,**»'
W'V.
.*«*^*«***lf*4*,.
^ t i' # i # ^■
^ % % ^ M^ ?>^ :*.- ^ <f ^' %■ * * f- 1^- !!• '{^ % f f <^ ^
*,v^*^v, *»**.*■*'
>'■»■
|
H |
|
|
■ |
|
Kl-
rrtrt
^^^A^V^
^^■jm
^'^^'rP
#.,i.
i.m :'«s
t, M
"-■ c ., < «.
i ^' * * « f -i- ii
c#ji
I I I /r r f.% » » f I- It » Vs * *' » pV» |. i ■ * * r #■
r>
ril
•Li^
mmm^^^^^mw^^^^??p'^^/^,w?S'*'
W ^' #■ ^. !^ # ^
■1" «'
■
HARVARD UNIVERSITY
Library of the
Museum of
Comparative Zoology
c~-
Brigham Young University Science Bulletin
MUS. CO MP. zooi_:
LIBRARY
OCT 2 9 1971
HARVARD UNIVERSITY
TICKS Of JAPAN, KOREA, AND THE RYUKYU ISLANDS
by
Noboru Yamaguti Vernon J. Tipton Hugh L. Keegan Seiichi Toshioka
BIOLOGICAL SERIES — VOLUME XV, NUMBER 1 AUGUST 1971
BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN BIOLOGICAL SERIES
Editor: Stanley L. Welsh, Department of Botany,
Brigham Young University, Prove, Utah
Members of the Editorial Board:
Vernon J. Tipton, Zoology Ferron L. Anderson. Zoology Joseph R. Murdock, Botany Wilmer W. Tanner, Zoology
Ex officio Members:
A. Lester Allen, Dean, College of Biological and Agricultural Sciences Ernest L. Olson, Chairman, University Publications
The Brigham Young University Science Bulletin, Biological Series, publishes acceptable papers, particularly large manuscripts, on all phases of biology.
Separate numbers and back volumes can be purchased from Publication Sales, Brigham Young University, Provo, Utah. All remittances should be made payable to Brigliam Young University.
Orders and materials for library exchange should be directed to the Division of Gifts and Exchange, Brigham Young University Library, Provo, Utah 84601.
Brigham Young University Science Bulletin
TICKS OF JAPAN, KOREA, AND THE RYUKYU ISLANDS
by
Noboru Yamaguti Vernon J. Tipton Hugh L. Keegan Seiichi Toshioka
BIOLOGICAL SERIES — VOLUME XV, NUMBER 1 AUGUST 1971
Financial support for Volume 15, Number 1 was furnished by the U.S. Army Medical Research and Development Command under Grant No. DADA 1 7-71 -G-9 340.
TABLE OF CONTENTS
ABSTRACT 1
INTRODUCTION 1
MATERIALS AND METHODS 2
A MAP SHOW INC GEOCRAPHICAL LOCATION OF
JAPAN, KOREA, AND THE RVUKYU ISLANDS (Fig. 1) 3
KEY CHARACTERISTICS OF TICKS (Figs. 2-4) 4
ILLl'STRATED KEY TO THE GENERA OF TICKS OF
JAPAN, KOREA AND THE RYUKYU ISLANDS (Fig. 5) 7
FAMILY ARC:ASIDAE 2
Ke\ to the .species of the hunily .\rgasidae 2
Genus Arg(;.« 8
Argas japoniciis Yamaguti Clifford, and Tipton, 1968 8
Argas vespcrtilionis ( Latreille, 1802) 11
GeniLs OrnithodoTOs 15
Ornithodoros capetuis Neumann, 1901 15
FAMILY IXODIDAE 18
Genius Amblyomma 18
Kev to the species of the genu.s AnMijomma 18
Amhhjommn geocmydae ( Cantor, 1847 ) 18
Amhlyommti tiitidum Hirst and Hirst. 1910 23
Ambhjoninui testudinarium C. L. Koch, 1844 25
Genus Boophilus 30
Boophihis microplus ( Canestrini. 1888 ) 30
Genui Dcrviuccntor 36
Genus Haemaphysalis 42
Kev to the species of the genus Haemaphy salts 42
Hacmaphys(dis camptinuhita \\'arburton, 1908 43
Haemaphysali-s conciniui C. L. Koch. 1844 49
Haemaphysalis sp. {H. cornigera group) 54
Haemaphyscilis doenitzi Warburton and Nuttall, 1909 59
Haemaplu/sulis fUiva Neumann, 1897 59
Hacmaphi/S(dis formoscmis Neumann, 1913 68
Haernapliysalis fujls:imi Kitaoka, 1970 68
Haemaphysalis hystricis Supino, 1897 77
Hacmaphi/salis jajioidca \Varburton, 1908 83
Hacmaptu/salis kitaokai Hoogstraal, 1969 87
Haemaphysalis longicornis Neumann, 1901 94
Haemaphysalis megaspinosa Saito. 1969 100
Haemaphysalis pental-igi Pospelova-Shtrom, 1935 105
Haem'.iphysalis wellingtoni Nuttall and Warburton. 1908 110
Genus Ixodes 112
Key to the species of the genus Ixodes 112
Ixodes acutitarsus Karsch, 1880 113
Ixodes angustus Neumann, 1899 115
Ixodes gramdalus Supino, 1897 119
Ixodes lividus C. L. Koch, 1844 124
Ixodes motws])itiosiis Saito, 1967 129
Ixodes nipjioiiensis Kitaoka and Saito, 1967 129
Ixodes oiatus Neumann, 1899 135
Ixodes persulcatus Schulze, 19.30 142
Ixodes philipi Kairans and Kohls, 1970 148
Ixodes sigtuitus Binila, 1895 150
TABLE OF CONTENTS (contimud)
Ixodes simplex simplex Neiiniaiin, 1906 155
Ixodes tanuki Saito, 1964 158
Ixodes turdiis Nakatsuji, 1942 161 |
Ixodes uriac White. 1852 161 j
Ixodes vesj>ertilionis C. L. Koch, 1844 165 I
Geniis Rhipicephalus 170 I
Rhipiccpludus sanguineus Latreille. 1806 170
OOl'liTFlH. HIiCORDS 175 I
I
ACKNOWLEDGMENTS 177 •
LITERATURE CITED 178
APPENDIX 1. NOTATIONS ON THE LITERATURE CITED 185
APPENDIX 2. COLLECTION RECORDS 186
APPENDIX 3. LIST OF MAPS AND TEXT FIGURES 219
APPENDIX 4. INFORMATION ON SPECIMENS ILLUSTRATED 222 ,
APPENDIX 5. CORRECTED SCIENTIFIC NAMES OF HOSTS '
RECORDED IN THE LITERATURE , 227
L,
TICKS OF JAPAN, KOREA, AND THE RYUKYU ISLANDS
by
Noboru Yamaguti,' Vemon J. Tipton,- Hugh L. Keegan,' and Seiichi Toshioka^
ABSTRACT
This publication is a revision of Ixodid Ticks of Japan, Korea, and the Ryukiju Islands by Keegan and Toshioka ( 1957 ) . The Hck fauna of these areas is represented by 36 species in the genera Ar^fl.y, Ornithodoros, Ambhjomma, Boo- philus, Dermacentor, Haenmphij sails, Ixodes, and Rhipicephalus. Three species of the family Argasidae and 10 species of Ixodidae were not treated in the original edition. The additional information contained in this revision was de-
rived from field collections and a thorough re- view of the literature. Keys are provided for the identification of genera and species. For each species we have given synonymy, a brief diag- nosis, geographical distribution, hosts, and, if known, the biology and relationship to disease. Collection data for material actually examined or recorded in the literature are given in chart form at the end of the paper. Illustrations of 65 adults and 53 immatures are provided.
INTRODUCTION
The importance of ticks as vectors of viral and rickettsial diseases of man has been re- viewed by H(X)gstraal (1966, 1967a, 1967b). Ticks are also important vectors of pathogens affecting domestic and game animals. The man- ual, Ixodid Ticks of Japan, Korea, and the Rt/u- ktju Islands by Keegan and Toshioka ( 1957), rep- resented a significant contribution to our knowl- edge of tick vectors of disease. Inasmuch as there have been several biological and taxonomic studies of the tick fauna of Japan published since 1957, and to a lesser extent Korea and the Ryukyu Islands, Colonel Keegan suggested that Tipton and Yamaguti revise the original paper. We have supplemented information from the literature with our own collecting and life cycle studies at the 406th Medical Laboratory. Some additions and changes have been made in the format, such as the inclusion of the argasid ticks, a diagnosis for each species, an illustrated key to the genera, and a map for each species showing distribution based on all collection records avail-
able. We have also chosen to present the collec- tion data in chart fonn rather than as a part of the text. The format used by Elbl and Anastos ( 1966) has been used as a guide in the prepara- tion of the collection data chart.
Much of the infoniiation contained in this paper has been copied verbatim from the origi- nal 1957 manual. The order in which the authors are listed does not infer that Keegan and Toshioka have made any less contribution than the other two authors, and as a matter of fact this paper would hardly have been possible without the original manual as a starting point. Furthennore, we received constant encourage- ment and assistance from Colonel Keegan and Dr. Toshioka.
As in the 1957 manual, the generic classifica- tion given by Anastos (1950) and Hoogstraal (1956) has been followed. We have also relied heavily on the assistance of Mr. Glen M. Kohls ( Rockv Mountain Laboratory, Hamilton, Mon- tana), Dr. Harry Hoogstraal (Naval Medical
*[)eparlnient of Parasitology. Tokyo Women's Medicil (College. Tokyo. Japan.
-Departnient nf 7.noIogy. Rrigliani YoiinR Universily. I'rovo. Utah.
'ncpartnifnt of Prpveiitive Mi'dii inn. Sdionl of MedMine. University of MississiiJpi. Jatksoii. Mississippi.
'Department of Kntomologw -HHuli .MeJital I.aboratory. U.S. Army ,\Iedical Command. .\P0 San Francisco. 06543, US.\.
Bhigham Young University Science Bulletin
Research Unit No. 3, Cairo, Egypt), and Dr. Shigeo Kitaoka (National Institute of Animal Health, Tokyo).
Taxonomieally important striictnre.s are labeled in Fig. 2 to 4. A detailed discussion of tick morphology is not given in this paper. Such infomiation may be obtained from texts of medi- cal entomology or the excellent monographs of Nuttall and Warburton (1908, 1911, 1915).
Synonymy listed under the specific name in- volves only those papers in which specimens from Japan, Korea, or the Ryukyu Islands were examined or discussed by those authors. Though there have been several papers dealing with specimens from Taiwan ( Formosa, previously controlled by Japan before World War II), they were purposely excluded as sources of synonyms and literature, except in those cases where data contained were pertinent to the populations in the areas discussed. Hence, the synonymy pre- sented here is less extensive than that of other monographic papers. Literature containing bio- logical information germane to this paper is included.
With practicality in mind a brief diagnosis of each species is given rather than a detailed description. Keys to the immature forms are not included, but many illustrations of larvae and nymphs, as well as adults, which were not pro- vided in the publication of Keegan and Toshioka (1957) have been added. We believe this will be beneficial to field workers concerned with tick biology.
At the time of the writing of this paper (1970), the political boundary bet^veen Japa- nese and American administration of the Ryukyu Islands lies between Amami Gunto and Okinawa Gunto, the former having already reverted to Japan. In 1972, the remaining islands in Oki- nawa and Sakishima Gunto ( including Miyako and Yaeyama Retto) will return to Japanese administration, probably as Okinawa Prefectiire. Even though Amami Oshima is politically Japa- nese, for the purpose of geographical clarity, it is shown in Appendix 2 as being part of the Ryukyu Islands. It is administered by Kago- shima Prefecture in Kyushu.
MATERIALS AND METHODS
Most of the material examined as a part of this study was collected during extensive field trips on the four main islands of Japan and on islands of the Ryukyu Archipelago. A few speci- mens were provided by one of us (Tipton) in connection with field excursions made to Korea.
Engorged females were kept in vials until oviposition, and larvae were fed on ears of rab- bits so that laboratory life cycle studies could be initiated. Unengorged females ( freshly collected from the field) were likewise reared in the laboratory. These procedures enabled us to properly associate adults with immature forms. Tentative identifications were recorded on cards with collection data. One card per host was prepared. Specimens from each lot were sent to Mr. Glen M. Kohls, Rocky Mountain Labora- tory, for confinnation of identifications.
Unmounted specimens collected in Japan, Korea, and the Ryukyu Islands and preserved in 70? alcohol were used as a basis for most of the illustrations. Often it was necessary to re- move legs and spiracular plates in order to get a clear view of contour and chaetotaxy. In some instances the capitulum of the adult and whole body of the nymph or larva were mount- ed temporarily in Hoyer's solution on a cavity slide for detailed examination of the chaetotaxy and dentition of the hypostome.
Illustrations were prepared by the artists named in the acknowledgments.
Family Argasidae Canestrini, 1890
Nonscutate "soft ticks." Sexual dimorphism very slight. Integument of adults and nymphs mammilated, wrinkled, leathery, granulated, or with tubercles. Capitulum of adults and nymphs in camerostome on ventral side; apart from anterior margin of body. Palpal articles sub- equal, leglike, free, never fused in all stages. Porous areas absent. Eyes present or absent; when present, on the supracoxal folds. Spiracles of adults and nymphs usually anterior to coxae IV. Pulvilli usually absent or nidimentary in adults and nymphs, whereas functional in larvae. Number of nymphal stages variable. Type genus: Argas Latreille, 1796.
Key to the Species of the Family Argasidae
1. Suture between dorsal and ventral surface of body absent; dorsal surface mammilated;
on seabirds (Fig. 9, 10) . Ornithodoros capensis
Suture between dorsal and ventral surface of body definitely present; dorsal surface
finely wrinkled Genus Argas 2
2. R()d\ iicarK- round; on bats (Fig. 7. 8) Argas vesjwrtiUonis
Body longer than wide; on swallows (Fig. 5, 6) A. japonicus
Biological Slhies, Vol. 15, No. 1
Ticks
Maf I. Map of Japan. Kore;i and the Rynkyii LslaiuLs.
Bricham Young University- Science Bulletin
capi tulum cervical pit
cervical groove punctation
scutum
lateral groove -
porous area scapula lateral carina cervical groove - -
scutum marginal groove
pregenital plate - genital aperture
median plate epimeral plate
adanal plate anal plate
hypostome
internal spur
coxa I
coxa II
coxa III -coxa IV -external spur
spiracular plate "genital aperture
genital groove -anus
anal groove
Dorsal view of female
Ventral view of female
Fig. 1. Key characteristics of ticks - 1 (genus Ixodes)
Bi()i,oc:iCAL Series, Vol. 15, No. 1
TicK.s
eye
ornate marking
fovea
festoon
internal spur external spur
accesory shield
adanal shield
postanal -median groove
Female of genus Amblyomnia (dorsal view)
Male of genus Rhipicephalus (ventral view)
hood
camerostome
cheek
— eye
capitulum genital aperture
eye
suprocoxal fold spiracle
anus
preanal groove
transverse postanal groove
genus Ornithodoros
Fig. 2. Key characteristics of ticks - 2.
Bhigham Younc University Science Bulletin
article III
article II
-- article I basis capituli --porous area- cornua
-external profile
__ postero-external juncture
--dorsointernal setae
-hypos tome
article IV
infrainternal or ventrointernal setae
-article I
-auricula
basis capi tul i
genus Ixodes
genus Haemaphysal is
apicoventral spur or hook
posthypostomal seta
pul villus
claw
hump
Mailer's organ
macula
- goblet
Hypostome
Tarsus I
Spiracular plate
Fig. 3. Key characteristics of ticks - 3.
BioLotiiCAL Series. \*oi,. 15. No. 1
Ticks
Family Arqosidoe
Capitulum on under side of body. Scutum absent.
Family Ixodidoe
Capitulum at anterior end of body. Scutum or dorsal shield present. (Short in? Long in «f )
Usually with definite sutural line separating dorsal and ventral sur- faces; body margin distinctly flattened and structurally differ- ent from remainder of integument. /
Sutural line absent. Body margin structurally not different from remainder of integument.
Anal groove contouring the anus Anal groove contouring the anus posteriorly, festoons present. anteriorly, festoons absent.
Argos
Ornithodoros
Eyes present. Second segment of Palpi not projecting later- ally beyond the basis capituli.
Eyes absent. Scutum inornate. Second segment of Palpi usually projecting laterally beyond the basis capituli except H. kitaokai?
Palpi long and slender, longer than the basis capituli. Scutum ornate.
Palpi short and broad, not longer than the basis capituli.
Haemaphysalis
Basis capituli hexagonal
Amblyommo
I
Palpi very short and ridged dorsally and laterally. Coxa I with two very short spurs. Festoons absent.
Basis capituli rectangular. Scutum ornate. Coxa IV of male much larger than other coxae.
Palpi not unusually short, not ridged. Coxa I with two long spurs. Festoons present.
Dermacentor
Coxa I
BoQphilus
Coxa I
? J
tL kitaokai
Fio. 4. Illustrated key to the genera of ticks of Japan, Korea, and the Ryiikyu Islands.
