0 2 9 1 3 5 6 4 7 8 1 8 9 9 1 2 6 66 2 5 3 4 6 1 3 9 1 0 0 8 1 4 2 6 6 7 7 1 9 1 0 7 1 3 3 7 6 2 3 31 5 6 4 3 1 0 0 8 2 4 2 7 0 7 7 1 8 9 3 1 2 0 6 1 220 APPENDIX III - CONTINUED
OTU CHARACTERS 6 5 8 6 8 7 a s 8 9 9 0 9 1 9 2 . 9 3 9 4 9 5 9 6 9 7 9 8
UTAH 4 2 4 3 0 6 1 3 7 1 0 4 9 0 4 5 6 6 7 1 1 3 9 5 1 2 2 3 8 6 0 UTAH 5 2 0 3 3 5 4 3 1 9 8 9 8 3 8 5 8 6 7 1 4 8 1 1 1 2 3 6 6 1 UTAH 6 2 9 3 7 7 1 3 8 1 1 2 1 0 3 4 2 6 3 7 6 2 2 1 0 3 1 3 1 5 1 8 0 UTAH 7 2 9 2 6 5 3 3 8 1 0 1 8 8 4 2 6 5 6 6 1 8 8 3 1 2 2 4 3 7 2
insectivore HOST GROUP
CARN 1 1 5 4 3 4 9 3 3 1 0 4 7 4 6 9 9 7 8 4 2 4 1 0 8 1 1 8 4 4 7 2 NSPB 1 2 4 3 4 5 0 3 9 9 4 1 0 6 5 1 1 0 0 8 1 2 1 9 1 1 3 2 3 1 7 4 NSPB 2 2 5 3 0 4 6 3 7 1 0 0 1 0 2 5 3 8 2 7 7 2 3 9 4 1 2 7 4 2 7 5 NSPB 3 2 3 3 1 5 3 3 4 9 7 9 4 5 6 9 4 7 0 2 4 9 4 1 2 0 2 6 8 1 TALP 1 2 1 3 8 5 8 3 8 9 5 8 4 4 1 7 4 8 1 2 6 1 0 5 1 2 7 3 1 6 0 TALP 2 1 7 3 4 6 0 4 0 8 7 9 6 3 7 6 9 7 6 2 4 1 0 1 1 4 5 4 1 8 5 TALP 3 1 9 3 4 6 0 4 0 8 1 7 7 4 0 7 3 7 5 1 3 1 1 1 1 3 1 4 3 7 5 TALP 4 2 4 3 1 4 9 3 3 1 0 2 8 2 4 2 7 4 7 3 2 5 1 0 1 1 2 2 2 6 8 9 TALP 5 2 0 3 2 6 1 3 3 9 5 7 5 4 5 1 0 0 6 7 2 0 8 5 1 0 1 2 1 3 3 TALP 6 2 0 3 0 5 1 3 5 9 3 7 3 3 8 7 3 6 2 1 5 9 0 1 1 4 2 2 4 0 TALP 7 1 9 3 9 5 7 3 0 9 6 8 7 4 8 7 3 6 7 2 1 8 2 1 0 2 2 3 3 7 TALP 8 1 2 3 7 4 0 3 2 9 0 6 9 4 2 5 9 6 4 2 1 8 7 9 9 3 2 4 5 TALP 9 2 2 3 5 5 1 3 4 9 8 9 6 4 3 6 1 7 4 2 3 9 4 1 2 1 3 0 5 2 PALU 1 2 0 2 2 5 2 3 3 1 0 2 8 2 4 7 6 8 9 5 2 2 1 0 3 1 2 5 3 8 6 5 OBSO 1 2 7 4 7 6 3 5 0 1 0 3 7 5 4 7 6 8 7 6 2 4 1 1 1 1 2 5 3 8 7 7 OBSO 2 2 6 4 2 6 9 4 9 1 0 4 7 5 4 2 6 9 6 7 2 2 1 0 9 1 2 5 3 7 7 8
CARN IVORE HOST GROUP
STAF 1 2 4 4 9 4 2 3 1 7 6 5 8 3 9 6 6 7 2 2 2 8 5 1 0 2 3 3 9 5 STAF 2 2 4 4 6 44 3 4 7 9 6 0 4 2 6 5 7 2 2 3 8 6 1 0 6 3 4 9 3 STAF 3 2 5 54 5 0 3 5 9 5 6 7 3 7 5 6 6 9 2 4 8 7 1 0 6 26 9 9 STAF 4 2 3 5 1 3 9 3 1 8 7 5 8 4 0 6 1 7 1 2 2 8 5 1 0 0 2 7 9 7 221 APPENDIX III CONTINUED
OTU CHARACTERS 99 100 101 102 1 03 104 105 106 107 108 109 110 1 11 1 12 GEOMYID HOST GROUP
FEMU 1 54 148 .63 29 57 63 98 83 22 29 0 16 0 28 FEMU 2 50 141 70 30 56 54 95 75 27 20 0 17 0 26 FEMU 3 52 164 64 32 67 75 108 89 25 19 0 17 0 28 FEMU 4 52 149 69 38 63 70 89 91 30 21 0 16 0 29 GEOM 1 43 132 48 33 65 65 101 81 24 16 0 14 0 24 GEOM 2 38 122 45 32 58 61 86 70 21 17 0 13 0 20 LONG 1 50 126 54 38 62 55 98 82 20 23 0 18 0 21 LONG 2 57 1 15 54 37 67 58 100 81 22 17 0 17 0 21 LONG 3 49 104 50 41 70 57 122 82 22 18 0 12 0 19 THOM 1 58 136 52 39 72 63 101 92 22 31 0 18 0 29 THOM 2 48 121 51 40 71 59 109 97 29 20 0 12 0 26 HETEROMYIDHOST GROUP
HILL 1 39 100 34 34 63 42 87 78 18 17 0 16 0 20 HILL 2 42 113 40 32 53 49 88 86 15 15 0 15 0 17 HILL 3 39 106 39 31 52 46 90 82 18 17 0 17 0 19 h i l l 4 35 123 38 31 53 55 89 91 13 13 0 14 0 19 HILL 5 40 120 36 32 58 50 101 83 20 17 0 15 0 20 INCO 1 44 132 48 33 61 62 1 16 91 22 19 0 18 0 22 INCO 2 41 125 47 39 64 61 99 112 22 18 0 18 0 24 INCO 3 43 129 47 38 62 52 105 87 19 19 0 19 0 24 TRI A 1 41 136 40 36 60 48 97 109 27 27 0 21 0 32 TRI A 2 41 130 46 35 56 45 99 101 26 26 0 20 0 29 TRI A 3 37 131 47 37 66 51 89 1 08 27 26 0 28 0 31 SCIUR ID HOST GROUP
AFFI 1 48 161 66 39 67 58 1 1 1 109 29 22 0 20 0 32 AFFI 2 54 175 72 40 69 70 132 112 29 24 0 19 0 33 AFFI 3 50 169 69 43 71 60 131 1 19 27 23 0 20 0 33 AFFI 4 60 167 67 43 72 67 128 1 04 28 23 0 24 0 35 CYNO 1 60 166 69 43 79 62 122 1 15 25 33 0 18 0 35 INVA 1 72 166 73 36 73 74 121 107 21 28 0 20 0 30 INVA 2 62 162 74 40 77 61 124 1 10 20 25 0 20 0 28 ANGU 1 57 147 72 37 71 67 132 104 21 28 0 21 0 28 NSPA 1 33 116 46 35 67 52 1 15 80 14 13 0 14 0 19 NSPA 2 36 121 42 34 62 58 124 78 14 14 0 15 0 19 EUTA 1 57 162 70 41 67 64 137 99 28 28 0 18 0 33 EUTA 2 58 182 65 44 69 80 144 1 14 28 22 0 18 0 29 OCCI 1 59 159 66 43 73 58 133 1 07 27 24 0 21 0 32 PUNC 1 55 166 72 46 68 68 123 1 04 26 32 0 20 0 37 PUNC 2 48 178 69 40 62 69 140 1 1 1 27 32 0 19 0 36 PUNC 3 51 178 68 39 64 58 131 1 19 31 26 0 20 0 35 APPENDIX II! - CONTINUED