BnioHAM YouNO University Science Bulletin
Genus Argas Latreille, 1796
Body distinctly flattened dorsoventrally; dor- sal and ventral surfaces subequal in area. Body margin flattened even when depleted, consisting of radial striations or <|uadrangular plates. Su- tural line separating dorsal and ventral surface present. Integument leather)' or finely wrinkled, intermingled with small "buttons"; discs ar- ranged more or less radiallv. Eyes absent. Adults and nymphs similar. Parasitic principally on birds or l)ats. Type species: Argas reflexiis (Fabricius, 1794).
Argas japonictis Yamagnti, Clifford, and Tipton (Fig. 5 and 6)
Argas japoniciis Yamaguti, Clifford, and Tipton, 1968:453-459, Fig. 1-17; Uchikawa and Sato, 1968:157-161, Fig. 1, 1969:95-97; Saito, lijima, and Minai, 1969-39-41.
Argas .sp.: Inatomi and Yamaguti, 1960:17-18; Uchikawa, Sato, and Kugimoto, 1967:141- 151, Fig. 1-20.
Argas reflexus var. japonicus Yamaguti and Ina- tomi, 1961:142.
Argas persicus (not Oken, 1818): Hara, 1963: 123-125, Fig. 1.
Discussion:
The occurrence of this soft tick was first re- ported by Inatomi and Yamaguti ( 1960 ) from nests of the Japanese striated swallow, Hirundo dattrica juponica, which were built under eaves of the post office at Niimi City, Okayama Pre- fecture, Honshu. They gave a brief account of adult morphology. The following year, Yama- guti and Inatomi ( 1961 ) examined immature forms from this locality and considered this population to be a new variety of Argas reflexus, which they designated japonicus.
Hara ( 1963) reported the occurrence of Argas persicus from nests of the house martin at Agematsu, Nagano Prefecture. Uchikawa, Sato, and Kugimoto ( 1967 ) found a large number of specimens of Argas sp. in swallow nests built under eaves of the University Hospital of Shin- shu University, Matsuinoto City, Nagano Pre- fecture. Yamaguti, Clifford, and Tipton ( 1968 ) examined this material, including Hara's persi- cus. and indicated that all specimens were iden- tical to the new species, A. japonicus, described by them.
Diagnosis:
This species is similar to A. coolet/i Kohls and Hoogstraal, 1960, of western North America, A. lagenoplastis PVoggatt, 1906, of Australia, and
A. vulgaris Filippova, 1961, of eastern Russia, but adults may be differentiated from these species on the basis of number and distribution of setae on the tarsi, basis capituli, and anal valves. Larvae may be diftcrentiated on the basis of length, number, distribution, and stnic- ture of body setae (A. lagenoplastis); dentition of the hypostome and number and distribution of body setae on the posterior (juadrant (A. cooleiji; and dentition of the hypostome and size of the dorsal plate (A. vulgaris).
This swallow argasid is easily distinguished from other soft ticks of the Japanese fauna in that the body is ovate, the ventral "paired or- gans" are lacking, the mouth parts are not so close to the anterior margin of the body as in vespertilionis, and the tarsi have distinct sub- apical dorsal protuberances.
DlSTRIBUTIGN AND HoSTS:
To date this species is known only in Japan and Korea. Reported hosts are the Japanese striated swallow, Hirundo daurica japonica. and the Japanese house martin, Deliclion urbica da.njpus. Under experimental conditions A. japonicus will feed on chickens.
ArgdS japonicus
9 Collected and examined
O Collection records from
literature 3 Combination of above ft
O
Map 2. Known distribution of Argas jtiponUus.
Biological Series, Vol. 15, No. 1
Ticks
X A^niia.
Fic. 5. Argas japonicus, female and male.
10
Bhicham Young University Science Bulletin
-r A-^Ac
Fig. 6. Argas japnnicus, lana ani> nvTnph.
Biological Series, Vol. 15, No. 1
Ticks
H
Biology:
Uchikavva et al. (1967) and Uchikawa and Sato (1968, 1969) examined a natural popula- tion in Nagano Prefecture and studied a labora- tor\' colony led on chicken.s. Tiiey found the tick.s to be very active, and .some of them in- truded into the wards of the hospital (Shinshu UniversitA' Hospital) during the period from late March to September when the host birds were nesting. .■Ml de\eloping stages, as well as eggs, were found in the holes and crevices of mud used in construction of nests. After the host birds leave and migrate to southern regions, the ticks stav in the same niches and overwinter there. Uchikawa et al. (1967) state that the winter colonv consists of unfed specimens plus those which have fully digested a previous blood meal. Unfed larvae die within three weeks when kept at 30 C, but they survive until the following spring without a blood meal under natural conditions. There are at least two instars in the nvmphal stage, and both are often found in natural populations. In laboratory colonies 4th instar nymphs are seen only occasionally. Morphological differences among nymphs of dif- ferent stages are generally very slight. Uchi- kawa and Sato ( 1968 ) studied the morphological differences of these nymphs and found that the apicoventral setae on the tarsi change with molting, and this serves to discriminate the nymphal instars. In the laboratory Uchikawa et al. (1967) found that larvae, nymphs, and adults began to feed on chickens after a short preparasitic period of about 3 days. Larvae fed for 3-6 davs, and nvmphs and adults for 9 min- utes to an hour. At 30 C the postparasitic period for larvae was 5-12 days, for 1st instar nymphs it was 8-18 davs, and 2nd and 3rd instar nymphs which molted into adults it was 12-20 days. The period was prolonged at lower temperatures. Each female laid .36-200 eggs during a 5-12 day period, and the larvae hatched in 12-16 days.
At the 406th Medical Laboratory rearing ex- periments on chickens were undertaken under conditions somewhat different than those de- scribed by Uchikawa et al. (1967). Newly liatched larvae were reared individually, and some of them emerged to adults after passing through the 4th nvmphal instar. Most of the larvae developed to adults, but they remained in a postparasitic (|uiescent phase without ovi- positing for more than one year. After feeding, the bodv outline of the larvae changed and closelv approached the nvmphal shape. The details of rearing experiments will be sum- marized and reported elsewhere in the near future.
Disease Relationship:
There are .some reports of invasion of dwell- ings by this tick, and inhabitants have often complained of tick bites (Inatomi and Yama- guti, I960; Hara, 1963; Uchikawa et al., 1967; and Uchikawa and Sato, 1969). However, there is no definite evidence of biting by A. japonicus, and it may be that bites were caused by the fleas, FwntopstjUa setiger Smith or Ceratoplnjl- Itis farreni chaoi Smit and Allan, which were often associated with A. jajwnicus in swallow nests.
The relationship of this tick to human dis- ease is not known, but because it lives in close proximity to man the potential for transmission of disease to humans is real.
In 1960 Yamaguti found a nestling bird heavily infested with larvae and nymphs of A. japonicus. The bird appeared to be weakened to the point of death.
Argas vespertilionis (Latreille)
(Fig. 7 and 8)
Caris vespertilionis Latreille, 1802/1803:67-68;
Kishida, 1936:142. Argas vespertilionis: Kishida, 1927:986, Fig. 1899,
1947:975, Fig. 2775; Saito, 1955:7-12, Fig.
7-14; Kamo, 1962:173; Kishida and Asa-
numa, 1965:395, Fig. 214; Asanuma, 1965a:
124; Yamaguti, Clifford, and Tipton, 1968:
453. Argas sp. Ito and Saito, 1954:563-564; Saito,
1955:7-12, PI. II.
Discussion:
The first record of A. vespertilionis in Japan is probably Kishida's account in the 1927 edition of the Illustrated Encyclopedia of the Fauna of ]apan. He stated that this species occurs in Honshu, Shikoku, Kyushu, Korea, and the Ryu- kyu Islands and listed seven species of bats as host animal in the above areas. Kishida (1936) also reported his collection of Caris vespertilionis in Korea. Saito ( 1955 ) collected Argas sp. from bats in Niigata City and reared the species on the host, Pipistrelhis ahramus. He did not defi- nitely identify the tick, but he mentioned that his material was very close to A. vespertilionis. Al- though specimens were not available for study, morphological characters shown in photographs of the tick are similar to those of vespertilionis. Additional material h;is been collected from Pipistrelhis spp. at Fukuota, Kyushu, and Sap- poro, Hokkaido, and from Vespertilio superans at Shojo, Fukushima Prefecture, Honshu. Tlie lot from Ves])ertiUo superans was sent to Kohls and it was his opinion that the specimens differ
12
Bricham Young University Science Bulletin
Fic. 7. Argas vespertilionii, female and male.
Biological Series. Vol. 15, No. 1
Ticks
13
Fig. 8. Argas vespertilionis, lana.
14
Bhigham Young University Science Bulletin
only slightly in integumental characters from Egyptian vespertilionis and may well be the same species.
Hoogstraal ( 1956) considers the A. vesper- tilionis group to include European and African vesj)ertilionis, A. pussihis Kohls, 1950 on Palwan Island in the Philippines, and closely related populations of uncertain species status, and that this group ranges tliroughout the continent and island groups of the world except in the Americas.
Diagnosis:
This round bat-tick is the sole argasid species parasitizing bats in Japan and Korea and is readily distinguished from other argasids in this fauna. The Ixxly is circular, the anterior margin has a very slight anterior projection, in the peripheral area there are rectangular "cells" and a definite sutural line dividing the dorsal and ventral surfaces, there are distinct ventral "paired organs" posterior to the anus, and the mouth parts are close to the anterior margin of the body.
Distribution and Hosts:
Hoogstraal (19.56) gave the distribution and hosts of the vespertilionis group as England, the Netherlands, Sweden, Spain, Germany, Africa, Korea, China, the Philippines, and Ceylon. The group is also known to occur in southern India, Cambodia, Australia, France, Italy, Russia, and Japan. Sugimoto (19.36b) cited Sauter's collec- tion of A. vespertilionis on Taiwan. Almost all species of bats within the distributional range of A. vespertilionis may be parasitized by this tick, and all stages of the tick mav infest these bats. Both nyiuphs and adults occasionally at- tack man.
Biology:
Saito (1955) reared this species on the host bat in the laboatory but did not mention details of the life cvcle, except that the nvmphs fed at night, adults appeared about three months after passing through three nymphal instars at 25-.30 C, and that below 10 C the ticks of each stage went into hibernation.
Hoogstraal (1956) reported that A. vesper- tilionis was succ-essfully reared in his laboratory at 80-90 F and 40-50? relative humidity.
Eggs hatched 16-20 days after oviposi- tion and larvae fed as soon as 4 days after- wards. Larvae fed for 14-31 days, usually 17-19 davs, and molted 5-10 davs later. Nvmphs fed 3-4 davs alter molting and they usuallv fed twice, followed bv a molt S or 9 days after the first meal and 12-14 days
Maf .3. Known distribution of Ar<^iis lespcrtilionis.
after the second meal. Nymphs become re- plete in 20-50 minutes, usually .30-40 min- utes. Adult males may develop from the first nvmphal molt, but usually nymphs molt twice before becoming adults. Males and females may feed within 7 davs after molt- ing, and they feed for .30-40 minutes. No females oviposited within 6 months after the nvmphal-adult molt, even though with males continuously and given two to six blood meals. The first oviposition follows a blood meal bv about a week and appears to trigger a physiological release mecha- nism, because in several instances females have deposited fertile eggs three months afterwards with or without a meal.
Disease Relationshii':
Nvmphs and adults may attack man (several authors), and in Japan, Kamo (1962) reported one instance of a human lieing bitten by this tick. Reports indicate that mild itching may persist for several weeks. This tick may be a vector of a spirochete of bats, but conclusive supporting evidence is not available. The rela- tionship of this tick to human disease has not been studied.
Biological Sebies, Vol. 15, No. 1
Ticks
15
Genus Ornitlioduros C. L. Kocli, 1S44
Body more or less flattened. Dorsal surface usually con\'ex when fully fed. Body margin structurally not different from adjacent areas. Suhiral line absent. Capituluni suhtenninal or apart from anterior margin of the body. Hood, cheeks, and camerostoine present. Dorsal humps and protuberances prominent on tarsi. Eyes absent or present.
Type Species: Oniithodoros s(ivi<;^mii (Audouin, 1826).
Ornithodows capensis Neumann (Fig. 9 and 10)
Ornithodows talaje var. capensis Neumann,
1901:2.58. Ornithodows cajyensis Neumann, 1901; Kohls,
1957a: 89-90, Fig. 1; Keegan and Toshioka,
1957:23; Asanunia, 1960:94, 1965a: 124, Fig.
7.16.
DiSClTSSION:
The original description of this species was based on material taken from nests of penguins on islands off the coast of Cape Colony, South .\frica. Kohls (1957a) considers capensis to be distinct enough to warrant full specific status and to be readily distinguishable from talaje on the basis of morphological differences as well as host relationship and distribution. In Japan, O. cai>ensi.^ was first found by Asanuma associated with Ixodes signotus in July 1955 at Kabujima, Aomori Prefecture, Honshu. He did not, how- ever, refer to this soft tick in his two papers on Ixodes signatus published in 1955 and 1957. In his paper of 1960 Asanuma assembled the known information on O. ca})ensis in Japan up to that time and indicated he had collected this species in 1955. Later the tick was collected by 406th Medical Laboratory personnel at the same lo- calitv', as well as from several other localities.
Dl\gnosis:
This soft tick is the only species of the genus OrnitJiodoros parasitizing sea birds in this area and is easily recognized by the generic char- acters used in the key. As reported by Kohls (1957a), this species resembles O. amhhis, but differs from that species in having a more point- ed anterior margin of the body and cheeks with partially overlapping mouth parts.
Distribution .\nd Ho.sts:
O. caj)ensis is known to be widely distributed throughout the world. Kohls ( 1957a ) reviewed the distributional records of this species and listed the names of islands where the species is
known to occur. These include islands off the coasts of South Africa, southern Australia and Japan, and additional islands in the Atlantic, Pacific, and Indian oceans.
Amerson (1968) states: "It is presently known from 32 islands and at sea in the Central Pacific and immediate surrounding areas from 22 sea and shore bird species, as well as from man, the European rabbit, sea turtles (Celonia mijdas), and ground litter . . . Since 1901 O. capensis has been recorded aroimd the world in the tropical and temperate regions and has been associated with 29 species of sea and shore birds throughout the world (from published and unpublished records)." Amerson also gives a world distribution map of this species.
In Japan O. capensis has been found on the black-tailed gull. Lams crassirostris, the streaked shearwater, Calonectris Jeucomelas. and the an- cient auk, Synthliborainplnis antiqitus. Accord- ing to Asanuma (1960), O. capensis will attack domestic fowls.
Biology:
O. capensis has been reared on chickens at the 460th Medical Laboratory. Larvae fed for 5-6 days and molted to 1st instar nymphs 1-4
.M.-^p 4. Known distribution of Ornithodoros capensis.
16
BniGHAM Young Univehsity Science Bulletin
2.0 mm
a
0, 5 mm
0.5 mm
0.5mm
Fig. 9. OrnitJwdoros capensis, female and male.
Biological Series, Vol. 15, No. 1
Ticks
17
^i^>>^
'//^^y^y^^^
0.02 mm
- A^Wo
Fig. 10. Ornithodoros capensis, larva and nymph.
18 Brigham Young University Science Bulletin
days afterwards. Ist instar nymphs molted to (behind capituluin) in females, nymphs, and
2nd instar nymphs in 13-17 days without a blood larvae. C-'apituhim at anterior margin of body.
meal. These 2nd instar nymphs, when given a Porous areas present in female on dorsal side of
blood meal within 9 days after molting, molted basis capituli. Eyes present or absent; when
again to 3rd instar nymphs 14-18 days after the present, laterally on the scutum. Palpal articles
blood meal. There were at least three instars in not free; 4th article much reduced, inserted ven-
the nyinphal stage under laboratory conditions trallv on 3rd ;irticle. Spiracular plate present,
at approximately 25 C, but some 3rd instar posterior to coxae IV.
nymphs developed into 4th instar nymphs. De- „ „ i / i i_ n i-rnc
-., ' r 1 IT ' I 1 11 1 .11 I, Type Genus: /xor/e.s Latreillc, 1795.
tails of the life cvcle shall be reported elsewhere
at the conclusion of current rearing and life- Genus Ambhjomina C. L. Koch, 1844
^ ' Generally ornate. Palpi long; article II at
Disease Relationship: least twice as long as wide. Eyes and festoons
Unknown. present. Basis capituli variable in form; usually
roughly triangular or rectangular dorsally.
Family Ixodidae Murray, 1877 Adanal shield absent in male. Spiracular plate
,.,,.,„ ^ , ,. I . rousihlv triangular or comma-shaped.
Scutate hard ticks. Sexual dimorphism ^ . o i
marked; well-sclerotized scutum covering the Type Species: Amhhjomma cajennense (Fabri-
dorsal surface almost entirely in males, partially cius, 1787).
Key to the Males of the Genus Amhhjoinina
1. Hypostome with denticles of inner file almost equal with others; coxae II, III, IV with
subecjual external .spurs; on tortoises (Fig. 12) geoeimjdae
Hypostome with denticles of inner file much smaller than others; coxa IV with external spur much longer than those of coxae II and III; on larger wild and domestic mam- mals ( Fig. 17) testudinarium
Key to the Females of the Genus Amhhjomma
1. Hypostome with denticles of inner file much smaller than others; coxa IV with external
spur slightly longer than those of coxae II and III; on larger wild and domestic
mammals (Fig. 16) testudinarium
Hypostome with denticles of inner file almost equal with others; coxae II, III, IV with
subecjual external spurs 2
2. Coxa I with two spurs; body with thick setae; dorsal foveae unusually large; on tortoises
(Fig. 11) geoemijdae
Coxa I with single small spur; body without thick setae; dorsal foveae inconspicuous; on
.sea snakes (Fig. 15) nitidum
Amhhjomma geoemijdae (Cantor) and noted morphological characters, but he did
(Fig. 11-14) not mention the relationship betvveen the two,
, , , ,, ,„,~ r.^r, probably because specimens of "t'OCHii/f/we were
Ixodes oeoemyduci. .mor. 1847:608. \^^^ ,^^^,(.^^^^^ ^^^ l^^^ Later Kohls (1957b)
Amhhjomma maknjunum Neumann, 1908:14-16, discussed both species 'and showed mahnjamim
26, Fig. 9-10; Keegan and Toshioka, 1957:8, (^ ^e a svnonvm of oeocmiidac. PI. 4, 5; Kawashima, 1963:103; Asanuma, Specimens reported here were ail taken from
1965a: 107. tortoises on Kyushu and in the Hyukyu Islands.
D,o^.,oo,^», One specimen, a nvmph found on a human, was
iscussiON: 1,1 r ■ 1 .1-
„, . . , 1, , 1 , 1 ^, probably a stray from a tortoise examined the
I his tick, originally described by (>antor m '. ^ i/
IS47, was collected from the neck of Cxeocmifda ' '■^'
spinosa in Sumatra. Keegan and Toshioka Diagnosis:
(1957) collected males, females, and nymphs This tortoise tick is easily distinguished from
from tortoises on Kyushu and in the Ryiikyu nitidum and testudinarium by the combination
Islands. Anastos (1950) reviewed the synonymy of characters used in the key. The female of
and distribution of mahnianum and '^eoemrjdae this species is especially distinctive in that the
Biological Series, Vol. 15, No. 1
Ticks
19
Fig. 11. Amblyornma geoemydae, female.
20
Bhigham Young University Science Bulletin
Fig. 12. Ambh/omina geoemijdcw, inal
Biological Series, \'ol. 1.5, No. 1
Ticks
21
Tti. ■TrLc^^iu,
Fig. 1.3. Amblyomma geocmt/dae, nymph.
Brigham Young University Science Bulletin
'?n. Tm^-tiA^
0.2 mm
Fic. 14. Aml>li/omma gcocvujdae , larva.
BioLor.icAi, Series, Vol. 15, No. 1
Ticks
23
clor.suin hii.s a pair of large, adjacent foveae on the post.scutal area; the dorsal surface has thick, short setae; and the basis capituh has large porous areas.
DisTiuBUTiox .\Ni) Hosts:
Anastos (1950) stated that the species was known only from tortoises collected in Singa- pore, Malaya, Simiatra, and the Philippines. Kohls (1957b) also reported this species from Sarawak ex Varanus salvator, Ctjlemtjs dentata (as C. (Ihor), Ueosemijs spinosa (as Geoemijda spinosa), Te.studo cmtjs, and "stomach of fish," Pontius sealei.
|
Amblyomma geoemydae . ," ! |
|
|
^ Collected and examined * // |
|
|
O Collection records ^»^ 0 U literature Jv^ %^f\ |
|
|
CP Combination of above ^ // \ '^'^h^ |
|
|
ty / J jl^T"**^ " |
|
|
^ ** / o \ r'"-^^^'"^\2f* |
|
|
Okinawa Hfc // C^-y } (] Ky"^^ |
|
|
" ^ / '^^ ' |
|
|
Inomote /-v • <^ y^ ^7 \ |
|
|
1=-^^°°"^'^ ^AXt |
|
|
r-0 y ) ^ |
|
|
-^^^^^^^ |
|
|
^j^Xi^sV^' i \ y |
. |
|
t.^'fi^^'l/ |
A ^ |
|
^'\a' } |
/~i 1 |
|
i>-vTanegashima |
'"^^ |
|
' •' |
o 100 '->-\jr |
Map .5. Known distribution of Amhli/otnrmi ^cocmt/dtic.
Biology:
One of us (Toshioka) reared to adults, nymphs collected from Clemmys japonica in Tanegashima, Kagoshima Prefecture, 19 July 1955. In the laboratory the nymphs attached to the wet skin of the tortoise, Geoclemys reevesii, and molted on 1.3 September 1955.
Xadchatram (1960) used Testudo emij.s, Ctiora am})ou\en.sis (a box tortoise), and a guinea pig as hosts for his life-cycle studies. Testudo emtjs was the most suitable host for larvae. nvTnphs, and adults; the box tortoise was
a little less suitable host, and the guinea pig was an unsatisfactory host.
Laboratory life cycle of Amblyomma geoemydae (reared on Testudo emys) (from Nadchatram, 1960)
Stage or Activity Days
Female oviposited after 5
Female continued oviposition for 21
Larvae hatched out after 20-32
Larvae were fed after 24
Larvae fed for 8-20
Engorged larvae quiescent for 3-5
Nymphs emerged after 10-14
Nymphs were fed after 20
Nymphs fed for 12-28
Engorged nymphs quiescent for 5-10
Adults emerged after 25-30
Adults were fed after 14
Adults fed for 18-48
Number of days required to complete life cycle 185-271
Disease Relationship: Unknown.
Amblyomma nitidum Hirst and Hirst
(Fig. 15)
Amblyomma nitidum Hirst and Hirst, 1910: 304- .305.
Discussion:
The original description of this tick was based on specimens collected from a sea snake in the Solomon Islands. Sharif (1928) stated that there were specimens of all stages in the Indian Museum from Port Blair, Andaman Is- lands. Warburton (1932) described Amblyom- ma laticaudae as a new species taken from a sea snake Laticauda colubrina in Singapore. Kohls ( 1957b ) reported on Amblyomma sp. from L. colubrina from Singapore, and Audy, Nadcha- tram, and Lim (1960) also reported A. nitidum from the same host and locality. Audy et al. (1960) stated that Kohls later compared these specimens and listed laticaudae as a synonym of nitidum.
Two females collected from Laticauda semi- fasciatus on Ishigaki ( Is. ) were the first reported specimens of A. nitidum in the Ryukyu Islands.
Diagnosis:
The female of this sea snake tick is distinct from the other two Ambh/omma spp. included in this publication in that the coxae have very small external spurs; the lateral margin of the basis capituli is convex at the middle when viewed from the ventral side; the dorsum does
24
Bricham Young University Science Bulletin
Fig. 15. Amhhiommn nitUhim. female.
Biological Series, Vol. 15, No. 1
Ticks
25
not have marginal groo\'e.s; and the dorsal foveae are inconspicuou.s.
Distribution and Hosts:
The knowai distribution includes: Solomon Islands, ex sea snake (Hirst and Hirst, 1910); Andaman Islands, host unmentioned (Sharif, 1928); Singapore ex Laticauda coluhrina (War- burton, 1932, as A. laticimdae: Audy et al., 1960); and the first record from the Ryukyu Islands reported herein.
M.\p 6. Known distribution of Ambhjonima nitiduv
Biology:
It is of interest that this nocturnal sea snake spends the daytime resting, often in tightly-packed groups, in mangrove tree- holes or in rock crevices out of the water. Furthemiore, the genus Laticauda appears to have evolved from an elapid stem much later than the other sea snakes. This would explain not onlv the infestation by Am- hh/onuna, but also the infestation of the air- sacs by larvae of a peculiar trombiculid mite, Vatacarus ipoides Southcott ... It will be interesting to discover how A. nitidum is adapted to parasitizing this sea snake (Audyetal., I960).
Disease Relationship: Unknown.
Amblyomma testudinarium C. L. Koch (Fig. 16-19)
Amhlijomma testudinarium C. L. Koch, 1844: 226; Kishida, 1922a: 850-851, 856; Robinson, 1926:257; Sugimoto, 1937a: 317-323, 1937b: 612-613, PI. 4; Nakamura and Yajima, 1937: 174-175, Fig. 1-9 in PI. \IV; Itagaki, Noda, and Yamaguchi, 1944:1-97, PI. 17, 22, 1959: 1-118, PI. 17, 22; Keegan and Toshioka, 1957: 9, PL 6, 7; Kawashima, Kamo, and Miyazaki, 1960:77-80; Tanaka et al., 1960:69-70, Fig. 1-3; Nagahana and Matsuo, 1962:119-120; Asanuma, 1965a: 106-107, Fig. 7.3, 1965b:400, Fig. 226.
Ambhjomma ijajimai Kishida, 1935:137-138, Fig. 1-4; Nakamura and Yajima, 1937:176, Fig. 7, 8 in PI. I, Fig. 5-10 in PI. XV; Itagaki, Noda, and Yamaguchi, 1944:1-97, PI. 18, 22, 1959:1-118, PI. 18,22.
Discussion:
Anastos (1950) and Kohls (1950) have re- viewed the extensive synonymy of this widely distributed species. The first report of this tick from Japan (Kishida, 1922a) was based on a specimen collected from a pig in Tokyo. Sugi- moto ( 1937a, 19.37b ) reported two collections of larvae from the frog, Microhtjh fissipes, and from a green frog (sp?) on Ishigaki, an island in the Ryukyu archipelago. These two lots were not collected by Sugimoto but submitted to him by the collector. Considering the host-parasite relationship of this tick, it is very questionable that A. testudinarium was parasitic on a frog.
In his description of Ambhjomma ijajimai Kishida (19.35) included illustrations of venter, scutum, spiracular plate, and the 1st tarsus of the holotype female. He gave only diagnostic characters to separate it from A. testudinarium, and a precise description of the holotype was given later by Nakamura and Yajima (1937). The holotype specimen labeled as "1291 l.VII. 1935. Taito-cho, Hinan-ku (Formosa), Pasicau, from mammae of water buffalo, Yajima Am- hh/omma ijajimai Kishida" was examined by Kohls, and it was his opinion that A. ijajimai is is conspecific with A. testudinarium Koch. A. Ijajimai was also reported by Itagaki et al. ( 1944, 1959), and all of these collections are from water buffalo, Buhalus hubalis, on Formosa.
Diagnosis:
This large tick is easily distinguished from nitidum and geoemijdae in that the denticles of
26
Brigham Young University Science Bulletin
Fig. 16. Amhli/omma testudmiirium. female.
BrOLOGiCAL Series, \'ol. 15, No. 1
Ticks
27
Fic. 17. Amhlyommii testudinarium, male.
28
Brkjham Youno University Science Bulletin
^^r-
^^
'M.. ^ti»»^
Fic. 18. Atnhlyomma testudinarium . nymph.
Bk)i,oc.icai. Sehies, Vol. 15, No. 1
Ticks
29
KiG. 19. Amhlyoinrna tcstudinarium, larva.
30
Bhigham Young University Science Bulletin
the inner file are much smaller than those of the other files; the external spur of coxa I is longer than the internal spur; the spur of coxa IV is longer than those of coxa II and III which arc short and blunt in the male, broad and ridgelikc in the female; tarsi III and IV have two distinct subapical ventral spurs; the dorsum has a pair of small foveae which arc well separated in the female.
Distribution and Hosts:
Kohls ( 1957b) has listed the distribution and hosts of this species. It is widely distributed in the Far East and is known to occur in the fol- lowing areas: Indi;i, Bunna, Ceylon, Indonesia, Borneo, Malaya, Indochina, Formosa, the Philip- pines, and Japan. Hosts recorded by Sugimoto ( 1937a), Nakamura and Yajima (1937), and also by Kohls ( 1957b ) are usually larger animals such as water buffalo, horse, cow, wild boar, wild pig. goat, dog, tiger, rhinoceros, deer, tapir, and human.
Amblvomma testudinanum
^ Collected and examined
O Collectpon records (fom
literature ^ Coinljination of above
Map 7. Known distribution of Amhhjommii tcsttiili- imriiim.
BlOLOCY:
According to Anastos (1950) all stages in the life cycle are known, and this tick occurs on
a wide variety of wild as well as domestic ani- mals. One of the unengorged females collected by 4()6th laboratory personnel from Amami Oshima was placed on a rabbit's ear in the hiboratory on 6 November 1967, fed for a week, and increased in size to 23 mm in length and 21 mm in width. After engorgement this tick died on 26 November 1967 without laying eggs.
Disease Relationship:
"Krijgsman and Ponto (1932) stated that this species transmits piroplasmosis and anaplas- mosis, and Sharif (19.3S) considered this species as a possible disease vector" (Anastos, 1950). In Japan, however, its disease transmission capa- bility is not known. Japanese workers have re- ported that it attacks humans.
Genus Boophilus Curtice, 1891
Inornate. Palpi short, compressed, ridged dorsallv and laterally. Basis capituli hexagonal dorsally. Eyes present. Festoons absent. The males with adanal and accessory shields. Anal groove obsolete in the female, faintly indicated in the male. Caudal process present or absent in the male. Spiracular plate round or oval in both sexes. Coxa I bifid.
Type Species: Boophihis annulatus (Say, 1821).
Boophilus microplus (Canestrini) (Fig. 20-23)
HaemapIiysaUs micropla Canestrini, 1887:104- 105, 110, PI. 9, Fig. 3, 3a-d, 5, 5a-b.
Rhipiceplwhis atinulatus cauclatus Neumann, 1897:413, Fig. 42.
Uwboophilm austraUs Fuller, 1899:389-392; Ki- shida, 1939a:541-544.
Boophihts annulatus cauclatus: Tokishige, 1911;
Ogura, 19.36:75, Fig. 1-6 in Pi. IV. Boopliilus sp.: Nuttall and Warburton, 1915:
433. BoopfiUus cauclatus: Minniiig, 1934:25-27; Ki-
shida, 1927:985; Itagaki, Noda, and Yama-
guchi, 1944:1-97, Pi. 15, 19, 1959:1-118, PI.
15, 19; Kitaoka and Yajima, 1958a: 135-147,
1958b: 148-162, 1958c: 179-188. Ihohoophilus sinensis Minning, 1934:25-27;
Schulze, 19.3.5:2.^; Kishida, 1939a: .541-544;
Nakamura and Yajima, 1937:168-175, PI.
XIII. Vroboophilus shariji Minning, 19.34:1-48; Kishi- da, 1939a:541-.544. Palpohoc^philus minningi Kishida, 1936:140-142.
Fig. 1-9; Nakamura and Yajima, 1937:168-
175, PI. XI.
Biological Series, Vol. 15, No. 1
Ticks
31
Margawpits aiitiulatus dtistralis: Sugimoto, 1936c: 582-583, 1937b:599-601.
Margaropus annulatus caudatus: Sugimoto, 1937h:6()l-603.
Palpohoophihis hrachijuris Kishida, 1939a:538- 552, PI. I, II; Shigemori, Aso, and Yajima, 1953:290-293.
LJroboopJiilus cdtiihiltis: Ki.shida, 1939a: 54 1-544; Nakaimira and Yajima, 1937:168-175, PI. XII, 1942b:34-39, Fig. 1-3 in PI. II; Shigemori, .\,so. and Yajima, 1953:290-293.
Boophilus microj)his: Keegan and To.shioka, 1957:9-10, PI. 8, 9; Kitaoka, 1961a:85-95, 1961b:96-104, 1961c: 105-112, 1967:18-21; Ki- taoka and Yajima, 1961a:41-52; Kitaoka and Morii, 1963:.32-35, 1967b: 126-129.
Discussion:
Thi.s tick occurs in Japan, Korea, and the Ryukyu Islands and is apparently the only spe- cies of the genus Boophilus found in these areas. It is known under a great variety of names in the Japanese literature. This is partially due to the fact that most Japanese workers have fol- lowed the classification of Minning ( 1934, 1935, 1936). In this paper we have accepted the opinions of Anastos (1950) and Hoogstraal ( 1956) in the classification of species of Boophi- lus. These workers reject the subdivisions of the genus and additional species designations pro- posed bv Minning and are of the opinion that only a careful study of the vast Boophihis ma- terials available will resolve current inconsisten- cies. Thev list three species: B. decoloratus (C. L. Koch, 1844), B. annulatus (Say, 1821), and B. microj>his (Canestrini, 1887) to which must be added B. kohlsi Hoogstraal and Kaiser, 1960. Of these only microphis has been shown to occur in Eastern Asia. Sugimoto ( 19.36c, 1937b) reported Margaropus annulatus australis and M. a. caudatus ex cattle, horse, and dog from Kumamoto, Oita, and Kagoshima prefec- tures, and also from four islands of the Ryukyus, hut in the light of our collection these may have been misidentified. Specimens also have been erroneously identified as Boophilus annulatus caudatus l)y Tokishige (1911) and Ogura (19.36), both of which were cited bv Sugimoto (19.37b). Kishida (1929) also reported B.^/nni/- latus caudatus from Okinawa and Tanegashima (Is.). The holotype of B. (Palpohoophilus) minningi Kishida, 19.37. as well as other speci- mens under these various names, were examined bv Keegan and Toshioka, and none of them dif- fered significantly from microplus.