CHARACTERS 9 9 1 0 0 1 0 1 1 0 2 1 0 3 1 0 4 1 0 5 1 0 6 1 0 7 1 0 8 1 0 9 1 1 0 12
5 7 1 4 4 7 0 4 6 6 6 7 2 1 1 5 1 1 3 2 9 2 6 0 2 0 3 2 5 8 1 5 6 5 2 3 6 6 2 7 4 1 2 4 1 1 0 2 0 2 4 0 1 8 2 3 5 3 1 4 7 6 6 3 4 6 1 6 8 1 2 2 1 0 8 2 3 2 0 0 1 8 2 3 4 8 1 6 0 5 5 3 8 6 2 7 3 1 2 6 1 0 0 2 4 1 9 0 1 8 2 3
HOST GROUP
5 9 1 3 0 3 9 3 1 5 1 7 2 1 0 4 1 0 4 2 3 2 0 0 2 0 2 5 4 1 1 2 7 4 6 3 1 4 8 6 4 I 1 1 9 4 2 7 2 0 0 2 0 2 7 4 1 1 1 9 4 0 3 1 5 2 6 1 1 0 9 l O I 2 8 1 9 0 1 9 2 7 4 0 1 2 3 4 5 2 9 5 3 4 8 9 4 8 1 2 0 1 8 0 1 5 2 4 4 5 1 2 1 5 2 3 7 6 6 5 5 I 1 2 8 7 2 3 1 9 0 1 8 2 8 5 3 1 1 5 5 0 3 0 5 6 5 1 1 1 5 8 4 2 3 1 8 0 1 7 2 4 5 0 1 5 0 5 8 3 3 7 4 7 0 1 2 4 1 0 7 2 9 I S 0 1 9 2 3 5 5 1 5 5 6 1 4 1 7 8 6 3 1 3 0 1 1 3 3 0 2 3 0 2 0 3 3 4 6 1 3 2 5 6 3 6 7 0 5 8 1 2 2 1 1 5 2 8 2 1 0 1 7 2 3 4 4 1 4 6 6 2 3 7 6 3 6 2 1 3 1 1 1 2 3 2 2 3 0 1 3 2 5 5 4 1 4 3 5 2 3 5 5 6 6 8 1 1 4 1 1 5 2 9 2 2 0 1 8 3 0 7 2 1 7 9 7 2 4 6 7 8 8 9 1 6 4 1 5 0 3 3 2 4 0 2 1 3 1 6 3 1 8 4 8 0 4 4 8 1 8 1 1 7 2 1 2 3 3 3 2 7 0 2 2 3 2 6 1 1 7 3 6 4 3 9 8 1 7 6 1 3 9 1 3 5 3 7 2 2 0 1 9 3 5 5 1 1 5 7 6 7 4 0 6 4 6 8 1 2 9 1 2 3 3 2 2 6 0 2 0 3 1 6 0 1 7 8 7 1 4 8 7 6 7 4 1 3 7 1 2 3 3 1 2 7 0 2 1 3 2 6 7 1 6 6 7 2 4 1 7 9 8 1 1 3 0 1 1 2 3 0 2 4 0 2 0 3 0 4 9 1 5 8 5 9 3 8 6 3 5 6 1 4 0 1 1 5 2 8 2 3 0 2 0 3 3 5 3 1 7 1 7 1 4 5 7 2 7 1 1 4 7 1 3 3 3 1 2 6 0 2 1 3 3 6 3 1 7 3 7 6 3 8 8 3 7 5 1 5 7 1 2 9 3 3 2 2 0 2 0 3 2 5 5 1 7 0 6 9 3 3 5 9 6 1 1 3 7 1 2 0 3 1 2 4 0 2 1 3 3 4 6 1 5 0 6 0 3 9 7 0 6 5 1 3 9 1 1 1 3 0 2 3 0 1 9 3 1 5 0 1 5 8 6 8 4 0 6 6 6 6 1 3 5 1 4 3 3 1 3 3 0 1 9 3 0 5 3 1 6 1 6 1 3 8 6 4 7 1 1 2 6 1 2 5 3 1 2 4 0 1 9 2 9 5 1 1 5 8 6 6 4 2 6 0 7 1 1 3 0 1 3 6 3 0 2 4 0 1 9 3 2 4 9 1 5 8 6 6 3 7 7 4 6 4 1 2 3 1 2 2 2 6 1 9 0 1 5 2 6 5 9 1 6 6 6 6 4 5 7 9 7 9 1 3 2 1 2 7 3 0 2 3 0 1 7 3 0 5 . 2 1 5 6 5 9 3 8 7 4 7 8 1 2 2 1 2 5 3 0 2 6 0 2 1 3 5 5 7 1 6 1 5 7 4 4 7 7 6 5 1 3 9 1 2 1 2 2 2 3 0 2 0 2 9 5 6 1 6 1 6 0 4 4 7 8 7 1 1 5 0 1 1 7 3 2 2 6 0 2 0 3 4 5 9 1 6 5 6 1 3 9 7 1 6 8 1 4 1 1 0 6 2 8 2 4 0 2 0 3 2 4 0 1 1 1 3 7 2 8 4 3 5 4 9 7 9 0 2 3 1 8 0 1 9 2 7 3 8 1 1 3 4 1 3 2 5 1 5 2 1 0 3 8 5 2 2 1 8 0 2 0 2 6 3 7 1 1 1 ■ 3 9 3 2 4 9 S 3 9 4 9 1 2 4 1 8 0 1 9 2 7 3 9 n o 4 4 3 0 5 1 5 3 1 0 2 8 2 2 1 1 6 0 1 8 2 3 5 1 1 3 9 5 5 3 3 5 2 6 1 1 1 2 8 6 2 1 2 8 0 1 8 2 3 5 3 1 4 6 5 8 3 8 6 5 6 1 1 1 4 1 0 3 2 2 2 4 0 1 9 3 2 5 8 1 4 7 5 9 3 5 6 0 6 3 1 2 4 9 8 3 1 2 4 0 1 9 3 2 4 8 1 4 4 6 2 3 3 5 9 6 5 1 0 7 9 3 2 8 2 2 0 1 9 2 9 223 APPENDIX III - CONTINUED
OTU CHARACTERS 9 9 t o o 1 0 1 1 0 2 1 0 3 1 0 4 1 0 5 1 0 6 1 0 7 1 0 8 1 0 9 1 1 0 1 1 1 1 1 2
UTAH 4 5 1 6 0 1 4 2 3 3 6 7 5 4 1 2 5 1 0 4 3 0 2 3 0 2 0 0 3 4 UTAH 5 4 9 1 3 3 5 2 3 5 6 3 6 3 1 0 8 8 9 2 7 1 9 0 1 6 0 2 7 UTAH 6 5 1 5 6 1 5 6 4 0 7 0 6 3 9 3 1 2 6 3 0 2 6 0 1 9 0 3 5 U T A H 7 4 8 1 4 4 5 5 3 4 6 5 5 4 1 1 4 I 0 4 2 9 2 2 0 1 3 0 3 0
insectivore H O S T G R O U P
CARN 1 5 2 1 6 3 6 1 4 0 7 1 9 3 1 1 9 1 1 8 3 1 3 0 0 1 3 3 3 9 NSPB 1 5 5 1 4 3 5 2 3 1 7 8 7 1 1 2 3 1 0 5 2 7 2 5 0 1 7 0 3 3 NSPB 2 5 3 1 4 3 5 5 3 7 7 7 6 7 1 3 4 9 3 3 0 2 3 0 2 0 0 3 4 NSPB 3 5 6 1 4 5 5 1 3 7 7 4 7 5 1 1 1 1 0 3 3 0 2 6 0 1 6 0 3 1 1 5 1 TALP 1 4 2 4 9 4 0 7 9 6 9 1 6 1 8 4 3 3 2 7 0 1 0 2 3 7 TALP 2 5 3 1 5 2 6 0 3 1 8 1 6 9 1 4 5 1 1 5 2 9 2 8 0 9 2 3 3 TALP 3 5 4 1 5 3 5 5 2 9 6 8 7 4 1 2 5 1 0 4 2 9 2 8 0 1 1 2 3 5 6 0 T A L P 4 6 1 1 5 9 3 5 5 8 6 5 1 3 0 1 1 3 2 6 2 2 0 1 4 1 3 3 T A L P 5 4 9 1 4 0 5 1 3 5 6 2 6 1 1 2 0 8 9 2 4 2 0 0 9 2 2 4 6 TALP 4 5 1 4 1 5 0 3 2 5 9 6 1 1 2 3 8 1 2 4 2 0 0 9 2 2 6 7 TALP 5 4 1 4 4 4 3 3 6 6 5 6 4 1 3 0 8 8 2 4 2 0 0 9 2 2 6 8 TALP 4 3 1 2 6 4 6 3 4 6 1 6 6 1 2 2 8 1 2 7 2 2 0 1 0 2 2 7 TALP 9 4 6 1 7 3 5 4 4 2 7 4 6 8 1 2 5 9 5 3 2 2 4 0 1 6 1 3 1 5 5 PALU 1 1 4 7 5 1 3 8 6 9 7 2 1 2 2 1 0 0 2 2 2 9 0 1 1 2 3 1 OBSO 1 4 0 1 4 3 6 3 4 0 6 5 6 5 1 3 0 1 0 1 3 0 2 3 0 1 6 0 2 9 OBSO 2 5 0 1 5 1 5 8 3 7 6 2 6 7 1 4 0 1 0 7 3 0 2 2 0 1 8 0 2 9
CARNIVORE HOST GROUP
S T A F 1 3 0 1 4 9 6 3 3 2 5 8 6 1 1 0 1 1 1 4 2 3 2 2 3 1 8 0 3 0 S T A F 2 3 4 1 4 5 6 0 3 0 6 1 7 1 9 1 9 2 2 4 2 4 I 2 1 0 3 4 STAF 3 3 4 1 3 6 5 3 4 2 6 5 7 3 9 7 1 1 3 2 4 1 9 2 2 0 0 2 7 STAF 4 2 7 1 2 4 5 2 3 3 5 9 6 3 8 3 1 0 3 2 2 1 7 3 1 6 0 2 2 zzh APPENDIX III - CONTINUED
OTU CHARACTERS 1 1 3 1 1 4 1 1 5 1 1 6 1 1 7 1 1 8 1 1 9 1 2 0 1 2 1 1 2 2 1 2 3 1 2 4 1 2 5 1 2 6
GEOMYID HOST GROUP
FEMU 1 2 8 2 2 1 1 8 3 2 0 2 0 1 2 8 2 0 1 1 2 1 0 1 7 FEMU 2 3 3 1 1 3 1 1 2 1 7 1 9 1 2 1 1 7 1 1 3 9 1 4 FEMU 3 3 0 2 1 6 1 3 3 1 9 2 2 1 2 3 1 7 1 1 2 1 1 1 7 FEMU 4 3 2 2 1 5 1 4 2 1 9 2 0 2 1 8 1 5 1 1 0 a 1 7 GEOM 1 2 5 1 2 9 1 7 2 2 0 1 9 1 2 4 1 7 1 1 5 1 0 1 6 GEOM 2 2 3 1 2 0 1 2 1 1 8 1 8 1 2 0 1 2 1 1 2 9 1 3 LONG 1 2 2 3 1 6 1 6 2 2 1 2 2 3 2 4 2 2 2 9 9 1 6 LONG 2 1 8 3 1 2 1 2 2 1 9 2 0 3 2 2 1 9 1 7 8 1 3 LONG 3 1 8 3 9 a 2 1 7 1 9 3 1 9 1 5 2 7 5 1 3 THOM 1 2 4 2 1 9 1 6 2 2 4 2 3 1 2 5 1 9 1 1 6 1 0 1 6 T H O M 2 2 3 2 1 0 9 3 1 8 1 9 1 2 2 1 1 2 1 0 7 1 6
HETEROMYID HOST GROUP
HILL 1 1 9 1 2 4 1 5 I 1 6 1 8 1 1 3 1 1 2 7 5 1 4 HILL 2 1 9 2 1 5 1 2 2 1 6 1 2 2 1 0 8 1 7 6 1 1 H I L L 3 1 9 2 1 7 1 0 1 1 7 1 7 2 1 2 8 1 6 4 1 3 H I L L 4 1 4 2 1 5 1 2 1 1 8 1 2 2 1 0 1 1 2 5 6 1 3 h i l l 5 2 1 2 2 3 1 7 1 1 7 1 7 2 1 8 1 5 2 8 6 1 3 INCO 1 2 3 2 1 4 1 1 2 2 0 1 8 2 1 5 1 1 1 1 0 5 1 8 INCO 2 2 0 2 1 7 1 4 2 2 0 1 3 2 1 3 1 0 2 1 0 6 1 6 INCO 3 1 9 1 1 6 1 0 1 2 0 1 7 1 1 2 1 1 1 7 6 IS T R I A 1 1 9 1 1 7 1 0 1 2 2 2 3 1 1 5 1 0 1 1 1 6 2 1 T R I A 2 1 8 1 2 1 1 4 2 2 3 1 9 1 1 9 1 1 1 1 0 6 1 9 T R I A 3 2 1 1 1 9 1 3 2 2 4 2 1 1 1 8 1 1 1 1 1 6 2 1
S C I U R I D HOSTGROUP
AFFI 1 2 2 3 1 4 1 1 0 2 3 2 3 3 1 4 1 1 1 8 9 1 8 AFFI 2 2 5 3 1 6 1 6 0 2 4 2 4 3 1 3 1 5 1 1 0 1 0 2 0 AFFI 3 2 4 3 1 2 1 0 0 2 7 2 4 3 1 1 1 0 1 7 7 1 8 AFFI 4 1 9 3 1 2 1 0 0 2 7 2 3 3 1 1 1 0 1 6 7 2 0 CYNO 1 2 2 1 2 0 1 5 0 2 9 2 4 1 2 0 1 3 1 1 3 9 1 7 I N V A I 2 7 1 2 2 1 7 0 2 9 2 2 1 2 1 1 2 1 1 3 9 1 8 INVA 2 2 6 1 2 1 1 4 0 2 7 2 0 1 2 2 1 3 1 1 2 8 1 6 ANGU 1 2 6 1 2 0 1 3 0 2 5 2 3 1 2 2 1 3 1 1 4 6 1 8 N S P A 1 2 5 0 0 0 0 1 8 1 7 0 0 0 1 1 6 8 NSPA 2 2 3 0 0 0 0 2 0 1 8 0 0 0 1 1 6 1 0 EUTA 1 2 2 2 1 0 9 0 2 2 2 3 2 1 2 1 3 1 9 6 1 8 E U T A 2 2 3 2 a 6 0 2 3 2 3 2 8 8 1 9 6 1 7 OCCI 1 2 7 3 1 6 U 0 2 2 2 3 3 1 7 1 2 1 1 3 8 1 9 PUNC I 2 7 3 1 5 1 2 0 2 6 2 3 ' 3 1 4 1 3 1 1 0 8 2 0 PUNC 2 3 0 3 1 4 1 0 0 2 8 2 4 3 1 3 1 0 1 1 0 8 2 0 P U N C 3 3 0 3 1 6 1 3 0 2 7 2 5 3 1 4 1 2 1 1 1 9 2 2 225 APPENDIX 111 - CONTINUED
OTU CHARACTERS 113 114 115 116 117 118 1 1 9 120 121 122 123 124 125 126
P U N C 4 2 3 3 1 4 1 0 0 2 8 2 3 1 2 1 0 1 9 6 1 9 T O R U 1 2 8 3 8 1 3 0 2 1 22 1 6 1 0 1 1 4 8 1 6 T O R U 2 2 6 3 7 1 7 0 2 1 2 0 1 6 1 2 1 1 5 8 1 6 T O R U 3 2 7 3 4 9 0 2 2 21 1 3 1 0 1 1 6 9 1 6
MUROID HOST GROUP
B I S E 1 1 3 1 9 8 0 2 2 20 1 1 0 8 1 6 4 1 8 B I S E 2 2 0 1 10 1 0 0 2 1 21 1 8 7 1 5 5 1 9 B I S E 3 1 7 1 9 7 0 2 2 21 1 9 8 1 6 5 1 8 B R E V 1 1 7 0 2 7 0 1 9 1 7 1 6 4 I 8 5 1 6 B R E V 2 2 4 0 4 7 0 2 1 1 9 1 6 7 1 9 6 1 6 B R E V 3 2 7 0 3 6 0 1 9 1 8 1 7 5 1 1 0 6 1 3 I S A B 1 2 1 1 1 3 1 0 0 2 3 2 6 I 1 6 1 0 1 1 0 7 1 6 I S A B 2 2 7 1 1 2 1 2 0 2 5 3 0 1 1 5 1 2 1 1 0 8 1 8 I S A B 3 2 8 1 1 4 1 0 0 1 9 2 4 1 1 4 1 0 1 9 7 1 7 I S A B 4 1 9 1 1 4 1 1 0 2 1 2 7 1 1 6 9 1 1 1 7 1 9 1 S A B 5 2 2 1 1 4 1 0 0 2 4 2 4 1 1 5 1 1 1 9 7 1 7 I S A B 6 2 2 1 1 7 1 1 0 2 7 2 9 1 1 5 1 1 1 9 7 2 1 I S A B 7 2 8 1 6 1 2 0 2 9 2 8 2 1 7 1 2 1 1 3 8 2 1 I S A B 8 2 9 1 1 7 1 2 0 2 7 3 4 I 2 0 1 4 1 1 1 8 2 0 I S A B 9 2 3 1 2 5 1 5 0 2 6 2 4 1 2 2 1 5 1 1 2 6 2 0 I S A B 1 0 2 1 I 2 1 1 5 0 2 6 2 6 I 2 0 1 3 1 1 4 8 1 9 I S A B 1 1 2 4 1 1 8 1 2 0 2 4 2 7 I 1 9 1 2 I 1 2 7 1 8 I S A B 1 2 2 3 1 2 5 1 6 0 2 4 2 6 1 2 0 1 5 1 1 1 8 1 9 I S A B I 3 2 3 I 2 0 1 5 0 2 7 2 8 I 1 9 1 3 1 I 1 7 1 9 I S A 3 1 4 1 9 1 1 5 1 1 0 2 6 2 6 1 1 8 1 2 1 1 1 9 2 0 I S A B 1 5 2 0 1 2 2 1 1 0 2 6 2 7 1 1 9 9 1 8 6 2 2 I S A B 1 6 2 1 1 2 0 1 2 0 2 3 2 4 1 1 8 1 1 I 9 6 1 9 I S A B 1 7 2 8 1 2 3 1 2 0 2 5 2 6 1 2 0 1 3 1 1 2 7 2 0 ISAB 18 2 6 1 2 2 