Diagnosis:
Anastos (1950) and Arthur (1960) .stated
there are only three valid species of Boophilus in the world, but B. kohlsi Hoogstraal and Kaiser, 1960 should be included as the fourth species. Although these species are similar in general appearance, B. microplus may be distinguished by the following combination of characteristics: the male has a distinct caudal appendage ( this serves to discriminate niicro])lus, decoloratus, and kohlsi from annulatus), and the dentition of the hypostome is 4/4 in both sexes {micoplus and kohlsi), while 3/3 or rarely 3.5/3.5 in decoloratus: in the female, the inner margin of the palpal basal segment is concave, while this segment has an inner bristle-bearing protuber- ance in decoloratus and kohlsi.
Hoogstraal (19.56) states that microplus males have adanal shields in which the inner margin does not reach the body margin, but in Japanese specimens this spurlike projection often goes beyond the posterior body margin, and the tips may be seen from the dorsal side.
Distribution and Hosts:
Anastos (1950) and Hoogstraal (1956) give detailed infomiation concerning distribution and hosts of B. microjilus. It is known to occur in Central America, South America, Australia, the Orient, southern Florida, and Africa. In the Orient it is known from Fonnosa, Indonesia, the Philippines, New Guinea, Borneo, Burma, India, Assam, and small islands of Southeast Asia. Oc- casionally it has been found on domestic chick- ens and in Africa has been collected from a lion. Other hosts recorded are cattle, horses, buffalo, sheep, deer, and dogs. Kishida ( 1927 ) adds the pheasant as a host of this species.
Biology :
Boophilus microplus is a one-host tick; the larvae attach to the host and remain thereon until maturity. Molting and mating occur on the host. "Engorged females leave the host from 35 to 149 days after having attached as larvae and there may be from two to three generations a year in South Africa" (Lounsbury, 1905, cited by Hoogstraal, 1956). "Wilson (1946) observed no seasonal periodicity of adults in Nyasaland. He found larvae with nymphs and adults on cattle only once. Nymphs and adults were usually found together. . ." (Hoogstraal. 1956).
Kitaoka and Yajima (19.58c) calculated the duration of each stage on bulls which were ex- perimentally infested with batches of larvae throughout the year. "Fifty percent of the larvae and nymphs molted 6.6 and 14.0-15.6 days after infestation, respectively. Fifty percent of engorged female ticks dropped in 22.1-24.2 days after infestation. Adult males migrated to search
32
BmcHAM Young University Science Bulletin
Jll' . ^■/i*.^<.^>'cCa^
Fio. 20. Boophilus microplus, female.
Biological Series, Vol. 15, No. 1
TlCK.S
33
Fig. 21. Bnophiluf microplus, male.
34
Brigham Young Univebsitv Science Bulletin
1 0 mm
'hi. "TtuaoJ^
Fio. 22. Boophilus micropltts, n\inph.
Biological Series, \'ol. 15, No. 1
Ticks
35
0,5 m m
•%.;?t*»«^
Fic. 23. Boophilus microplus, larva.
36
Bhicham Young University Science Bulletin
for females after ecdysis and feeding for 2-3 days. The duration of the whole parasitic stage was influenced by seasonal temperature to a nuicli less degree, and the longest duration varied from 27-32 days in all seasons in Tokyo." A series of excellent physiological studies on this species (as B. caudatus and /nicro/j/i/.v) has been worketl out bv Kitaoka and Yajima ( 1958a,
1958b, 1958c) and Kitaoka (1961a, 1961b, 1961c).
Rabbits were used as host animals for a rearing experiment at the 4()6th Medical Labora- tory. An engorged female ( collected from a black calf from Ishigaki [Is.], 8 February 1967, Tipton, 67-R-0011)Maid 2,318 eggs from 14 February to 3 March 1967.
Laboratory life cycle of Boophilis microphts ( reared on ears of rabbits ) Piuise Date Remarks
Female oviposited on Larvae hatched on Larvae placed on host Earliest nymph emerged on Earliest adult emerged on Engorged adults dropped on
14 Feb.-3 Mar.
17 Mar.-31 Mar.
4 Apr.
13 Apr.
21 Apr.
1 May-5 May
Rate of hatch 98.4%
93 females, 78 males through 5 days
Mai* 8. Known distrihntioii of hoophiliis miiroi>lus.
DisE.\sE Relationship:
This species is one of the most important pests of cattle, and a vector of organisms patho- genic to domestic animals in the Americas, Asia, and Australia. These pathogens are Aimplasma marginiile, Babesia hi<^cmina, B. berbera, B. ovis.
Borrelia spp., Coxiella burnetii, Haenuitoxenus velifertis, and NuttaUa equi. Babesia bigemina is the causal agent of the disease known under the various names of Texas cattle fever, red water fever, splenic fever, bloody murrain, Mexi- can fever, and tick fever.
Genus Dermacentor C. L. Koch, 1844
(Fig. 24-2.S)
Usually ornate. Palpi short, broad, or mod- erate. Basis capituli rectangular dorsally. Eyes and festoons present. Males with coxa IV much larger than others. Ventral plates or shields ab- sent in males. Coxa I bifid in both sexes. Spi- racular plate ovate or comma-shaped.
Type Species: Dcnmuvntor rcticuhitus (Fabri- cius, 1794).
We did not attempt to determine the species of the genus Dermacentor for the same reason that was given by Keegan and Toshioka ( 1957). Whereas adequate infonnation on other genera has been published bv Japanese workers since 1957, verv little is available tor the genus Der- macentor. Kohls (1967) stated that specific detenninations within the genus will continue to be difficult until more extensive studies on in- traspecific variation in the Far East have been made. We followed his suggestion and included only the collection records derived from the literature in chart fomi in Appendix 2.
At least four species have been reported as occurring from Korea and Japan bv Kishida (1922a, 19.36), Nakamura and Yajima (19,37), Yajima (1942), and Itagaki, Noda, and Yaina- guchi ( 1944, 1959). They include D. auratus. D. coreus, D. recticulatus. and D. variegatus. Arthur
BiHLOcicAi. Skiues, Vol. 15, No. 1
Ticks
37
K Tf'^J'^'iAAAAa^
Fin. 24. Dermaccntor sp., female.
38
Brigham Young University Science Bulletin
X'^^n^-^j-aA^
Fig. 25. Dcrmtirentnr sp., male.
Biological Series, Vol. 15, No. 1
Ticks
39
9>Uy^^»AsJtA.
Fig. 26. Dermacentor sp., female.
40
Brigham Young University Science Bulletin
X-^yi-^^yOtji^uy.
Fic. 27. Dennacentor sp.. male.
Biological Series, \'ol. 15, No. 1
Ticks
41
Fig. 28. Dennacentor sp., male.
42 BmciiAM VouNG University Science Bulletin
(I960) synonymized D. varieguttis under D. al- Sexual diniorpliisiii sliglit. Palpi usually conical
hipictus. with 2nd article salient laterally beyond width
The illustrations used in Keet^an and Toshi- of basis capituli (except H. kitaokai). Basis
oka (1957) arc reproduced without change, capituli rectangular. Ventral plates absent in the
They were made from specimens identified and male. Platelike dorsal spur present on trochan-
loaned by Dr. Asanuma (D. auratus and D. ter 1. Spiracular plates usually comma-shaped
coreus) and Dr. Yajima (D. reticulatus). in males, round or oval in females. Posterior
margin of coxa I never bifid.
Genus Haemavhii sails C. L. Koch, 1844 _ „ ,, i 7. ■ r^ 1
' •' Type Species: Haenmplnjsam conctnna C. L.
Inornate. Eyes absent. Festoons present. Koch, 1844.
Key to the Males of the Genus HaemapJtijsaJis
1. Palpal segment III with dorsal retrograde spur 2
Palpal segment III without dorsal retrograde spur 5
2. Palpal segment III with dorsal spur at middle (Fig. 55 and 66) 3
Palpal segment III with dorsal .spur not at middle (Fig. 38 and 77) 4
3. Palpal segment II with lobelike, anterointemal projection dorsally; ventroposterior mar-
gin fomiing an angle at middle; lateral groove absent (Fig. 55) . hijstncis
Palpal segment II without such projection as above; ventroposterior margin slightly
convex; lateral groove present (Fig. 66) longicornis
4. Coxa IV with two long .spurs; palpal segment III with a dorsal spur extemolaterally
(Fig. 38) - - -- cornigera group
Coxa IV with a short spur; palpal segment III with a dorsal spur on internal margin
(Fig. 77) - wellingtoni
5. Palpal segment II long, clavate, not salient laterally; suture between II and III not clear;
spiracular plate about twice as long as wide (Fig. 62) kitaokai
Palpal segment II slightly salient laterally with round contour posterolaterally, scarcely
beyond basis capituli; suture between II and III well defined (Fig. 47) formosensis
Palpal segment II dLstinctly salient laterally beyond basis capituli; suture between II and
111 well defined 6
6. Palpal segment III clawlike, strongly curved inward at tip, overlapping when closed
(Fig. 34) '^concinna
Palpal segment III otherwise 7
7. Coxa IV with long internal spur 8
Coxa IV with short internal spur 9
8. Color light yellow; length 2.02-2.97 (avg. 2.49 mm), breadth 1.37-1.63 (avg. 1.51 mm);
spur on coxa IV long, sharply pointed, usually straight, approximately Y3 the length
of coxa (Fig. 43) jlava
Color yellowi.sh-l)rown; length 2.89-3.70 (avg. 3.30 mm), breadth 1.43-2.14 (avg. 1.80 mm); spur on coxa IV stout, wide at base, usually curved, less than half the length of coxa (Fig. 70) vwgaspinosa
9. Coxa I without spur; on Ryukyu black rabbit (Fig. 74) pcntaJagi
Coxa I with internal spur 10
10. Posterior half of palpal segment II prominently flared, fomitng bell-shape; junction of segments II and III definitely notched, tarsi with strongly dorsal hump; on dog
(Fig. .'30) caiiipanulata
Posterior half of palpal segment II otherwise; junction of segments II and III without
definite notch; tarsi tapering (Fig. 59) japonica
^(11. iuitmna is not Imiuded in llic almve key.)
Key to the Females of the Genus }IaemaphysaJi,s
1. Palpal .segment III with dorsal retrograde spur 2
Palpal segment III without dorsal retrograde spur 4
2. Palpal segment II strongly salient extemolaterally; palpal segment III with long \entral
Biological Series, Vol. 15, No. 1 Ticks 43
spur; hvpostome 4 4 (Fig. 37) , .. cornigera group
Palpal segment II moderately salient extemolaterally; palpal segment III with moderate
ventral spur; hypostome 5/5 (Fig. 54, 65) 3
3. Palpal segment II with an anterointemal lobelike projection dorsally (Fig. 54) htjstricis
Palpal segment II without such a projection ( Fig. 65 ) longicornis
4. Palpal segment II long, parallel-sided, not salient laterally; comua absent; lateral
grooves absent; hypostome 3/3 (Fig. 61) kitaokai
Palpal segment not parallel-sided, at least slightly salient laterally; coniua present; lat- eral grooves present, hypostome 4/4 or more 5
5. Palpal segment II slightly salient laterally with round contour posteroextemally, sairce-
ly beyond basis capituli; comua present; lateral grooves present; hypostome 4/4 in
anterior half, 5/5 in posterior half (Fig. 46) formosensis
Palpal segment II salient laterally, distincdy beyond basis capituli; comua present;
lateral grooves present; hypostome more than 4/4 _ 6
6. Posterior half of palpal segment II strongly salient or flared extemolaterally forming
boot-shape (Fig. 41) or bell-shaped (Fig. 29) _.__ _ .. 7
Palpal segment II salient laterally but not fomiing such shape as above 8
7. Palpal segment II protruding extemolaterally at base at right angle to the axis, boot-
shaped; comua very slight; tarsi tapering (Fig. 41) _ doenitzi
Posterior half of palpal segment II prominently flared, bell-shaped; external profile with a definite notch at junction of palpal segments II and III; comua short but well defined; tarsi humped (Fig. 29) campanulata
8. Coxa I without spur; external profile of palpal segment II with a short sahence, point-
ed, slightly directed extemolaterally at the posteroexternal juncture; on Ryukyu black
rabbit ( Fig. 73) . pentalagi
Coxa I with a short spur; external profile of palpal segment II nearly straight or slight- ly bulged externally at the posteroexternal juncture 9
9. Coxae I-III with short, blunt, subequal spurs; coxa IV with a spur longer than others 10
Coxa I with a spur distincdy longer than others 11
LO. Color light yellow; coxa IV with a spur slightly longer than others; length 1.89-1.92
(avg. 1.90 mm), breadth 1.19-1.28 (avg. 1.2.3 mm)"(Fig. 42) _. flava
Color brownish; coxa IV with a strong, triangular spur distinctly longer than others; length 2.99-3.48 (avg. 3.31 mm), breadth 1.88-2.21 (avg. 2.03 mm) (Fig. 69)
inegaspinosa
11. External profile of palpal segment III longer than that of II; spurs on coxae II, III, IV
short but distinctly extend beyond posterior coxal margins (Fig. 58) japonicn
Extemal profile of palpal segment III subequal to that of II; spurs on coxae II, III
blunt, scarcely extend beyond posterior coxal margins (Fig. 33) . ^ concinna
Haemaphysalis campanulata Warburton based on numerous specimens taken from Chi-
(Fig. 29-32) nese dogs in Mongolia by M. L. Hearn. He
Haeinapht/salis campamdaia Warburton 1908: '''l^o ^^'^^ed that the type specimens agreed in
508-514, Fig. 5, 6; Nuttall and Warburton, "^^^^"^^ ^^* *<^ specimens in the British Mu-
1915-491-492 Fig 431-4.32- Kishida 19'?2a- ^^"'" named H. flava by Neumann. Warburton
852; Nakamura and Yajima,' 19.37:155 PI. VI, ( ^908) did not mention the collection data of
1942b:37 Fig. 8 in PI II Fig. 1-5 in PI III- ^^^ ''^"«'''- ^^^ ^^^ '^^ presumably rs tube No.
A.sanuma, 1942:17-26, 1947b :972, Fig. 2768, 137 (see Nuttall and Warburton, 1915, p. 408),
1956a: 111-112, 1956b:400, Fig. 225; Itagaki, ^^^ich he detemiined to have contamed flava.
Noda, and Yamaguchi, 1944:1-97, PI. 4, 1959: campanulata, and japonica. Subsequendy, Nut-
1-118, PI. 4; Shigemori, Aso, and Yajima, J'^" '''f^ Warburton (1915) stated that War-
1953:290-293; Keegan and Toshioka, 1957:12, burtons types of japonica (from a tube labeled
p] ]7 ^8 "No. 173 H. flava )'■ were found on Nemor-
haedus crispus, now known as the Japanese
Discussion: serow, Capricorni.s crispus, at Hondo, Japan, by
Warburton's original description (1908) was the Duke of Bedford's collectors (see discussion
'In Nullall and Warburton, 1915, p. +08, //. Ilava. this number is 1)7.
■44
BlUOHAM ^OUNG UnIVEHSITY SCIENCE BULLETIN
F. T5UT5U.MI
Fig. 29. Ilucmaphijsiilui carnpdnuUita , ft-niiilc.
BlOLOOICAI. ShHIEN. \'()1,. 15, No, 1
Ticks
45
FTSUTSUhl
Fig. 30. Haemaphysalis campamihita, male.
46
HiticHAM Young UNrvF.RSiTY Science Bulletin
F"ic.. 31. Haemaphysulis cainpumilatn. nymph.
Biological Series, Vol. 15, No. 1
Ticks
47
Fic. 32. Haemaphysalis campamdata, larva.
48
BniGHAM Young University Science Bulletin i
of //. japonica). If it is true that all of the specimens in tube N. 173 (No. 137?) were col- lected from the same host and locality, this col- lection data of japonica might apply to that of campanulata. Tlicrefore, Warburton's reference to the specimens of campanuhtu, whicli had been mixed with jlata in the tube, might well be the first record of this species from Japan.
Nuttall and Warburton (1915) examined Warburton's Mongolian specimens and reported additional samples from Japan. They were col- lected from dogs in Tokyo and Yokohama and from a house rat in Tokyo.
Diagnosis:
Thi.s common dog tick is distinct from other haeinaphysalids in that palpal article II is strong- ly salient laterally; the posterior margin is flared upward to become "bell-shaped," hence the name "campanulata-" the junction of article II and III is deeply notched; the tarsi are humped. Disthibution' and Hosts:
This species is known from China, Mongolia. Manchuria, Japan, and Korea. Hosts include cows, horses, dogs, rats, and man. One lot sub- mitted to us (collection, 41-J-(X)2; see Appendix 2) contained .33 females, 11 males, and 4 nymphs, but it seems doubtful that so great a number of ticks were taken from a human host. There are no records of this species from the Ryukyu Islands.