1 2 0 2 3 2Q 1 2 0 1 1 I 1 3 7 2 0 I S A 3 1 9 2 6 1 2 0 1 3 0 2 4 2 6 1 2 0 1 3 1 1 2 7 2 0 I S A B 2 0 2 0 1 1 7 - 2 0 1 9 2 4 I 1 5 1 0 1 10 7 1 7 L A T I 1 2 1 1 1 3 1 0 0 2 6 2 4 1 1 9 1 2 1 1 2 9 1 9 L A T I 2 2 4 1 1 5 1 2 0 2 7 2 5 1 1 8 1 4 1 1 3 8 1 8 L A T I 3 2 3 1 1 6 1 0 0 2 3 22 1 1 7 1 3 1 1 3 9 1 8 L A T I 4 2 6 1 1 7 1 1 0 2 6 2 7 1 1 9 1 4 1 1 3 8 2 0 L A T I 5 1 7 1 1 9 1 2 0 1 5 2 3 1 2 2 1 3 1 1 3 9 2 0 N E O T 1 2 0 1 1 0 8 0 2 0 IQ I 1 0 7 1 8 6 1 6 N E O T 2 2 2 1 1 0 1 1 0 2 1 IQ 1 1 0 9 1 1 0 7 1 6 N E O T 3 1 9 1 9 7 0 2 2 1 9 1 9 7 1 9 6 1 8 N E O T 4 2 1 1 5 1 0 0 2 0 IQ 1 1 1 8 1 8 7 1 5 T A R S 1 2 3 1 8 0 0 2 3 20 1 1 1 8 1 8 7 1 6 U T A H 1 2 8 1 0 7 0 2 8 21 1 1 1 8 1 8 6 1 8 U T A H 2 2 5 1 9 7 0 2 3 2 3 1 1 4 8 1 1 0 6 1 8 U T A H 3 2 6 1 9 8 0 2 1 22 1 1 2 9 I 1 2 8 1 5 226 APPENDIX III - CONTINUED
OTU CHARACTERS 1 13 114 1 15 116 1 17 118 19 120 121 122 123 124 125 126
UTAH 4 24 1 1 0 8 0 24 23 1 13 10 1 9 6 19 UTAH 5 24 2 1 1 8 0 20 21 1 10 8 1 8 6 IS UTAH 6 19 1 10 9 0 24 24 1 11 10 1 1 1 7 18 UTAH T 23 1 10 9 0 23 21 1 13 10 1 1 1 5 16 INSECTIVORE HOST GROUP
CARN 1 24 1 15 8 0 25 22 1 19 11 1 20 10 19 NSPB 1 IS 1 28 15 0 28 28 1 23 13 I 13 a 17 NSPB 2 20 1 27 14 0 27 23 1 24 13 1 16 9 18 NSPB 3 22 1 22 12 0 27 24 1 21 12 1 13 8 17 TALP 1 23 1 21 12 0 22 27 1 21 12 1 15 9 22 TALP 2 21 1 20 14 0 20 23 1 17 11 1 15 10 19 TALP 3 17 1 22 14 0 27 28 1 23 14 1 12 9 19 TALP 4 23 1 22 15 0 24 24 1 21 15 2 9 8 18 TALP 5 20 1 23 1 1 0 18 22 1 19 1 1 1 1 1 7 16 TALP 6 18 1 23 I 1 0 17 22 1 21 13 1 1 1 7 15 TALP 7 21 2 19 13 0 19 23 1 21 11 1 1 1 8 17 TALP B 19 1 19 10 0 19 21 1 18 11 1 1 1 6 14 TALP 9 24 1 23 17 0 23 26 1 23 14 1 13 9 22 PALU 1 21 1 21 12 0 22 24 1 22 13 1 1 1 8 18 OBSO 1 15 3 9 U 0 23 25 3 10 12 1 8 8 16 OBSO 2 23 3 8 1 1 0 26 23 3 12 18 2 9 6 17
CARNIVORE HOST GROUP
STAF 1 20 4 15 15 2 32 28 4 19 15 2 8 7 17 STAF 2 16 4 19 16 3 26 29 4 21 13 2 8 7 20 STAF 3 21 4 15 12 3 31 27 4 15 15 2 5 6 17 STAF 4 14 4 17 16 3 28 27 4 17 15 2 7 6 15 227 APPENDIX III - CONTINUED
OTU CHARACTERS 127 120 129 130 131 132 133 134 135 136 137 138 139 140 GEOMYID HOST GROUP
FEMU 1 0 1 1 1 0 0 1 1 67 63 54 80 91 43 FEMU 2 0 1 1 1 0 0 1 1 60 57 58 72 90 45 FEMU 3 0 1 1 1 0 0 1 1 60 59 61 83 86 45 FEMU 4 0 1 1 1 0 0 1 I 65 63 66 94 93 51 GEOM 1 0 0 1 0 0 0 I 1 57 51 44 70 72 33 GEOM 2 0 0 1 0 0 0 1 1 62 49 49 74 83 37 LONG 1 0 0 1 0 0 0 1 1 61 55 50 73 81 40 LONG 2 0 0 1 0 0 0 1 1 63 55 47 72 80 37 LONG 3 0 0 1 0 0 0 1 1 50 53 54 70 68 37 THOM 1 0 0 1 0 0 0 1 1 77 65 68 81 95 44 THOM 2 0 0 1 0 0 0 1 1 54 55 61 86 73 38 HETEROMYID HOST GROUP
HILL 1 0 0 0 0 0 0 1 1 51 52 47 69 69 33 HILL 2 0 0 0 0 0 0 1 1 52 55 47 69 72 33 HILL 3 0 0 0 0 0 0 1 1 50 46 46 71 65 31 HILL 4 0 0 0 0 0 0 1 1 61 50 48 72 72 33 HILL 5 0 0 0 0 0 0 1 1 60 53 48 78 75 37 INCO 1 0 0 0 0 0 0 1 1 57 61 55 83 73 38 INCO 2 0 0 0 0 0 0 1 I 50 59 49 73 67 26 INCO 3 0 0 0 0 0 0 1 1 63 58 54 82 82 35 TRI A 1 0 0 0 0 0 0 1 1 58 60 53 86 82 36 TRIA 2 0 0 0 0 0 0 I 1 64 63 52 81 78 39 TRI A 3 0 0 0 0 0 0 1 1 61 59 53 89 83 37
SCIURID HOST GROUP
AFFI 1 2 0 1 0 1 0 1 1 63 67 62 90 93 43 AFFI 2 0 0 1 0 1 0 1 1 85 72 66 98 1 09 53 AFFI 3 0 0 1 0 1 0 1 1 66 71 60 94 98 48 AFFI 4 0 0 1 0 1 0 1 1 75 72 64 98 1 03 45 CYNO I * 1 0 1 0 2 0 1 1 74 75 63 93 98 46 INVA 1 1 0 • I 0 2 0 1 1 71 73 67 97 1 00 47 INVA 2 2 0 1 0 ■ 2 o 1 1 83 68 69 100 101 45 ANGU 1 2 0 1 0 2 0 1 1 63 70 67 94 89 47 NSPA 1 3 0 0 0 0 0 1 1 61 47 41 63 87 37 NSPA 2 3 0 0 0 0 0 1 1 61 59 42 62 83 39 EUTA 1 2 0 1 0 1 o 1 1 62 72 63 89 93 44 EUTA 2 2 0 1 0 1 o 1 1 61 67 68 100 80 40 OCCI 1 2 0 1 0 2 0 1 1 61 72 59 102 86 48 PUNC 1 1 0 1 0 2 0 1 1 87 76 66 105 1 17 53 PUNC 2 1 0 1 0 2 0 1 1 80 76 73 108 108 49 PUNC 3 1 0 1 0 2 0 I 1 89 84 69 108 1 15 56 228 APPENDIX III - CONTINUED
OTU CHARACTERS 127 128 129 130 131 132 3 134 135 1 36 137 138 139 140
PUNC 4 1 0 I 0 1 0 1 1 56 64 61 96 80 40 TORU 1 1 0 0 0 1 0 1 1 56 67 60 82 82 42 TORU 2 1 0 0 0 I 0 1 1 52 62 58 77 81 38 TORU 3 1 0 0 0 1 0 1 1 64 69 52 85 94 34
MUROID HOST GROUP
BISE I 0 0 0 0 0 0 1 0 63 61 51 68 82 33 BISE 2 0 0 0 0 0 0 1 0 63 56 41 72 73 33 BISE 3 0 0 0 0 0 0 1 0 59 55 45 69 78 33 BREV 1 1 0 0 0 0 0 1 I 51 50 55 75 79 28 BREV 2 1 0 0 0 0 0 1 1 48 52 60 79 73 32 BREV 3 1 0 0 0 0 0 1 1 53 51 60 79 68 33 ISAB 1 0 0 0 0 0 0 1 1 60 69 61 94 83 46 ISAB 2 0 0 0 0 0 0 1 1 72 70 S3 96 101 47 ISAB 3 0 0 0 0 0 0 1 1 62 63 53 82 84 42 ISAB 4 0 0 0 0 0 0 1 1 57 62 60 100 78 41 ISAB S 0 0 0 0 0 0 1 1 74 64 46 82 92 41 ISAB 6 0 0 0 0 0 0 1 t 76 71 71 108 94 40 ISAB 7 0 0 0 0 0 o 1 1 83 78 61 104 111 50 I SAB 8 0 0 0 0 0 0 1 1 67 80 71 101 97 51 ISAB 9 0 0 0 0 0 0 1 1 69 70 61 92 93 45 I SA B 10 0 0 0 0 0 0 I 1 81 72 66 1 12 100 50 ISAB!1 0 0 0 0 0 0 1 1 72 75 66 103 100 49 I SAB 12 0 0 0 0 0 0 1 1 66 67 60 89 88 46 I SA B 13 0 0 0 0 0 0 1 1 73 70 67 104 95 52 I SA B 14 0 0 0 0 0 0 1 1 82 79 59 92 112 51 I SA B 15 0 0 0 0 0 0 I 1 70 64 60 89 80 33 I SA B 16 0 0 0 0 0 0 1 1 69 64 58 91 92 44 ISAB17 0 0 0 0 0 0 1 1 64 65 58 87 88 44 ISAB 18 0 0 0 0 0 0 1 1 59 67 61 89 84 45 ISAB19 0 0 0 0 0 0 1 1 76 78 61 92 92 46 ISAB20 0 0 0 0 0 0 1 1 61 69 59 91 87 39 LATI 1 1 0 1 0 0 0 1 1 61 69 53 71 S3 41 LATI 2 1 0 1 0 0 0 1 1 66 73 64 91 89 38 LATI 3 1 0 1 0 0 0 1 1 82 74 59 87 79 35 LATI 4 1 0 1 0 0 0 1 1 63 74 64 98 92 49 LATI 5 1 0 1 0 0 0 1 1 78 67 59 91 99 44 NEOT 1 1 0 0 0 0 0 1 0 53 SO 44 69 76 29 NEOT 2 I 0 0 0 0 0 1 0 65 49 42 64 80 33 NEOT 3 2 0 0 0 0 0 1 0 48 53 45 71 70 31 NEOT 4 1 0 0 0 0 0 1 0 56 51 47 67 78 33 TARS 1 2 0 1 0 t 0 1 1 66 66 53 75 86 41 UTAH 1 2 0 1 0 1 0 1 1 72 61 59 83 101 41 UTAH 2 2 0 1 0 1 0 1 1 61 61 61 82 89 40 UTAH 3 2 0 1 0 1 0 1 1 60 62 61 82 83 41 229 APPENDIX III - CONTINUED
OTU CHARACTERS 127 128 129 130 131 132 133 134 135 136 137 138 139 140
UTAH 4 2 0 1 0 1 0 1 1 61 61 55 78 81 37 UTAH 5 2 0 1 0 0 0 1 t 57 56 53 76 71 38 UTAH 6 1 0 1 0 1 0 1 1 78 65 58 82 102 39 UTAH 7 1 • 0 I 0 1 0 I 1 59 59 55 79 83 38
INSECTIVORE HOST GROUP •
CARN 1 1 0 1 0 2 1 1 1 78 67 56 80 102 48 NSPB 1 2 0 1 0 0 o 1 1 67 63 50 85 89 43 NSPB 2 2 0 1 0 0 0 1 77 63 48 87 99 53 NSPB 3 1 0 1 0 0 0 1 1 79 65 50 78 1 03 45 TALP I 2 0 1 0 0 0 1 1 70 67 58 89 87 46 TALP 2 2 0 I 0 0 0 1 1 66 64 55 83 92 43 TALP 3 2 0 1 0 0 0 1 1 81 66 50 94 100 49 TALP 4 2 0 1 0 0 0 1 1 67 60 50 79 88 43 TALP 5 2 0 1 0 0 0 1 1 70 57 47 79 83 46 TALP 6 2 0 1 0 0 0 1 1 73 60 45 74 93 45 TALP 7 2 0 1 0 0 0 1 1 67 61 49 79 85 47 TALP 8 3 0 1 0 0 0 1 1 63 55 45 79 83 31 TALP 9 2 0 1 0 0 0 1 1 81 62 50 06 105 41 PALU 1 3 0 1 0 0 0 1 1 67 66 55 89 81 41 OBSO 1 0 0 0 0 0 0 1 1 75 62 49 79 87 41 OBSO 2 0 0 0 0 0 0 1 1 79 62 52 91 94 43 CARN IVORE HOST GROUP
STAF 1 4 0 t 0 0 0 0 1 61 60 55 73 85 45 STAF 2 4 0 1 0 0 0 0 1 75 61 53 79 96 49 STAF 3 3 0 1 0 0 0 0 1 61 58 57 70 83 46 STAF 4 4 0 1 0 0 0 0 1 61 49 55 71 77 31 230 APPENDIX III - CONTINUEO
OTU CHARACTERS 141 142 143 144 145 146 147 148 149 150 151 152
GEOMYID HOST GROUP
FEMU 1 49 80 74 43 43 38 83 69 37 53 52 102 FEMU 2 50 69 61 42 39 37 76 61 33 51 47 94 FEMU 3 53 80 70 50 51 48 88 72 41 68 61 106 FEMU 4 53 83 73 41 51 50 89 68 35 70 63 110 GEOM 1 39 71 62 23 33 33 62 51 23 45 44 80 GEOM 2 40 70 61 33 33 31 67 60 24 45 44 85 LONG 1 41 67 69 41 37 37 67 62 31 51 47 86 LONG 2 35 67 69 36 36 29 66 61 28 47 43 87 LONG 3 44 46 58 31 40 39 62 58 30 54 50 86 THOM 1 47 71 82 41 47 41 81 75 32 53 57 105 THOM 2 50 66 65 36 46 44 75 60 28 62 58 97
HETEROMYID HOST GROUP
HILL 1 31 64 58 27 31 30 62 46 23 42 42 78 HILL 2 41 57 61 32 34 33 64 55 23 47 46 87 HILL 3 38 61 52 24 37 34 67 47 31 48 47 80 h i l l 4 40 70 60 30 37 33 70 51 24 51 50 91 h i l l 5 37 72 64 31 35 33 66 53 24 43 42 80 INCO 1 42 61 60 33 39 37 70 51 27 47 47 89 INCO 2 41 57 55 22 37 34 67 50 22 44 50 78 INCO 3 40 75 61 29 39 37 70 61 27 51 49 93 TRI A 1 44 80 64 25 43 39 85 54 25 53 53 108 TRI A 2 38 78 70 33 39 31 75 62 26 48 44 100 TRIA 3 44 63 60 31 42 38 83 55 28 53 51 105 SCIURID HOST GROUP