Map 9. Known jxniiiltita.
listribution of llucinaplu/sc:li.\
Biology: stages in the life cycle may be found on the
Both immature and adult ticks are very com- ground around dog kennels. Asanuma (1942b)
mon on Japanese dogs, even in urban areas. All has provided an excellent account of the life
Generation
and Stage
Ph;i
Laboratory life cycle of H ae ma j)]u/ sails campanulata (reared on rabbits)
Period in Days
Remarks
|
F, |
Adult |
Feeding |
9-14 |
Females were placed on host with males on 29 Nov. 1967. |
|
F, |
Adult |
Postparasitic ( Preoviposition ) |
4-7 |
|
|
F, |
Adult |
Oviposition |
12-24 |
Egg number: 826-1,749 |
|
F.. |
Egg |
Incubation |
25-29 |
10,126 F:; larvae hatched from |
|
F. |
Larva |
Feeding |
2-10 |
10.693 eggs. Lar\ae were placed on host 29 Jan. lf)6.8. |
|
F, |
Larva |
Postpartsitic |
10-13 |
|
|
F., |
Nyiuph |
( Premolting ) Feeding |
3-7 |
|
|
F, |
Nymph |
i'ostparasitic (Premolting) |
11-27 |
Ticks were reared in facilities at the 406th Medical L;iboratorv in whicli temperature was main- tained at approximately 25 C and relative humidity fluctuated from 10-70%.
Biological Series, Vol. 15, No. 1
Ticks
49
cvcle of this species.
Rearing was attempted at the 406th Medical Lahoraton'. A single female was obtained from a dog in Chiha Prefecture, 12 June 1967, and 12 females of the F, progeny were fed on rahbit ears.
Disii.^sK Relationship: Unknown.
Haema))In/salis concinna C. L. Koch (Fig. 3:3-36)
Ihiemaphiisalis concinna C. L. Koch, 1844:237- 238; Nuttall and Warburton, 1915: 452-458, Fig. 387-393; Kishida, 1922a:852, 1936:142; Ogura and Takada, 1927:202-204, PI. XII; Yajima. 1942:499-510, Fig. 4; Itagaki, Noda, and Yamaguchi, 1944:1-97, PI. 7, 1959:1-118, PI. 7; Keegan and Toshioka, 1957:13; Asa- numa, 1965a:lll-112.
lliwmaphijsalis Jiirudo L. Koch, 1877:786; Neu- mann, 1897:341; Keegan and Toshioka, 1957: ,30.
lUicmaphijsaUs concinna var. kochi Neumann, 1905:239.
llaemaphijsalis concinna kochi: Newmann, 1911: 111.
Discussion:
Neumann (1897) at first referred his speci- mens from Japan to //. liirudo Koch, then subse- quently (1905) renamed them H. concinna var. kochi, realizing that the type had been lost. The specimens were three females collected by Hil- gendorf ( Berlin Museum ) and two females from a dog at Saga collected by Yamaguchi. Neu- mann (1911) raised this variety to subspecies, but later Nuttall and Warburton (1915), having examined Neumann's specimens in the Berlin Museum, svnonvmized the subspecies under concinna. Koch's original description was not available, but judging from the French quota- tion by Neumann (1911)," it is very adequate for detennining the species. As cited by Keegan and Toshioka' (1957), Nuttall and Warburton (1915) condemned the name Jiirudo. Conse- quentlv. Neumann's hirudo (1897) might be the first reliable record of concinna from Japan, provided we follow Nuttall and Warburton (1915).
Ogura and Takada (1927) reported H. con- cinna on cattle and horses at Tokachi, Kushiro, Nemuro, and Kitami, ;dl on Hokkaido. Kishida (1936) reported collection of this species by M. Akasawa in 1928 from North Korea. Itagaki et al. (1944, 1959) also recorded a collection
from a horse in Hamgyong-pukto, Korea. Ya- jima ( 1942 ) reported it from horses in Aomori Prefecture. During the preparation of this paper, one female and three males loaned by Dr. Kita- oka were available for study.
Pomerantzev ( 1950 ) , who gave a detailed discussion of the morphology, biology, and dis- tribution of the species, states that "it possibly covers all of eastern and southeastern Asia (Japan, Kokhinkina)."
Diagnosis:
The male of this species is very distinct in that the tips of palpal articles III are curved inward to form "pincers" which overlap when closed. Tarsus W is tapered and without a dorsal protuberance. In the female the palpal profile is very similar to flava and japonica, but may be distinguished from jlava in that the spur on coxa I is longer than the others, and from japonica in that the external profile of palpal article III is subequal to II, palpal article II has 3-4 infraintemal setae, and spurs on coxae II, III, and IV are wide, blunt and extend scarcely beyond the posterior margin of the coxae.
Distribution and Hosts:
The distribution of this species includes Western Europe, the Balkan peninsula, and the USSR. According to Pomerantzev, adults of this species parasitize large domestic and wild ani- mals, while nymphs and larvae are found on a wide variety of small and large mammals and birds, and occasionally on reptiles. Olsufjev and Petrov (1960) state that larvae and nymphs are parasitic on rodents and small mammals. Both adults and nymphs have been known to attack man.
Biology:
The biology of this three-host tick has been studied by Russian workers, and Pomerantzev ( 1950) has briefly summarized the life cycle under laboratory conditions. The engorged fe- male begins oviposition in 4-16 days and lays more than 1,000 eggs over a .30-32 day period. Larvae begin to appear in 16-30 days and feed for 3-6 days. Larvae molt to nymphs in 16-25 days, and nymphs feed 8-16 days or more. The cycle can be completed in the laboratory within 120-160 days. Larvae and nymphs can be starved for more than 13 months. Pomerantzev suggests that the life cycle in nature may take 1.5-2 years. Mel'nikova (1958) observed the life cycle in nature and concluded that it would take 3-4 years for completion. He also found
= Iic
ussim tlors,Tl prosque
tide, palpes tourts,
jM-'ine plus longc que larf^c — I.. 9 mm; la 7 iimi
male inconnu.
50
Brigham Young Univebsity Science Bulletin
Fig. 33. Uacmapht/salis coiwinna, female.
Biological Series, Vol. 15, No. 1
Ticks
51
/Uotio
Fig. 34. Haemaphysalis conciniia, male.
52
Bhigham Young University Science Bulletin
■^TL, 7n.i*i>y^
Fig. 35. Ilacnuiphi/salis conciutui, nymph.
Bioi.ocicAL Series, Vol. 15, No. 1
Ticks
53
0,5 fTim
'TK Ttu^cJu.
0.7 mm
Fic. 36. Haemaphysalis concinna, larva.
.54
BiiicHAM YouNc: University Science Bulletin
Mai- 10. Known cli.stTibulion of Huevuipluisalia con- ciitrui.
that optiinuni conditions for development occur in the middle mountain zone ot oak forest and inter-alpine valleys (400-800 meters above sea level) covered with broadleaf forests where hu- midity is high. According to Cemy and Kra- tochvilova-Kralova (1963), H. concinna in C/ekoslovakia occurs in southern Moravia and southern Slovakia, chiefly on goats. Active adults are found mainly in May and June. He also observed that the entire life cycle took three years.
Disease Relationship:
This species is reported to be a proven vector of the causal agent (rickettsia) of tick typhus in the USSR and expcrimentidly transmits the virus of spring-summer encephalitis. Pavlovskv and Soioviyev ( 196.3) demonstrated the capabil- ity of transmission of the virus by bites of the adults as well as transovarian transmission to its progeny. Ticks collected with a flannel-flag har- bored the agent of tularemia according to Ol- sufjev and Petrov (1960).
Remarks:
Nuttall and Warburton (1915) and Pome-
rantzev (19.50) described the dentition of the female hypostome as 5/5 or 6/6, but in the specimen examined by us the dentition was 4/4.
Haeinaphijsalis sp. {H. cornigera group) (Fig. 37-40)
llaemaplnjsalis cornigera Neumann, 1897:350- 352, Fig. 16-17; Keegan and Toshioka, 1957: 13, PI. 19, 20; Asanuma, 1965a: 11 1-112, 1965b:. 399, Fig. 223.
Haemaphiisalis cornigera var. taiwana Sugimoto, 1936a:"l-ll, PI. I-IV; Sugimoto, 1936d:.336- 346, PI. I.
Haenuiplujsalis ias Kishida, 1936: (? unpub- lished); Nakamura and Yajima, 1937:160, Fig. 10, 11 in PI. VIII, Fig. 1 in PI. IX; Nakatsuji, 1942:295; Yajima, 1950:197-200, Fig. 1-4.
Discussion:
This species is known in the Japanese litera- ture as H. ias Kishida, 19.36. Actually, it seems that Kishida not only did not publish a descrip- tion of ias but did not even publish a reference to this name in 19.36. Apparently he had made a verbal reference to the five male specimens taken from a cow on Miyake ( Is. ) as a new species. In the following year Nakamura and Yajima (1937) published a description of the male of this tick and referred to it as H. ias Kishida, 1936. Yajima (19.50) later described the female and immature stages of the species. This material was also taken from a cow on Miyake (Is.). Yajima also mentioned adult ticks collected from cattle on Hachijo (Is.). Nakatsuji (1942) reported occurrence of ias on a dog on Miyake (Is.), but he did not give details. Kee- gan and Toshioka ( 1957 ) examined the holo- type male and other material loaned bv Dr. Kishida, Dr. Asanuma, and Dr. Yajima. They stated that all of the specimens were taken from cows on Hachijo (Is.) and Miyake (Is.) and that they were unquestionablv cornigera. Since there is still some doubt about the status of H. ias Kishida, 19.36, the Japanese population in this paper is treated tentatively as a part of the cornigera group until there is opportunitA' to further study the problem.
Diagnosis:
This is the only haemaphvsalid species in the Jap;niese fauna in which the male has an excep- tionally large coxa IV with two long spurs. Palpal article II is prominenth" salient laterallv in both sexes. In the male the ventroposterior margin of palpal article II undulates irregularly, giving it the appearance of having two spurs. The dorsal retrograde spur on palpal article III
Biological Series, Vol. 15, No. 1
Ticks
55
F TSUTSUM
Fig. 37. H aemaphysalis sp. (H. comigera group), female.
36
Brigham Young University Science Bulletin
F.TSUTSUMl
Fig. 38. Uaemaphysalis sp. ( H . comigera group) . male.
Biological Sebies, \'ol. 15, No. I
Ticks
57
0.5 mm
\^
'Tn. Tnu.aJu.
Fig. 39. Hucmaphysalis sp. (H. cornigera group), nymph.
58
R„,n„»v, Y..rNr. Univkbsitv Science Bulletin
Fic. 40. H aemaphysalis sp. (//. cornigero group), larva.
Biological Series, Vol. 15, No. 1
TicK.s
59
i.s slightly lateral rather than medial. Tarsi III and IV have small midventral spurs and api- coventral spurs.
DlSTHlBUTIOX .\ND HoSTS:
Anastos (1950) discussed the distribution and hosts of H. cornigera. It is widely distribut- ed in Southeast Asia from Bunna through the East Indies and Taiwan. It has not yet been reported from the Ryukyu Islands, and the only records from Japan are those from the Izu Is- lands off the coast of Honshu now included in Tokvo-to. Recorded hosts for cornwera are cat- tie, deer, water buffalo, pigs, man, and a wide variety of smaller mammals and birds. Speci- mens reported from Japan have been taken from cattle, and on one occasion from a dog.
Mai' U. Known distribution of Hacmuphysalis .sp. H. cornigera group).
Biology: Unknown.
DisE.\sK Relationship: Unknown.
Haenuiphtjsalis doenitzi Warburton and Nuttall (Fig. 41)
Uaeina))lujsaU.y doenitzi Warburton and Nuttall, 1909:64-65; Keegan and Toshioka, 1957:18, PI. 28; Saito, 1959b: 193-209.
Discussion:
This species was originally described from material collected from a water-hen in Singa- pore in 1909. A female tick taken from a pheas- ant at Akune, Kagoshima Prefecture, 10 January 1956, by Toshioka was the specimen illustrated in Keegan and Toshioka (1957, PI. 28). This specimen was listed as species incertae sedis in that manual because Kohls had expressed the opinion that it was very similar to H. doenitzi except for the 5/5 dentition of the hypostome (original description of doenitzi, 4/4). Hoog- straal ( 1969a ) has expressed the opinion that laboratory rearing will be necessary in order to resolve the problem involving forms in the Ori- ental tropics with 4/4 dentition versus temperate forms with 5/5 dentition.
Diagnosis:
H. doenitzi is a very small species. In the female, the boot-shaped palpal article II is sharply salient laterally and at right angles to the axis of the basis capituli; palpal article III is subtriangular from a dorsal view, without dorsal spur; the cornua is poorly developed; the hypostome has 5/5 dentition (4/4 according to Nuttall and Warburton, 1915); and coxae I-IV have very short, blunt spurs.
Distribution and Hosts:
In addition to the original record of H. doenitzi from a water-hen in Singapore, this species was recorded from Indochina (Touman- off, 1944) without the specific locality or host being mentioned. In Japan, Saito ( 1959b ) col- lected immature forms and adults from pheas- ants and a male from vegetation. In his labora- tory he fed H. doenitzi on a domestic fowl, Callus gallus domesticus.
Biology: Unknown.
Disease Relationship: Unknown.
Haenwphiisalis flava Neumann
(Fig. 42-45)
Haemaphysalis flava Neumann, 1897:333-336, Fig. 3 (in part), 1901:260 (in part); Donitz, 1905:129-130; Warburton, 1908:510-512, Fig. 1, 2; Blanchard, 1909:151; Nuttall and War- burton, 1915:408-410, Fig. 342, 343; Kishida, 1922a: 846-854, 1927:985, Fig. 1897, 1936: 142,
60
Bhicham Younc University Science Bulletin
K^>U9«4jJLk.
Fic. 41. Ilacmaphysalis doenitzi, female.
Biological Series, Vol. 15, No. 1
Ticks
61
Map 12. Known distribution of Haei}uiphi/salis docnitzi.
1947:972, Fig. 2767; Sugimoto, 1935:23-24.
Fig. 5 in PL I, Fig. 2 in PI. II, Fig. 4 in PI.
Ill, 1936c:580, 1937a:308; Nakamura and Ya-
jima, 1937:157; Yajima, 1942:. 503-509, Fig. 3;
Nakatsuji, 1942:294; Itagaki, Noda, and
Yamaguchi, 1944:1-97, PI. 5, 1959:1-118, PI.
5; Shigemori, A.so, and Yajima, 1953: 290-293;
Asanuma et al, 1955:127-128; Asanuma,
1956a: 91-96, 1965a: 111-112, Fig. 7.7, 1965b:
400, Fig. 224; Asanuma and Sakurai, 1958:
28-39; Keegan and Toshioka, 1957:14-15, PI.
21, 22; Ohara et al., 1959:8-17, Fig. 1-6, PI.
I-XIV; Saito, 1959b: 193-209, 1960a: 303-321,
1960b: 189-239, Fig. 1-37, 1962a: 127-146,
1962b: 147-159; Honma and Ohara, 1961:1-7;
Saito, Ohara, and Unagami, 1960: 323-329;
Saito and Ohara, 1961:1-32; Saito et al., 1965:
14.3-159. Ilaeuuiphysalis flava var. armata Neumann,
1905:237-238. Haemaphtisalis flava armata: Neumann, 1911:
112; Kishida. 1922a:S52. Ilaemaphiisalis i Kishida, 1922a: 852-853, PI. 26;
Nakamura and Yajima, 1937:148. Uacmtqyhijsalifi orientalis Ki.shida, 19.34. ( noincn
nuiltmi); Keegan and To.shioka, 1957:14.
Haemapbysalis tcatanabei Yajima, 1942:500-502, Fig. 1; Keegan and Toshioka, 1957:30.
Discussion:
Neumann's original description (1897) was based on material collected from dog, horse, cow, hare, and vegetation in Japan. However, as pointed out by Warburton ( 1908 ) , this ma- terial (type series) contained at least two dis- tinctly different species. They were H. flava and H. campanulata. Moreover, Warburton ( 1908 ) found japonica mixed with flava and campanulata in a tube labeled as No. 137 "H. flava," identified by Neumann himself, which had been deposited in the British Museum. More confusingly, Donitz (1905) pointed out that H. neunumni Donitz, 1905 (now considered to be a synonvm of H. longicornis Neumann, 1901), was found with flava in the lot which Neumann sent him.
Neumann ( 1897 ) originally described the male of flava as having a long spur on coxa IV, and in 1901 he mentioned the presence of a short-spurred form of flava (Warburton, 1908: 513). Subsequently (1905), he considered his original long-spurred fonn to be a variety {flava var. armata) and in 1911 made the short-spurred fonn the type of H. flava flava. This was criti- cized by Warburton ( 1908 ) . To make the sitiia- tion clear, Warburton ( 1908 ) redescribed the true flava ( = Neumann's long-spurred form) based on one female and one male given him by Donitz, which were taken from a dog at Ise and from cattle or a horse at Hiroshima.
In the same paper Warburton established japonica ( = Neumann's short-spurred forni) and campanulata (see discussion of H. campanulata and H. japonica). Nuttall and Warburton (1915) repeated Warburton's original description and added measurements and collection records of Indian specimens. However, the specimens from India are now considered to be a separate spe- cies, H. indoflava.
Kishida ( 1922a ) described H. i. the holotype of which was a male collected from a wild boar on Mt. Kiyosumi, Honshu, by Aoki, 30 October 1909. Although the holotvpe specimen was not available for study, both the description and the illustrations agree with the morphological char- acters of H. flava. Asanuma ( 1969 ) concurs with this opinion.
Keegan and Toshioka (1957) examined the holotype of H. orientally- Kishida, 19.34, taken from a dog at Tokyo, and listed it as a synonym of flava. Since the original description was not published, orientally is apparently a nomen nudum.
62
BiiicHAM Young University Science Bulletin
Yajiina (1942) described H. ivatanabei based oil two females taken from a horse at Okiinaka- yama, Ivvate Prefecture, Honshu, 16 October 1939, by Y. Watanabe. One of the holotype series, No. 2535 (the date cm the label was 16 VIII 1939), mounted on a slide, was loaned to us for study through the courtesy of Dr. Kitaoka, and it is our opinion that //. wcildiiahei is con- specific with //. flava.
Diagnosis:
This species resembles H. japonica but mav be distinguished from that species in that in both se.xes the spur on coxa IV is longer than the others; in the male the spur is straight, sharply pointed, and approximately as long as the coxa; in the female it is slightly longer than the other short, blunt, subequal spurs on coxae I-III. This species is commonly found on dogs, whereas there are no collection records of ja- ponica from dogs.
Distribution and Hosts:
According to the available literature, this species is known only from Japan and Korea. In Japan it has been collected on all four princi- pal islands. Nuttall and Warburton (1915) re- ferred to records from dogs, wild boar (S»/.s- cristatus), and sheep in India; however, Dhanda and Bhat (1968) consider the materials from India to be H. indoflava. They state that there is no evidence that H. (//.)' flava Neumann occurs in India. Adult H. flava have been taken most frequently from hares and dogs, but they have also been found in considerable numbers on cows, horses, wild boar, deer, and bear (Ursus thihetanus japonicus). Asanuma et al. (1955) listed hosts for each stage and indicated that bird hosts were parasitized onlv bv imma- ture fonns. Immature fomis are also found on the same hosts as adults and on small rodents. Saito (1959b) and Saito et al. (1965) al.so studied occurrence of this tick in Niigata Pre- fecture including Sado Island, and they found fhiva from animals other than those on Asanu- ma's list. We have identified specimens as flava from the following prefectures: Honshu - Ao- mori, Iwate, Fukiishima, Niigata, Ibaraki, Tochi- gi. Tokyo, Kanagawa. Chiba, Shizuoka, Nagano. Aichi, Shiga, Kyoto, Nara, Mic, Hiroshima, and Shimanc; Kyushu - Miyazaki, Kagoshima, and Kumamoto; and Shikoku - Ehimc and Kagawa.
Biology:
The biology of H. flava has been studied bv Asanuma ( 195'6a), Asanuma and Sakurai ( 1958) in Chiba Prefecture, and by Saito (1959b) in Niigata Prefecture. Asanuma found that on
Hjemaphysalis flava
^ Collected and examined
O Collection records from literature
(J Combination of above
v^)OMivagi '^Fukusfimia
Shimane Hirosfiim.
^i\iiigdta(^ \fFukusfiii Shiga /Na^no "^°'^^'9;
^Kvo.o\ />no-, r^ff"":'!'
y *Tokushima
) Kumamoto FMiyazaki
YamanashiA: ^
. i, ""fJt^Chiba (J^^^anagawa
■ (jMivake le .Aichi
Tokyo
Hachl]o
Map 1.3. Known distribntion of llaemaphijsalia flava.
hares, adults were most abundant in late autumn and winter. Saito observed a similar trend in Niigata Prefecture, except for the fact that there was a peak in March instead of winter, and adults disappeared in summer. Nymphs were found active on hares throughout the year, but generally adults were most abundant from au- tumn to winter. Larvae were found most fre- (luently in November in Niigata Prefecture and in summer in Chiba Prefecture. Asanuma (1956a) studied the laboratory life cycle of H. flava. Eggs of H. flava hatched in 24-25 days when maintained at a constant tempeature of .30 C and at 80-90? relative hmnidity. Eggs de- posited early in May hatched in 28 days at room temperature, while eggs deposited in late May hatched in .32 days. Duration of larval attach- ment to the host varied from 2-13 days. Most lar\ae dropped from the host on the third, fourth, or fifth day after attachment. After de- tachment from the host, larvae entered a post- parasitic period which varied in length accord- ing to temperature. At a constant temperature of 30 C and at 1005? relative humidity, duration of the postparasitic period was from 9-15 days (average 11 days). At 25 C, it was from 14-21 days. When kept at room temperature, after
Biological Series, Vol. 15, No. 1
Ticks
63
Fig. 42. Haemaphy salts flava, female.
64
HiiiGHAM VouNC University Science Bulletin
K.9><^^aA(xhA.
Fig. 43. Hacmaphi/salis fltiva, male.
Biological Series, Vol. 15, No. I
Ticks
65
0.2 r
Tyi. Triu-o^
0.2 mm
Fic. 44. Hacnuipht/scili.s jlava, nymph.
66
Brigham Younc University Science Bulletin
Fig. 45. Haemuphi/salK fhivii, larva.
Biological Series, Vol. 15, No. 1
TiCK.S
67
dropping off the ho.st, it was three month.s for those detaching in February; for those detach- ing in June, it was 17 days; for those detaching in September, it was 17 davs; for those detach- ing in October, it was about 32 davs; for those detaching in late November it was 5 months, and the larvae molted the following May. Nymphs remained attached to the host for 2-13 days. A large number fell from the host on the fifth or sixth dav. Again the postparasitic period varied in length at different temperatures. It was 16 days under laboratorv conditions, at a constant temperature of 30 C and 100% relative humidity. At room temperature this period was consider- ably longer. For nymphs which engorged in February, it was about three months; in July it was 16 davs; and in September it was 21 days. The length of time necessarv for the female to engorge varied from 9 to 28 days in several experiments. Engorged females varied consider- ably in weight. Duration of the preoviposition period was 5-90 davs and was correlated with changes in temperature. Females which fell from the host in December did not oviposit until
the following spring, but as the weather became wanner this period decreased in length. In March it was about one month, in May it was about 10 davs, and in July it was about 6 days. In general, heavier female ticks deposited more eggs than lighter females. Partially fed females were also observed to lay eggs. At a constant temperature of 30 C the ovipositing period lasted 12-22 days. The number of eggs produced daily reached a maximum several days after the be- ginning of the period. From 80-90% of the eggs were deposited in the first half of the oviposition period.
Asanuma also stated that some ticks which were removed from hosts by scratching or forced to leave because of the death of the host before they were fully engorged could oviposit or feed again on a new host. This observation is of epidemiological importance in that such ticks are capable of mechanical transmission of tula- remia to new hosts.
In the 406th laboratory, ticks were fed on rabbits' ears and the following results were obtained :
Generation and Stage
Phase
Laboratory life cycle of HaemaphysaUs flava (reared on rabbits)
Period in Days
Remarks
P Adult
P Adult
P Adult
F. Egg
F, Larva
F, Larva
F, Nymph F, Nvmph
Feeding
Postparasitic ( Preoviposition ) Oviposition
Incubation
Feeding
Postparasitic (Premoldng) Feeding Postparasitic ( Premolting)
|
7-11 |
3 unengorged females |
|
(67-J-0294) were placed |
|
|
on host on 23 Dec. 1967. |
|
|
11-24 |
|
|
8-27 |
Egg number: 1,219-2,564 |
|
(average 1,937) |
|
|
23-34 |
5,.550 larvae hatched |
|
from 5,811 eggs. |
|
|
3-5 |
Larvae were placed on |
|
host on 1 March 1968. |
|
|
13-19 |
|
|
3-5 |
|
|
17-20 |
The rearing experiment extended from 23 December 1967 to the middle of April 1968. Labora- tory conditions were similar to those for HaemapJitjsaJis campanulata.
(1956a) conducted a U.S. Government spon-
DiSE.\SE Rel.\tioxship :
From a medical viewpoint, //. flava may be the most important tick in Japan because of its probable role in the epidemiology of tularemia on Honshu. The distribution of this tick in J.ipan has been discussed by many Japanese acarologists (see Appendix 2). Asanuma
sored study on the biology of H. flava and on its efficiency as a vector of tularemia under ex- perimental conditions.
Japanese literature contains only two records in which tularemia was definitelv transmitted to man by tick bite (Ohara and Ichikawa, 1962).
68
Bbigham Young I'niversity Science Bulletin
Asanuina ct al. (1955) carried out a series of experiments to determine the effectiveness of //. flava as a vector of tularemia. Adult and immature //. flava were fed on rabbits infected with tularemia. Ticks which had fed were pooled according to stage of development and sex, and material from each pool was inoculated into mice and guinea pigs. AH experimental ani- mals became infected with tularemia, and the etiological agent was recovered from the heart blood of each of these animals. Positive results were also obtained with adults which were in- fected as nymphs and with nymphs which had been infected as larvae. Studies on transovarial transmission were not attempted.
Saito (1959a, 1962b) removed ticks from wild hares which had died of tularemia and found them to be infected with Pasteurellu ttiki- rensLs: He also observed that infected ticks were able to transmit P. tularensis from host to host, but he could not isolate the causal agent of tula- remia from ticks collected in the field by the flagging method.
HaemaphtjsaJis jormosensis Neumann (Fig. 46-49)
HaemaphijsaUs formoscmis Neumann, 1913:135- 1.37; Kishida, 1922a: 852.
Discussion:
This species has not been previously reported from Japan, Korea, or the Ryukyu Islands, but Neumann (1913) and Ogura (1936) reported it from dogs in Taiwan, and Sugimoto (1936c) cited these records. Other papers by Sugimoto published in 1935-19.37 include records from Taiwan but not from Japan or the Ryukvu Is- lands. Kishida (1922a) included H. jormosensis in his key, but he did not give data on hosts and locality. Thus, the occurrence of this tick in Japan was in doubt until it was collected by 406th personnel from wild boar and the Ryukyu black rabbit on Amami Oshima.
Diagnosis:
This species is easilv distinguished from other haemaphysalid ticks bv the following com- bination of characters: the palps are blunt; pal- pal article II is very slightly salient laterallv but extends scarcely beyond the basis capituli, and the lateral margins are almost parallel, with a round posteroexternal margin; palpal article III has ;i blunt tip (dorsal view) and a ventral spur but no dorsal spur; the hvpostome dentition is 6/6 in the male ;md 4 4 in the female.
female, probably from either Ili/strix bengalensis or Urstis torquattis, in Bunna. Neumann's (1913) record was from dogs in Taiwan. Toumanoff (1944) reported //. formosensis collected from deer in Haut-Chlong, Indochina.
DiSTHIBUTION AND IIoSTS:
Nuttall and VVarburton
1915) collected a
A/ah 14. Known distrilnition of Hiicnuiplii/niilit jonno- scn.sis.
Biology: Unknown.
Disease Relationship: Unknown.
Haevuiphijsalis fujisana Kitaoka (Fig. 50-53)
llacmuplujsaUs fiijisano Kitaoka, 1970:73-81, Fig. 1-2.5.
Discussion :
The relationship between //. jujisami and other HaciiuiphysnUs species which parasitize cattle in Japan re(|uires additional study.
//. fitjisdiui is related to //. ronciniui Koch, 1844, but the male of //. concinna is e;isily sepa- rated from //. fujisana in that palpal segment III is chelate. Tlie female closely resembles //. con- cinna in general appearance, but "the palpi ot the
Biological Sehies, Vol. 15, No. 1
Ticks
69
1/1/ \l y \jg.
P'lo. 46. HacmaphysalLt fnrmosensis, female.
70
BniCHAM VouNC University Science Bulletin
Fig. 47. Haemaphysalui formosensis, male.
Biological Sehies, \'ol. 15, No. 1
Ticks
71
0. 2 mm
Fic. 48. Haenuiphysalis formosensls, nymph.
72
BnicHAM Young University Science Bulletin
Kic. 49. Ilaemaphi/sdlis formosensis, larva.
Biological Series, Vol. 15, No. 1
Ticks
73
Fig. .50. Haemaphijsalis fujisana, male. (After Kitaoka. 1970)
74
Brigham Voung University Science Bulletin
Fig. 51. Jhicmaphij.tnlis fuji.santi, female. ( After Kitaoka, 1970)
Biological Series, \'ol. 15, No. 1
Ticks
75
Fig. 52. Haemaphysalin fujisaiui, nymph. ( Alter Kitaoka, 1970)
76
Bricham Young University Science Bulletin
Fic. 53. Iltwmaphi/sali.s fiijisdiui. lar\a. ( After Kit. ink, i, 1970)
Biological Series, Vol. 15, No, 1
Ticks
77
adult of the new .species have a .small, abrupt basolatcral salience that is lacking in //. con- ciniui. The iininature form has more salient palpi than the adult" ( Kitaoka, 1970).
This species is also similar to //. japonica in tliat the spur on coxa I is longer than those of coxae II-IV. But in both sexes the dorsal view of palpal segment II differs from U. japonica. In //. fttjisana, palpal segment II has an external profile uliich is longer (male), and subequal to or slightly longer (female) than that of segment III, and the basolateral (posteroexternal) junc- ture is more sharplv ridged; vcntrallv palpal seg- ment II bears 3 infraintemal setae. The spira- cular plate of the male is longer than wide and without a dorsal process.
In H. jafwiiica, on the contrary, palpal seg- ment II has an external profile distinctly shorter than that of segment III, and the basolateral juncture is somewhat blunt with a dull angle; N'entrally it bears more (5-6) infraintemal setae. The spiracular plate of the male is wider than long and with a dorsal process.
DlSTRIBUTIOX AND HoSTS:
This species has been collected only from
\I \r 1.5. Known distribntioii of lliiciniiphi/suli.-; jiiji.siinii.
Holstein calves near the foot of Mt. Fuji, near Fujinomiya, Shizuoka, Japan.
Biology:
Kitaoka ( 1970 ) reared the progeny of one engorged female. About 250 eggs were deposit- ed during a period of nine days. Eggs began to hatch in 17 days. Larvae fed for 3 to 4 days and molted II to 18 days later at 25 C and in 8 to 10 days at 30 C. Nymphs fed for 3 to 4 days in August and 4 to 5 days in September. Nymphs molted in about 15.9 days (males) and 17.5 days (females) at 25 C.
Disease Relationship: Unknown.
Haemaphiisalis Injstiicis Supino
(Fig. 54-57)
Haemaphi/salis hi/stricis Supino, 1897:251-253; Kishida, 1922a: 852; Sugimoto, 19,37a: 31 1-313, 19.37b: 609; Asanuma, l947b:973. Fig. 2771, 1965a: 111; Keegan and Toshioka, 1957:15, PI. 23, 24; Kawashima, 1963:103; Hoogstraal, Trapido, and Kohls, 1965:467-480, Fig. 1-38.
Haemaplit/salis bisj)inosa: Neumann, 1901:261- 262; (Not Neumann, 1897).
Haemapliiisalis nishhiamai: Sugimoto, 1935:29- 33, Pl.'l-III, 19.36c: 58 1 -582, Pi. II, 19.37a:317, 1937d:287-290, PI. Ill; Nakamura and Yajima, 1937:148.
Hacmaphiisalis iuasakii Sugimoto, 1937:610, PI. III.
Discussion:
Sugimoto (1937a, 19.37b) reported the oc- currence of //. hiisfricis in Yonakuni, one of the Ryukyu Islands, but made no mention of the host. Kishida (1922a) included this species in his key but gave no collection details. Asanuma (1947b) gave a brief account of this species in the Illustrated Encijclopedia of the Fauna of Japan. A collection by 406th personnel includes four lots from wild boar on Ishigaki (Is.) and Iriomote (Is.) in the Ryukyus, but in more re- cent surveys this species was collected from dogs, pigs, and the Ryukyu black rabbit on Amami Oshima, Kagoshima Prefecture.
H. nishit/amai was originally described by Sugimoto, who examined 13 males taken from a wild boar, a dog, and a man on Taiwan. Subse(|uently (19.37a), he listed an additional male from a dog, and in 1937 he redescribed the male and described the female. He stated (1937d) that the male of nishUjamm was very close to lujslricis but distinct from it in that palpal segment II has "a fairlv strong ventral
78
Bbigham Young University Science Bulletin
Fic. 54. Hacmnphi/sali.s hi/stricis, female.