AFFI 1 45 80 78 40 42 39 78 77 35 53 51 104 AFFI 2 51 93 101 50 43 41 89 90 37 59 53 113 AFFI 3 48 82 79 44 42 41 80 73 43 54 53 1 1 1 AFFI 4 49 88 76 43 42 42 82 77 39 58 56 111 CYNO 1 53 89 83 41 46 42 81 74 36 59 56 105 INVA 1 50 91 83 47 47 48 84 89 41 61 61 111 INVA 2 55 94 85 42 49 44 95 81 35 63 62 115 ANGU I 52 75 76 44 43 45 82 71 39 58 58 103 NSPA 1 27 60 72 31 27 28 54 66 29 31 23 64 NSPA 2 27 56 70 34 27 27 50 64 30 32 26 70 EUTA 1 48 76 76 40 42 41 78 73 37 58 52 107 EUTA 2 50 72 74 37 41 41 87 70 33 61 56 112 OCCI 1 SO 82 74 43 45 42 80 70 39 61 58 109 PUNC 1 55' 94 82 52 51 47 94 82 44 64 61 123 PUNC 2 50 89 95 so 48 44 95 89 43 67 61 124 PUNC 3 54 98 82 50 49 47 94 89 45 65 61 122 231 APPENDIX III - CONTINUED
OTU CHARACTERS 141 142 143 144 145 146 147 148 149 150 151 152
PUNC 4 50 94 75 33 42 39 77 64 32 59 53 99 TORU 1 45 6 6 74 40 37 37 63 67 36 50 50 81 TORU 2 43 61 69 39 37 37 62 67 34 50 49 81 TORU 3 39 82 72 39 34 30 71 61 35 50 49 95
MUROID HOST GROUP
BISE 1 33 70 6 6 35 26 26 65 6 6 33 40 39 79 BISE 2 28 65 65 28 23 30 S3 51 24 44 39 83 BISE 3 30 64 65 31 33 24 62 62 23 39 38 81 BREV 1 34 60 69 26 27 25 58 6 8 27 39 39 75 BREV 2 38 58 6 6 29 24 30 59 60 27 45 41 78 BREV 3 41 61 55 32 32 31 50 56 30 48 42 75 ISAB 1 48 74 78 39 42 39 76 74 35 59 55 10 0 ISAB 2 45 8 6 91 42 39 38 05 83 39 58 53 1 12 ISAB 3 41 70 70 39 38 37 70 6 6 34 54 49 90 ISAB 4 49 67 64 37 41 41 80 6 6 31 61 57 107 ISAB 5 37 61 71 39 29 31 67 70 35 45 37 89 ISAB 6 61 101 8 6 38 50 50 75 83 33 69 6 6 114 ISAB 7 50 92 82 48 39 30 85 90 41 61 50 1 16 ISAB 8 59 84 85 50 50 50 89 87 45 76 70 130 ISAB 9 49 84 80 42 41 42 81 70 33 59 57 1 0 2 ISAB 10 50 90 87 44 46 45 81 6 6 38 69 61 I 14 I SA B 11 50 73 83 48 44 43 78 79 39 58 58 94 I SAB 12 49 81 76 42 42 43 73 69 35 58 59 104 I SA B 13 53 91 85 51 48 48 8 6 78 44 6 8 64 119 I SA B 14 46 87 81 55 39 33 03 89 42 59 53 109 ISABI 5 45 87 75 33 41 41 75 67 36 59 60 107 I SAB 16 48 79 76 42 42 43 73 74 36 59 50 100 I SAB 17 51 73 82 43 45 43 75 79 35 62 56 105 ISAB 13 51 75 74 42 41 45 71 71 36 63 61 110 I SAB 19 47 81 81 45 45 44 79 79 37 63 61 106 ISAB20 47 67 75 37 40 40 76 73 33 58 56 100 LATI 1 52 74 76 41 49 47 0 0 80 29 65 60 108 LATI 2 50 90 73 39 47 47 91 70 36 6 6 64 1 16 LATI 3 50 84 04 37 39 39 83 70 35 53 53 105 LATI 4 53 8 8 83 46 40 46 93 78 41 6 6 60 120 LATI 5 47 85 78 33 44 39 85 70 30 59 53 104 NEOT 1 33 63 63 26 31 29 56 57 25 41 39 75 NEOT 2 30 58 69 29 28 26 58 67 27 37 35 72 NEOT 3 37 61 60 27 30 31 51 54 30 42 41 76 NEOT 4 36 57 61 27 29 31 55 6 0 24 41 40 77 TARS 1 39 6 8 73 32 29 27 67 63 29 39 39 85 UTAH 1 46 80 71 43 39 37 72 73 38 51 49 95 UTAH 2 47 63 6 8 39 39 37 72 62 32 53 50 92 UTAH 3 47 67 71 37 40 38 71 62 32 53 49 93 232 APPENDIX III - CONTINUED
OTU CHARACTERS 141 142 143 144 145 146 147 148 149 ISO 151 152
UTAH A 47 83 69 33 39 38 72 61 26 50 45 92 UTAH 5 36 61 59 30 32 33 62 56 29 49 45 83 UTAH 6 43 81 92 45 39 39 77 87 41 55 52 104 UTAH 7 43 64 71 38 36 34 53 63 38 50 47 75
INSECTIVORE HOST GROUP
CARN 1 47 60 83 50 33 39 70 78 42 52 49 8 6 NSPB 1 44 67 77 43 38 39 67 75 37 49 50 82 NSPB 2 41 78 96 46 37 31 77 87 45 48 42 91 NSPB 3 44 78 87 47 27 26 71 83 37 40 38 87 TALP 1 49 72 74 41 41 42 6 8 78 38 55 53 94 TALP 2 49 69 78 46 44 42 75 72 42 60 53 91 TALP 3 43 84 94 46 43 42 78 69 41 53 55 97 TALP 4 41 72 81 43 39 39 72 6 8 39 49 49 89 TALP 5 38 6 8 74 40 37 34 64 63 37 47 44 80 TALP 6 36 69 82 45 33 31 71 62 39 45 39 84 TALP 7 40 69 70 41 36 33 64 67 35 47 41 80 TALPB 37 67 76 33 35 34 6 6 59 28 48 43 79 TALP 9 39 78 95 40 38 35 79 89 39 47 47 94 PALU 1 45 69 72 36 41 41 72 62 33 53 52 83 OBSO 1 47 78 74 39 34 29 69 6 8 33 45 46 99 OBSO 2 53 82 84 41 33 33 82 77 33 50 45 1 0 2
CARNIVORE HOST GROUP
STAF 1 42 64 62 41 41 39 67 55 34 58 53 8 6 STAF 2 43 6 8 73 44 43 40 67 60 35 55 51 83 STAF 3 45 64 69 42 43 41 72 67 35 56 56 93 STAF 4 47 65 6 8 29 34 40 6 6 67 26 50 51 82 Flyura 1, Fhenopram obtained from the Condensed Ratio Correlation (CRC) matrix.