Biological Series, Vol. 15, No. 1
Ticks
79
Fig. .55. Haemaphysalis In/stricis, male.
80
Brigham Vounc UNivEHsiTi- Science Bulletin
Fic. 56. Hactnaphtjsali.'i hi/strici-i, n\Tnph.
Biological Series. Vol. 15, No. 1
Ticks
81
0 2mm
0.2 m m
Fic. 57. Haemaphij sails htfstricis^ larva.
82
Bbigham Young University Science Bulletin
rotrogrado spine" and "lateral grooves modinm, hunt, including one festoon" and the first leg is of dark color. However, these nioqihological characters are not enough to separate this species from hijstrici.'!, and his description and illustra- tions fit litistricis well. Hoogstraal et al. (1965) examined two males and two females of ni.slti- i/anuii from the Sugimoto collection in Taiwan and considered them conspecific with hi/stricis.
Sugimoto ( 19.37h) described H. iwasukii based on a male specimen taken from the bed- ding site of a wild pig on Ishigaki (Is.). Al- though the specimen was not available for study, the illustration and description indicate that iiLasdkii is probablv a svnonym of liijstricis. Recently. Hoogstraal et al'. (1965) studied the identity of liystricis, and they considered //. hvasakii Sugimoto, 19.37 to be a synonym of hijstrici.s.
Neumann (1901) erroneously described a species under the name of hispinosa using speci- mens of a different species. These specimens were not those from India, the type locality of hisj)inos(i. but others collected from China and Japan. The Japanese lot contained two female specimens (Hilgendorf Coll. Berlin Museum). According to Nuttall and Warburton (1915), Neumann misidentified them, and what he ( 1901 ) described as hhpinosa was possibly lit/s- tricis. The following year Neumann (1902) ex- amined Supino's type of hi/stricis. wrongly sy- nonvmi7.ed his])inosa imder Jitistricis, regarding them to be identical. This was criticized by Warburton (1908) and Nuttall and Warburton (1915), but if the specimens were actually hystricis, Neumann's report ( 1901 ) might be the first collection record of In/stricis from Japan.
Dl.\GNOSIS:
H. Injstrici.'i is similar to longicorni.i in having dorsal and ventral spurs on palpal article III, and palpal articl(> II is salient laterallv. How- ever, it mav be distinguished from /ongifor/iiv by the presence of a dorsal knoblike, anteroin- temal projection on palpal article II, and a me- dian projection on the ventroposterior margin of palpal article II. In the male there are no lateral grooves, and in the female the comua is short, the basis capituli is at least twice as wide as long with a depression in the middle of the porous area, and coxa I has a short spur.
Distribution .vno Hosts:
This species has previously been considered to be widely distributed in temperate and tropi- cal eastern Asia and on numerous nearby islands. According to Hoogstraal et al. (1965), the dis- tribution is restricted to a subtropical and tem-
perate belt extending from the eastern Hiina- lavas of India to the coasts of Vietnam and Fu- kien Fro\ince of China, and to Taiwan, Ishigaki, and Okinawa. Amami Oshima may be now in- cluded in this belt as a result of recent collec- tions. All previously published records from China, Ceylon, Macao, Celebes, Java, Borneo, and the Malay Peninsula are either definitely in error or highh' suspect. Authentic localities for ht/.^tricis lie between about 87° and 130° east longitude and 29° and 16° north latitude.
Adults of hystricis have a fairly wide host range.
Hoogstraal et al. (1965) listed hosts as hu- mans, domestic dogs, buffalo, wild boars (Sits scrofci). tigers, hog-badgers { Arctonyx coUaris), Teledu badger {Myclantts imtlticei)s), sambar deer (Cervus unicolor), and the Formosan bark- ing deer or muntijac ( Muiifiacus rcevesi micru- nis ) .
The often-(|uoted record of a single female from the spiny hill tortoise, Geoemyda s])itwsa, (Nuttall and Warburton, 1915:425) in "India" is considered (juestionable since G. spinosa does not occur in India.
Map 1(1 Known distribution ot Hiicnuiphi/scilis tii/xtriiis.
Biological Series, Vol. 15, No. 1
Ticks
83
Biology:
Ecologically tlii.s tick appears to he a forest- inhabiting species which occurs at low to inter- mediate ele\'ations. Newly molted females often have been observed crawhiig on birds by Hoog- straal et al. (1965), and it is suppo.sed that they seek a host unusuallv early, perhaps several days before they reach a developmental condition that
allows attachment and feeding; or else nymphs molt to adults on the host rather than on the ground. At the 406th Medical Laboratory nymphs fed on rabbits' ears dropped off the host after they had fully engorged. Seventy larvae collected on vegetation on Aniami Oshi- ma were reared to adults on rabbits' ears. These adults were used for subsequent life cycle studies.
Generation and Stage
Pliase
Laboratory lite cycle of Haemaphysalis hijstricis (reared on rabbits)
Period in Days
Remarks
P Adult
P Adult
P Adult
F. Egg F, Larva
F, Larva
F, Nvmph F, Nvmph
Feeding
Postparasitic ( Preoviposition ) Oviposition
Incubation Feeding
Postparasitic ( Premolting ) Feeding Postparasitic ( Premolting)
8-9
4 unen gorged females (67-J-0308) were placed on host on 29 Nov. 1967.
|
7-8 |
|
|
15-18 |
Egg number: 3,245-4,528 |
|
(average 3,953) |
|
|
32-34 |
15,698 larvae from 15,814 eggs |
|
3-5 |
Larvae were placed on host |
|
on 25 Jan. 1968. |
10 +
2-5 17-22
The rearing experiment extended from 29 November 1967 to the middle of March 1968, in the 406th Medical Laboratory mider conditions similar to those for Haemaphysalis campamihita.
DisE.\SK Relationship: Unknown.
Haemaphysalis japonica Warburton (Fig. 58-60)
UaemaphtisaUs japonica Warburton, 1908:512- 513, Fig. .3, 4; Nuttall and VVarlnirton, 1915:402-403, Fig. 336-337; Kishida, 1922a: 852; Itagaki, Noda, and Yamaguchi, 1944:1- 97, PI. 6, 1959:1-118, PI. 6; Keegan and Toshi- oka, 1957:16-17, PI. 27; Asanuma, 1958:279, 1965a: 112.
Uaemaphtjsalis flava flava Neumann, 1911:112; Xuttall and Warburton, 1915:402, 514.
llacmaiihijsalls jczoensis Ogura and Takada, 1927:205-206, PI. XV; Nakamura and Yajima, 1937:148.
Discussion:
As already mentioned under //. campanulata and H. flava, the type specimens of japonica were found together with flava and campanu- lata in tube No. 173 (No. 137 ?) at the British Museum. Warburton (1908) stated that japonica
was undoubtedly conspecific with the short- spurred fonii of flava mentioned by Neumann (1901). He did not give the collection data of the type. Subquently, Nuttall and Warburton (1915) based their discussion of japonica on Warburton's type specimens, collected from the Japanese serow, Nemorhaedtis crispus ( ~ Capri- cornis crispus), at Hondo, Japan, and 24 males taken from a roe-buck at Kansu, China. Hondo, the type locality, is a town on one of the Ama- kusa Islands, Kumamoto Prefecture, Kyushu.
It is apparent that Warburton misspelled the name as japonnica for japonica. There is evi- dence that this species has been mistakenly re- ported as H. concinna Koch, 1844. It has also been reported as H. jezoensis Ogura and Ta- kada, 1927, now considered to be a synonym of H. japonica. Keegan and Toshioka (1957) ex- amined specimens labeled concinna and jezoen- sis loaned bv Japanese workers, and found them to be japonica.
Nuttall and Warburton (1915) described H. japonica var. douglasi based on four males col- lected from roe-deer at Ten-an-fu. Shensi, north-
84
Bbiciiam Young Uni\ersitv Science Bulletin
Fig. 58. llacmaphysalis japonicu, female.
Biological Series, Vol. 15, No. 1
Ticks
85
Fig. 59. Haemaphysalis japonica, male.
86
Brigham Young University Science Bulletin
0, 1 mm
Fig. 60. Iliicmaplu/sdlis japonica, nymph.
Biological Series, Vol. 15, No. 1
TiCK.S
87
em China, and they .stated that it i.s distinctly .smaller, narrower in front with straighter sides and lateral grooves, and with a longer and sharper ventral spur on palpal article III. Pom- erantzev (1950) referred to dotigkisi as a sub- species and indicated that it inhabits the USSR. The relationship of japonica to japonica dougla.si is not clear at the moment, and further study will be necessary to elucidate this problem. Hoog- straal (196S) considers japonicii and japonica douglasi to be identical.
Dr.\CNOSis:
This species is close to //. flava and has been mistakenly reported as H. concinna because of the similaritv of the palpal profile. However, unlike flava. the spur on coxa I is longer than the other coxal spurs in both sexes, and especial- ly in the male, coxa IV does not have such a long, sharp spur as in flava. It is also distin- guishable from concinna in that the tips of palpal article III are not so strongly recurved inward to become "pincerlike" in the male; the external profile of palpal article III is longer than that of II, while thev are subequal in con- cinna; palpal artical II has 5-6 infraintemal setae; and the spurs on coxae II and III are short but apparentlv extend beyond the posterior margin of the co.xae. See diagnosis of H. con- cinna for comparison with that species.
Dlstribution and Hosts:
Recorded distribution in Japan is Hokkaido, .\omori. and Nagano prefectures. It is also known to occur in Korea and China. Pomerant- zev ( 19.50) gave the distribution of japonica as Southern Primor, USSR, in addition to the above, and the distribution of japonica doiiglasi as the USSR and northern China. Hosts record- ed in Japan and Korea are horses, cattle, Japa- nese serow, deer, dogs, and on one occasion, one nvmph from a wild hare. Pomerantzev ( 19.50 ) gave the hosts for japonica douoja.si as deer, bear, lynx, badger, hedgehog, cattle, dogs, and humans. Immature stages were found on small rodents, badgers, scjuirrels, deer, dogs, Mitstela sibirica. and birds.
Biology:
Unstudied in Japaii. Pomerantzev (1950) briefly cited the seasonal occurrence of japonica doug^Ia.si in the USSR. "From October to March, males are exclusively found on animals, often in large numbers; from March through April, they arc joined bv females which parasitize mainly during the 1st half of the summer, are found un- til October and cease to be found entirely in the winter. Young stages feed from June to August
M.'ii' 17. Known di.strihution of Haemaplu/salis japonica.
and often are found along with semi-mature ones." Somov and Shestakov (1963) describe the habitat of H. japonica dotiglasi as follows: ". . . inhabits foothills and valleys with rich grass growths and shrub thickets. On exposed slopes this tick species rarely occurs." (Translation through the courtesy of the U.S. Naval Medical Research Unit No. 3, Cairo, UAR. )
Disease Relationship:
Not known in Japan. In the USSR, japonica dou<ila.n is suspected of being a vector of the virus of Russian spring-summer encephalitis (Pomerantzev, 1950), Rickett.'iia sihericus (Si- berian Tick Typhus) has been isolated from H. japonica dotip^la.si by Russian workers (Somov and Shestakov, 1963).
HaemaphysaJis hitaokai Hoogstraal (Fig. 61-64)
HacmapJujsalis kitaokai Hoogstraal, 1969b:211-
221, Fig. 1-35. Haemaphijsalis amhifiiia Neumann, 1906:217
(not Neumann, 1901), 1911:109; Kitaoka and
Morii, 1967a: 145-152, Fig. 1-9. HaenuiphijsaU.'i inerniis: Nuttall and Warburton,
88
Brigham Young Univehsitv Science Bulletin
Fig. 61. Hucviaphysulis kitaokai, female.
Biological Series, Vol. 15, No. 1
Ticks
89
Fig. 62. Haemaphysalis kitaokai, male.
90
Bbigham Young Univehsity Science Bulletin
Fig. 6.3. Haemaphtjsalis kiUiokai, nymph.
Biological Series, Vol. 15, No. 1
Ticks
91
0 2 mm
0 J mm
F"iG. 64. Hiwmaphtjsolis kitaokai, larva.
92
Brigham Young University Science Bulletin
1915:367; Kishida, 1922a: 852; Sugiinoto, 1937a:313-.317, PI. II, Fig. 7, 8 in PI. VIII; Keegan and Toshioka, 1957:16, PI. 25, 26; Iwa.saki ct al., 1964:1-10; T.suchic, Iwasaki, and Okui, 1966:1-15; A.sanuma, 1965a:lll- 112. Haevuipln/sdlis inermis var. aponomoides War- burton, 1913:128-130, Fig. 8; Nuttall and Warhurton, 1915:367-369.'
Discussion:
This unic|uc Japanese species has been called //. tinibi<!^ii(i or //. incniii'; including //. inermis var. aponomoides. Neumann ( 1906 ) reported one female amhU^ua which was found together with H. flava and an undetermined Ixodes sp. in a lot at the British Museum. The lot was taken from Naeniorhacdus crisptis ( = Capri- cornis crisjtits), the Japanese serow. Nuttall and Warburton (1915) considered ambigua to be a synonyni of inermis. They did not see the fe- male specimen referred to as ambigua by Neu- mann (1906); therefore, they carefully cited Neumann's record in brackets under synonyms of inermis. This record was repeated by Kishida (1922a) and Sugimoto (1937a) without provok- ing any attention until Kitaoka and Morii (1967a) expressed doubt about the occurrence of H. inermis in Japan (see below).
Warburton (191.3) described a variety, iner- mis var. aponomoides, based on 2.3 females taken from the Himalayan Zebu of India, and Nuttall and Warburton (1915) stated that they found one female of the variety from a horse at "Fu- koka" (apparently misspelled for "Fukuoka"), Japan (Berlin Museum No. 173).
Sugimoto ( 1937a ) described a Formosan species as inermis based on eight females taken from Cervus unicolor stvinhoii on Formosa, giv- ing the distribution of the species as occurring also in Japan (see above).
Keegan and Toshioka (1957) identified the Japanese population as H. inerinis, and no fur- ther attempt was made to clarify the status of the Japanese specimen of inermis var. aponomo- ides reported by Nuttall and Warburton (1915).
Hoogstraal ( 1962), having examined the type of inermis var. aponomoides and also the tAvo Fonnosan specimens (RML lot 16603, off "wild deer," Su-ao, Formosa, sent by Dr. Sugimoto), raised the variety to the species H. aponomoides for the population at high altitudes in the Hima- layas and on Foniiosa, He (1962), however, did not see the female specimen ( Berlin Mu- seum No. 173) of inermi.^ var. aponomoides.
On the other hand, Kitaoka and Morii (1967a) indicated that there are structural dif-
ferences between specimens from the Japanese population and H. inermis of Eurasia, and they suggested that the Japanese material should be a species separate from valid Eurasian inermis. They tentatively adopted the name ambigua for Japanese material.
Hoogstraal (1969) subsequently examined the one female specimen ( Berlin Museum No. 173) and the two female andngita identified by Neumann ( 1906), and after comparing the new- ly collected Japanese specimens with numerous collections of the related Eurasian species, he concluded that the Japanese population repre- sents a distinct species and described it as //. kitaokai. He considers the female specimen of iiu;rmis var. aponomoides ( Berlin Museum No. 173) to be H. kitaokai and //. ambigua a syno- nym of H. inermis. He reexamined Sugimoto's Formosan inermis (RML lot 16603), which he once identified as ajionomoides (1962) and ten- tatively assigned them to //. kitaokai. The speci- mens were in poor condition.
Diagnosis:
This species is morphologically distinct from all other haeniaphvsalids in Japan. The uni(|ue characters are as follows: in both sexes the palpi are elongate as in Ixodes spp., palpal article I is short but visible from a dorsal view; palpal arti- cle II is not salient laterallv; palpal article III does not have either a dorsal or a ventral retro- grade spur; the basis capituli has no cornua; lateral grooves are absent; coxa I has an ex- tremely short internal spur. It is close to H. inermis Birula of southern Europe, southwestern USSR, and northern Iran, but can be distin- guished from that species in that there are three dorsointemal setae and a single ventrointernal seta on palpal segment II of the male, and these setae number two and one respectively in the female; the male hvpostome is 2.2 times longer than broad compared with three times longer than broad in inermi.s; in the nymphs and larvae the tarsi are moderateb- stout in kitaokai but iKirrowly elongate in inermis. and the coxa) spurs are rudimentary in inermis but discrete in kitaokai (Hoogstraal,' 1969b).
DiSTlUBMTION ."VNT) HoSTS:
This speci(>s has been collected only in Japan on deer and the Japanese serow in Mie, Shiga, and Kanagawa prefectures, but more commonlv on cattle in the Sanbe Pasture. Shimane Prefec lure. Yajimas single female specimen was from a cow in N;igasaki Prefecture. There are some records of inermis or inermis var. aponomoides, but occurrence of these species in Japan is con- sidered to be erroneous (as mentioned above).
Biological Series, Vol. 15, No, 1
Ticks
93
Haemaphysalis kitaokai
^ Collected and examined
O Collection records from literature
(J Combination of dbove
Map 18. Known di.stribution of Haemaphysalin kitaokiii.