233 i x a o 2> * tccn Z n cv ^nc n > O r» z co 5 on>" *n ifl*<--* * - oz o ■ oe
M £ Figure 2. Fhonogram obtained from the Condensed Basic Correlation (CBC) matrix.
235 '0.10 8.SO 8.10 0.20 0.0* 0 00 0.10 0.00 0.70 0 00 0.00 1.00 p m ~r | t ~tt r ]~t~r i i | i i i i |~i t i T[ i i n | t'n i | i ) n i | i i u | u u | 236
THOM
OIOU
N 0 PA AFFI PUNCOCCI AF FI CUT* AFFI
AFFI CVNO IHVA ANOU
I. AT I
TAPS
UTAH
OBIO
TONII
BACV
CARN
NSPB
TALP
PALO
I I I M I ) I I I I I 1.1-1-L I 1 I ]. I I I I I 1 I I I i I I i 1 1 1 1 l i l I i i i i.l . r ■ ■ I ■ . i i I 2 *0.10 0.00 0.10 0.30 0 30 0.40 0.10 0.00 D.T0 0.10 0.00 1.0 r F igu re 3* Pbenogran obtained from the Condensed Ratio D ista n ce (CRD) m a tr ix .
237 1.441 8.43 7 0 3«S O.JIT 8.333 0.317 8.313 0.3*7 0.103 0.1*3 0.113 O.OTT j I M I | J I I I |~1 I 11 | ‘ I I i I | 1 1 I I | I J I I j I M I | TT" 1 I | I I J I | I'1 I I | I I * I | 238
THOM
OEOM
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AFFI
FUNC AFFI IUTA OCCI PUNC AFFI c y n o IHVA AMOU CUT*
LATI
T A ll
UTAH
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OOAO
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BREV
BIO I
HEOT
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FALU
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D
11 11111111111111111 ■ 111111111111111111 11111111 111. 11 1111 3 c a l l t . l l l 1.111 0.107 0.111 0.101 0.111 0.111 0.101 0.1*1 0.111 O.OIT d Figure 4. Graph diagram of the STAF cluster; relationships based on a revised Condensed Ratio Correlation (revised-CRC) matrix. Solid straight lines = first-order vectors; dashed straight lines = second-order vectors; dotted straight linos - third-order vectors; and solid wavy lines = group connecting vectors.
Figure 5« Graph diagram of major groups "G” and "H"; relation ships based on a revised Condensed Ratio Correlation (revised-CRC) matrix. Solid straight lines = first- order vectors; dashed straight linos = second-order vectors; dotted straight lines = third-order vectors; and solid wavy lines = group connecting vectors.
239 2^0 STAF
rS T A f
PUNC?
'GEOU'
FEMU FEM
GEO. NEOT THOI THOI
LOHCT TR1A TniA INCCT LONQ>
TRIA LONGr
INCO
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HILL
HlLt? IS AQ *0 J
h ispm Figure 6. Graph diagram of major groups "I” and "S-M"; relation ships based on a revised Condensed Ratio Correlation (revised-CRC) matrix. Solid straight lines = first- order vectors; dashed straight lines = second-order vectors; dotted straight lines = third-order vectors; and solid wavy lines = group connecting vectors.
2bl 2^4-2
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/on£v\ v l > = '/drei I /.* ViW *' ^ © -
/ d r e v \ \ W "ST« - ftSAB\ V 4 ^ V-/ I A B \ —-* V 12 U / ^ * \ /iSAO^ 15 J N^IX^/tSAB\ 2 -^/%. J t A B / * © * ' T j r < F v \ © — * © j © N'V " -s > ' * AI SABA i > /toru \ /OBsSj
(AuaiAr ^ r e u J j T /CYNOj V- x _ y i X . f: y ® z 2" ^ **• A a l p V ; ; r > f ^ PALU n . ©
1 T C\L •i/jALP ©=©“© -^„.-'7 S © ©
6 Figure 7 Sami-diagramatic representation of the dorsal chaetotaxy in tho females of Hirstionyssus.
2^3 2b4
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j4 o
s4 ® r4 « zS d h a s 5 o
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R 6 ® Figuro 8. Kirstlonyssus isabellinus (Oudemans). Paraxial view of chelicera in the female. Scale = 50 microns.
Figure 9, Hirstionyssus isabellinus (Oudemans). Ventral view of gnathosoma in the female. Scale = 100 microns.
Figure 10. Klrstionyssns isabellinus (Oudemans). Ventral view of tarsus II in the female. Scale = 50 microns.
Figure 11. Hirstionyssus isabellinus (Oudemans), Ventral view of tarsus IV in the female. Scale = same as Fig, 10. ant av, av, Figure 12, Hirstion.yssns femurails Allrad. Ventral view of the female. Scale = 100 microns.
Figure 13. Klrstjonyssus femurails Allred, Dorsal view of the female. Scale = same as Fig. 12.
Figure 1^. Hirstionyssus femurails Allred, Vontral view of tarsus II in tho female. Scale = 50 microns.
Figure 15. Hirstionyscus femuralls Allrod. Ventral view of genu II in the female. Scale = same as Fig. 14.
2 47
Figure 16. Hirstionyssus geomydis (Keegan). Ventral view of the female. Scale = 100 microns.
Figure 17. rlirstionyssus goomydis (Keegan). Dorsal view o f th e fem ale. S cale = same as Fig, 16.
Figure IS. Ejrstionyssus ftp omydis (Keegan). Ventral view of tarsus II in the female. Scale = $0 m icrons.
249
Figure 19. Hirstionyssus thonomys Allred and Back. Ventral view of the female. Scale = 100 microns.
Figure 20. Hirstionyssus thonomys Allrod and Beck. D orsal view of the female. Scale = same as Fig 19.
Figure 21. Hirstionyssus thonomys Allrod and Bock. V e n tra l view of tarsus II in the fenalo. Scale = : 50 m icrons.
251
Figure 22, Hirstionyssus longichelae Allred and Beck. Ventral view of the female. Scale = 100 microns.
Figure 23, Hirstionyssus lonreichclae Allred and Eock, Dorsal view of the female. Scale = same as Fig, 22,
Figure 2^. Hirstionyssus longichelae Allred and Beck, Ventral view of tarsus IX in the female. Scale = 50 microns.
253
Figure 25. Hirsttonyssus hilli (Jameson), Ventral view of the female. Scale = 100 microns.
Figure 26. Uirstionyssns hilli (Janeson). Dorsal view of th e fem ale. Scale = same as Fig, 25*
255 256 Figure 27. Klrstionyssus "C" n. sp. Ventral view of the female. Scale = 100 microns.
Figure 28. Hirstionyssus "C" n. sp. Dorsal view of the female. Scale =; same as Fig. 27*
257 258 Figure 29. Hirstionyssus incomptis ("ads and Hightower), V en tral view of the female. Scale = 100 microns.
Figure 30. Hirstionyssus incomptis (Fads and Hightower), D orsal view of the female. Scale = same as Fig. 29.
259 260 Figure 31- Hirstionyssus triacanthus (Jameson). Ventral view of the female. Scale = 100 microns.
Figure 32, Hirstionyssus triacanthus (Jameson). Dorsal view o f th e fem ale. Scalo = same as F ig, 31*
261
Figure 33* Hirstionyssus "B" n. sp. Ventral view of the female. Scale = 100 microns.
Figure Hirstionyssus "B" n, sp. Dorsal viow of the fem ale, Scalo = same as F ig, 33*
Figure 35* Hirstionyssus "B" n, sp. Ventral viow of tarsus II in the female. Scale = 50 microns.
263
Figui'a 3^. Hirstionyssus talpae Zemskaya. Vontral view of the female. Scale = 100 microns.
Figure 37* Hirstionyssus talpae Zemskaya. Dorsal view of th e fem ale. Scale = same as Fig. 36 .
Figuro 38. Hirstionyssus talpae Zemskaya. Ventral view of tarsus II in the female. Scale = 50 microns.
265
Figure 39• Hirstionyssus affinis (Jameson). Ventral view of the female. Scale = 100 microns.
Figure 40. Hirstionyssus affinis (Jameson). Dorsal view of the female. Scale = same as Fig. 39.
Figure 41, Hirstionyssus affinis (Jameson). Ventral view of tarsus II in the female. Scale = 50 microns.
Figure 42, Hirstionyssus affinis (Jameson). Ventral view of tarsus IV in the female. Scale = same as Fig, 4l.
26 ?
I
Figure ^3* Hirstjonyssus "D" n. sp. Ventral view of the female. Scale = 100 microns.
Figure hh, Hirstjonyssus "D" n. sp. Dorsal view of the fem ale. S cale = samo as F ig . h-3.
Figure h-5. Hirstjonyssus "D" n. sp. Ventral view of tarsus II in the female. Scale = 50 microns.
Figure ^6. Hirstjonyssus "D" n. sp, Ventral view of tarsus IV in the female. Scale = same as Fig, 45*
269 2?0
J VJUu
\ Figure ^7- Hirstjonyssus occidentalis (Swing), Ventral view of the female. Scale = 100 microns,
Figure ^8. Hirstjonyssus occidentalis (Swing), Dorsal view of the female. Scale = same as Fig,
Figure ^9* Hirstjonyssus occidentalis (’.Swing), Ventral view of tarsus II in the female. Scale = 50 m icrons.
Figure 50. Hirstjonyssus occidentalis (Swing), Ventral view of tarsus IV in the female. Scale same as Fig.
271 ’viUkA', Figure 51. Hirstjonyssus eutamiae Allred and Beck* Ventral view of the fenale. Scale = 100 microns.
Figure 52. Hirstjonyssus eutatniae Allred and Beck. Dorsal view of the female. Scale = same as Fig. $1,
Figure 53. Hirstjonyssus eutamiae Allred and Beck. Ventral view of tarsus II in the female. Scale = 50 microns.
Figure 5^. Hirstjonyssus outamiao Allred and Beck. Ventral view of tarsus IV in the female. Scale = same as F ig. 53.
273
Figure 55* Klrstionyssus cynomys (Radford). Ventral view of tho female. Scale = 100 microns.
Figure 56. Klrstionyssus cynomys (Radford). Dorsal view o f th e fem ale. Scale = same as F ig. 55*
Figure 57* Klrstionyssus cynomys (Radford). Ventral view of tarsus II in the female. Scale = 50 microns.
Figure 58. Klrstionyssus cynomys (Radford). Ventral view of tarsus IV in the female. Scale = same as F ig. 57.
275
Figure 59. Hirstion.yssus utahensis Allred and Feck. Ventral view of the female. Scale = 100 microns.
Figure 60. Hirstionyssus utahensis Allred and Beck. Dorsal view of the female. Scale = same as Fig. 59.
Figure 6l. Hirstionyssus utahensis Allrod and Beck. Ventral view of tarsus II in the female. Scale = 50 microns,
Figure 62. Hirstionyssus utahensis Allrod and Beck, Ventral view of tarsus IV in the female. Scale = same as F ig. 6 l .
277
Figure 6 3 . Hirstionyssus latiscutatus (do 1-feillon and Lavoipierro), Ventral view of the fonale. Scale = 100 microns.
Figure 6^. Hirstionyssus latiscutatus (do r-feillon and Lavoipierro). Dorsal view of the fonale. Scale — same as Fig. 63 *
Figure 65* Hirstionyssus latiscutatus (de I'eillon and Lavoipierre). Ventral view of tarsus II in the female. Scale - 50 m icro n s.
279
Figure 66. Hirstionyssus ’Vft" n. sp. Ventral view of tho female. Scale = 100 microns.
Figure 6?. Hirstionyssus "A" n. sp. Dorsal view of tho female. Scale = same as Fig. 66,
281 232 Figure 63. Hirstionyssus torus Allred and Bock. Ventral view of the female. Scale = 100 microns,
Figure 69 . Iiirstjon.yssus torus Allred and Beck. Dorsal view of the female. Scale = same as Fig, 68.
Figure 70. Hirstionyssus torus Allred and Beck. Ventral view of tarsus IV in the female. Scale ~ 50 microns.
283 zeh
r r ®\ Figure 71, Hirstionyssus obsolctus Jameson, Ventral view of the female. Scale = 100 microns.
Figure 72. Hirstionyssus obsoletus Jameson. Dorsalview o f th e fem ale. S cale = samo as F ig. 71.
285 286
Y
§ Figure 73 • Ilirstionyssus jsabollinus (Cudenans), Ventral view of the female, Scale = 100 microns.
Figure 7^» Hirstionyssus jsaholljnus (Cudenans). Dorsal viei-J of the fem ale. S cale = same as F ig . 73•
237
Figure 75. Kir stionyssus broviseta Strandtmann and Korlan. Ventral view of tho female. Scale = 100 microns.
Figure ?6. Kirsti on.yssus broviseta Strandtmann and I'orlan. D orsal viev/ of tho female, Scale = same as Fig, 75*
239 290 Figure 77. Kirstionyssus neotonae (Fads and Rightovrer ). Ventral viev; of tho female. Scalo = 100 microns.
Figuro 78. Hirstionyssns nootomae (Fads and iiightower). D orsal viev; of the female. Scale =samo as Fig. 77
291 ZoZ Figure 79* Kirstionyssus bisotosus Allred. Ventral view of the female. Scalo = 100 microns.
Figure 80. hirstionyssus bisotosus Allred, Dorsal view o f tho fem ale, Scalo = same as Fig, 79•
293
Figure 81. Hirstionyssus Stafford! Strandtmann and Hunt Ventral view of the female. Scalo = 100 microns.
Figure 82, Hirstionyssus staffordi Strandtmann and Hunt D orsal view of the female. Scale = same as Fig. 81
Figure 83• Hirstionyssus staffordi Strandtmann and Hunt V entral view of tarsus II in the female. Scalo = f>0 m icrons.
295 296
\ REF IITCES CITED
Allred, D, K, 1957s-. A new species of mite, }Iirstionyssns bisetosus, from the nest of the desert woodrat, .’eotom.a lepida lepida Thomas (Acarina: Dermanyssidae), Proc. Entomol, Soc, ’Washington 59(2):83-89.
______, 1957b. Fites fonnd on mice of the genus Feromvscus in Utah, II. Family Dernanyssidae, Amer, Kidland Uatur, 57l2T^50“460.
______. 1962. Fites on grasshopper mice at the Levada Atomic Test Site. Great Basin Hatur. 2 2 (4 ):101-1 0 4 ,
______. 1963. Fites from pocket mice at the Levada Test Site (Acarina). Proc, Entomol. Soc, Washington 65 (3 ):231-233.
, and D E, Beck, 1966, Kitos of Utah mammals, Brigham Young Univ. Sci. Bull., Biol. Ser. 8(1):1-123.
______, and K, A. Goates. 1964a, Kites from v;oodrats at tho Levada Test Site. J. Farasitol. 5^(1)*171. A * ______, and K. A. Goates, 1964b. !E.tes from mammals at the Levada Test Site. Great Basin Latur. 24(2):71-73»
, and E, J. Roscoe. 1957* Parasitic mites in desert woodrat nests, with notes on free-living foi’ms. Trans, Amer. Kicroscop. Soc. 76(4)-.389-403.
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