Biology:
Iwasaki et al. (1964) observed the seasonal occurrence of this tick in a pasture at the foot of Mt. Sanbe, Sliimane Prefecture. There are two peaks of adult prevalence, one is a small peak appearing early in spring to June and another more p r o m i n ent peak from October to No- vember. No larval and nymphal fomis were found in the pasture or on cattle. Kitaoka and Morii ( 1967a) suggest this is possibly due to the fact that the immature forms are exceptionally rapid feeders and are attached to the host for a short period of several hours. Tsuchie et al. (1966) reported observations on overwintering of this tick. On 19 November 1964, they placed jars containing 60 engorged ticks and dried, fallen leaves out-of-doors, and found (|uiescent ticks 1 cm deep under the leaves on 18 Febru- ary the following year. The 53 females which had successfully wintered oviposited in early May. Ten specimens were observed and they laid 109-1,020 (average 617) eggs during the period from 5 May until 6 June, when the av- erage outdoor temperature began to rise to 15 C at the Sanbe Pasture.
Kitaoka and Morii (1967a) studied the bi- ology of this species at Kodaira, Tokyo, using material originally collected at Sanbe Pasture in November 1962. They suggested that the life cycle was probably completed in one year in nature. The following table is a summary of data from Kitaoka and Morii (1967a). Nymphs and adults were fed on calves' ears.
Generation
and Stage
Plia;
Laboratory life cycle of Haemaphijsalis kitaokai Period in Days
25 C
Outdoor
Remarks
|
p |
.\dult |
Feeding |
- |
8-15 |
In Nov. and Dec. 1962. Engorged on calves on 1 Dec. |
|
P |
.\dult |
Postparasitic (Preoviposition) |
5-7 |
94-102 |
|
|
p |
Adult |
Oviposition |
2.3-36 |
57-81 |
Laid by 14 females |
|
( No. of eggs ) |
(369-899) |
(117-959) |
at 25 C and 19 females outdoors. |
||
|
F, |
Egg |
Incubation |
28-33 |
40-51 |
|
|
F, |
Larva |
Feeding |
1:50-20:00 hrs |
In May, fed on mice, rabbits, or calves. |
|
|
F, |
Nymph |
Feeding |
3:30-6:00 hrs |
In June and July, fed on calves. |
|
|
F, |
Nymph |
Postparasitic (Premolting) |
24, 27 |
A female emerged after 24 days; a male, after 27 days. |
Thirt}' females and eleven males were col- laboratory, but only four semiengorged females lected in the field and brought into the 406th were fed on rabbits' ears. One female placed
94
Bmr.HAM Young ITniversitv Science Bulletin
on the rabbit on 8 November dropped off on 15 November and laid 823 eggs from 10 to 30 December from which 588 larvae hatched out beginning on 7 January. The F, progeny were fed on a rabbit's ear on 22 January, dropped off on 23 January, but all died before they molted. Another lot of F, larvae were put on a rabbit's ear at 1400 of I Fefiruai-v and were observed to have already dropped off by 0800 the next moniing. The rapid feeding of larvae was also observed, but only a single larva survived until it molted to a nvmph on 3 March. The room temperature in the laboratory was approximate- ly 25 C.
Disease Relationship: Unknown.
Haemaplnj salts longicornis Neumann (Fig. 65-68)
IIaei)ui))luj.siilis longicornis Neum;mn, J901;261, Fig. 2;' Hoogstraal, Roberts, Kohls, and Tip- ton, 1968:1197-1213, Fig. 1-36.
Haenuiphtjsalis netimanni Donitz, 1905:127-129, Fig. 4-6; VVarburton, 1908:508-519; Neu- mann, 1911:109; Kishida, 19.36:139, 1939b: 17-19, PI. 4; Trapido, 1965:160; Hoogstraal and Trapido, 1966:1192, Fig. 10-26.
Ihwnmphiisalis bispinosa (not Neumann, 1897); Nuttall and Warburton, 1915:426-4.33, Fig. 358-362 (in part); Kishida, 1922b:8.52; Ogura and Takada, 1927:204-205, PI. 14; Sugimoto, 1936c:580, PI. I, 1937b: 605-609, PI. II; Naka- mura and Yajima, 1937:151, PI. IV, 1942a:21- 33, 1942b:.34-39, PI. II, III; Nakatsuji, 1942: 294-295; Yajima, 1942:, 502-509, Fig. 2, 1963: 103; Itagaki, Noda, and Yamaguc^ii, 1944:1- 97, PI. 3, 1959:1-118, PI. 3; Asanuma, 1947a: 7-14, 1947b:97.3, Fig. 2770, 1958:279, 1965a: 111-112, Fig. 7.6, 1965b:399, Fig. 222; Shige- mori, Aso, and Yajima, 1953 :290"- 293; Namba, 1953:130-1.3,5, 1954:49-55, 1958:1-99, PI. I, 196.3a:21-28, 1963b: 103-114; Chikaki, Otake, and Miiira, 1936: .53-62; Keegan and Toshi- oka, 19.57:11-12, PI. 1.5-16; Kitaoka and Ya- jima, 195Sa: 1.35-147, 195Sb: 149-162, 1958c: 179-188; Kitaoka, 1961a:S5-95, 1961b:96-104, 1961c: 105-1 12, 1961d: 142-144, 1962:106-111; Nagahana and Mat.suo, 1962:119-121; Saito and Ohara, 1961:1-32, Fig. 19, 20, 26; Saito, 1962a: 127-146, Fig. 1-30;" Saito ct al., 1965: 143-159, Fig. 1-24.
Discussion:
For many years this tick has been referred to as //. bispinosa Neumann, 1897 in the Japa- nese literature. Hoogstraal and Trapido (1966) selected and studied a female neotype of Jmpi-
nosa in the Nuttall Collection (N167) in the British Museum and a paralectotype of neu- manni (N1425) in the Neumann Collection in the National Veterinary School, Toulouse. The former specimen was taken from southern Madras, India, and the latter was from Japan. Since then it has been thought doubtful that l)isj)inosa actuallv occurs in Japan. .All materials in the 406th collection have been examined by Kohls and Hoogstraal and identified as H. neti- manni. It is likely that the specimens in the Japanese literature are also netimanni, since our specimens were collected from the same host species and in essentially the same geographical localities. More recently Hoogstraal at al. (1968) compared numerous specimens from Australia, New Zealand, tropical Asia, northern China, Japan, and Korea. They confinned that specimens of "bispinosa" and "netimanni" from temperate northern China, Japan, Korea, and northeastern USSR are structurallv identical to those from Australia and New Zealand referred to as H. longicornis Neumann, 1901, which ap- pears to be a prior name applicable to these northern populations. We accept their opinion and use the name H. longicornis in this paper. Published reports indicate there are tvvo strains (bisexual and parthenogenetic) within the Japa- nese population. H. ])is])inosa Neumann, 189"?, a tropical Asian species, tvpically a bisexual spe- cies, is known from India, Ceylon, Pakistan, Nepal to Bumia, Thailand, and the Malay Peninsula.
Diagnosis:
H. longicornis is very close to H. bispinosa Neumann, 1897 but may be distinguished from that species on the basis of its larger size: male 2.47 mm (2..30-2.57); female 2.60 mm (2.0-2.8); while in bis))inosa. male, 2,0 mm; female, 2.2 mm in length; hvpostome with dentition of 5 5 (rarely 4/4 in female), bispinosa always with 4/4 dental formula; UMnphs with 3/3 dental fonnula, whereas bis])inosa with 2/2; nymphs without dorsal spur on palpal article III, but bispinosa has a bulging posterodorsal margin of palpal article III. //. longicornis may also be distinguished from H. hystriris on the basis of the following characters: well-de\el()ped comua; no dorsal knoblike projection on anterointenial margin of palpal article II; lateral grooves pres- ent in the male and shape of spiraciilar plate as illustrated.
DlSTBIBUTIOX AND HoSTS:
Neumann's original description (1897) was based on females from a cow. Kempsev. New South Wales, .Australia. .According to Hoog-
Biological Series, Vol. 15, No. 1
Ticks
95
Fio. 65. Hacmaphi/salis Inngicornis, female.
96
BniGHAM Young UNrvERsrrY Science Bulletin
Fit;. 66. Haenmphijsah.s longicornis, male.
BioLiKiicAL Series, Vol. 15, No. 1
Ticks
97
Fig. 67. Haemaphtjsalis longicomis, nymph.
98
Bhicham Vounc Univehsitv Science Bulletin
0.2 m m
■9n. '^TUoaAi.
0.2 mm
Fig. 68. Hacm(ii>)uistilix /orifjifonii,?, larva.
Biological Series, Vol. 15, No. 1
TiCK.S
m
straal et al. (1968), this species occurs in Aus- geographical locahty, and our collection con- tralia, New Zealand, New Caledonia, Fiji Is- Finns Kitaoka's finding ( 196Id) of predominant- lands, Tonga, Friendly Islands, Efate Island, ly parthenogeiietic populations in the north of New Hebrides, northern China (Peking), Japan, Japan and bisexual populations in the southern Korea, and northeastern USSR. They suggest islands. However, we have both parthenogenetie that the bisexual population is presumably pre- and bisexual populations from Oki Island. In dominant on the Korean peninsula. It may be the laboratory the bisexual strain readily con- noteworthy that reliable records of this spe- verts to parthenogenetie type of reproduction cies on the R)ukyu Islands cannot be found. The when the females are isolated from the males, distribution and sex ratio of this species in Japan Bremner ( 1959) found a ratio of approximately have been reviewed by Kitaoka (I96Id). I male to 400 females in the Australian popula-
H. longicornis parasitizes many animals from tion and no spermatozoa in all males dissected,
small to large mammals, and birds. Hosts re- irrespective of whether they were fed or unfed,
corded are cattle, horses, sheep, goats, deer, Excellent physiological and ecological studies
bears, pigs, foxes, racoons, badgers, cats, dogs, of this species (as bispinosa) were done by
rabbits, house sparrows, skylarks, thrushes, tiir- Kitaoka (1961d), Namba (1953, 1954, 1958,
kevs. ducks, chickens, pheasants, and humans. 1963a, 1963b), and Asanuma (1947a). Naka-
mura and Yajima ( 1942a ) studied the laboratory
life cycle. In the 406th Medical Laboratory both
iiiOLOG\ . parthenogenetie and bisexual strains were reared
This tick is known to be parthenogenetie. In on rabbits' ears, and their life cycle data is sum- Japan the ratio of males to females varies with marized in the following table.
Laboratory life cycle of Haemapluisalis longicornis
( parthenogenetie and bisexual populations ) (reared on rabbits)
Period Number of Days
Least Most Average
P B P B P B
Oviposition to hatching 24 38 31 39 27.5 38.6
Resting (larvae) 3 3 5 4 4.0 3.5
Feeding (larvae) 4 5 5 7 4.5 6.0
Drop-off to molt (larvae) 14 20 17 22 15.5 21.0
Resting (nymphs) 2 3 3 3 2.5 3.0
Feeding (nymphs) 5 4 7 5 6.0 4.5
Drop-off to molt (nymphs) 12 14 16 16 14.0 15.0
Resting (adults) . . 4 2 6 4 5.0 3.0
Feeding (adults) . 11 11 19 13 15.0 12.0
Drop-off to oviposition (females) 3 3 7 6 5.0 4.5
Oviposition (females) 11 17 27 18 19.0 17.5
Number of eggs 2,024 2,740
Egg hatch (fc) 94 96
P = Parthenogenetie population B — Bisexual population
Disease Rel-^tionshu': Re.marks;
This species is a proven vector of Coxiella The only known example of triploidy in ticks hiirnclii (Q fever) among Australian cattle and occurs in //. lonoicorni.s. Oliver and Bremner is also capable of transmitting the causal virus ( 1968 ) found the chromosome number to be 32 of Russian spring-summer encephalitis in north- or .3.3 in the obligate parthenogenetie female and em USSR. In Japan it is suspected of trans- 31 in the reproductively nonfunctional male in mitting bovine piroplasmosis. Experimental in- the Australian population. According to infor- fection of calves with Theileria nmtans by the mation supplied by Oliver and Tanaka (1968), bite of this species (listed as bispinosa) was the chromosome number of the Japanese par- reported by Ishihara and Ishii (19.56). Namba thenogenetic population is 32 or 33 in females (1963b) also referred to this species as an im- and 31 or .32 in males (t^vo males found among portant vector of T. nmtans in Japan. approximately 3,000 samples of females which
100
Bhiciiam Young University Science Bulletin
Haemaphysalis longicornis
^ Collecled jnd examnied
O Collection records from literature
(^ Combination of above
, Abash in
Kamikawa_ Ishfkar Shinbesh) HWr-V^^ Tokachi // _
Hiyania •V^'bur
Oshima
'( iAomori
flQpukushima Tochigi
•,'C^lwaTe
NiigalaQ/ oQFukushii (^ / mGumma
#Tottor, (^"^''O.^S^'"'"'' ^-^., ^\*-' ^ TokyoOQChiba
Tsushima y'riJ^Q^C^^ C\h,z^^Q 0,h,m3
W H.joshimaMto-ykayama H,,,:, lki*^£j:ukuoka 7^<J!>S V ji"<^^0,tay/ Tokushima Hach,|oko#^j
^ C^^0j5'^'"^'"o*o Hachijo
NagasakCq'lVI,y^^3l^,
Kagoshima
•QO
Aogashima^
Mat 19. Known clistribiitioii of Haciiuiphi/salis lon'^- icornis.
wore produced from ten batches of eggs), and in the Japanese bisexual population it is 22 in females and 21 in males. In parthenogenetic females from Cheju Do, Korea, 22 to 24 chromo- somes have been observed.
Haenuiphysalii megaspinosa Saito
(Fig. 69-72)
Uiiemaphiistilis uiegospinosa Saito, 1969:87-96, Fig. 1-16.
Discussion:
Saito. using the flagging method, collected adults of this species in a cedar forest of a moun- tainous area in Kanagavva Prefecture. Personnel of the 406th Medical Laboratory collected this species from a wild boar, deer, and Japanese serow. Three lots. J-()177, J-()179, and J-01S4, collected from a wild boar and two deer in Mie and Shiga prefectures, contained this species along with //. jlava. The female of H. mega- spinosa resembles //. flava: therefore, the rec- ords of fhivii from thes(> hosts which have been previously reported by Japanese acarologists might possibly include H. megaspinosa. Other
specimens were loaned for study through the courtesy of Dr. Kitaoka, and two females were found in Dr. Yajima's collection at the National Institute of Animal Health, Kodaira, Tokyo.
Diagnosis:
This species is close to //. fhiva in general appearance but may be distinguished in that the body color is brownish and darker, and the body size is remarkably larger in both sexes. In the male, the conuia is longer; the dentition of the hvpostome is 5 5 to 6/6; the posteroexternal juncture of palpal segment II is somewhat blunt, whereas in flava the angle is sharper, the junc- ture itself being a small tip; legs more robust; trochanteral spur I is much reduced; coxae I and IV have relatively shorter spurs, especially on coxa IV the spur is stronger, more or less curved, wider at base; tarsi II-IV have a triangular, spur- like ventral projection at approximately the mid- dle. The female of this species is quite similar to that of flava. However, palpal segment II differs ventrobasally as in the male (see illustra- tions), and coxa IV has a longer spur.
Distribution and Hosts:
Saito collected H. megaspinosa from vegeta- tion, but adults of this species were collected from a wild boar, deer, and Japanese serow {Capricornis crisjnts). This species is known at pre.'ent from Mie, Shiga, Miyazaki, and Kanaga- wa prefectures. Immature forms have not yet been collected from the field.
Biology:
This species fed on rabbits in the 406th Med- ical Laboratory', and the life cycle was studied. Compared with //. flava, it showed slower de- \elopment in each stage. Five engorged females (6S-J-0301 ) began oviposition on 29 to 31 March 1968 and laid batches of .368-2,077 eggs during the ovipositing period of 23 to 26 days. After the incubation period of 33 to 37 days, larvae hatched out on 3 to 6 May. The hatch rates of the two batches were 98.1^ and 96.9?. A con- siderable number of larvae from the two colo- nies were fed on a rabbit for life-cycle studies. In the first series of rearing experiments, larvae were put on the host 9 days after h;itching and nvmphs were put on the host 4 davs after molt- ing. In the second series, larvae were put on the host 2S days after hatching and the nymphs were put on th(" host 26 davs after molting. The feeding periods of larvae and nvmphs did not differ essentiallv between the two experimental series, and the results are sumnnirized as follows:
Biological Series, Vol. 15, No. 1
Ticks
101
0.5 mm
O.I mm
' Am<J-o
Fig. 69. Haerrmphysalis megaspinosa, female.
102
Bricham Young University Science Bulletin
2 0 mm
/W:^H
r. /\^i<,
V\c 70. Uui'muphysalix rneguspinosa, male.
Biological Series, Vol. 15, No. 1
Ticks
loa
^TTL ■nti^.
0.2 mm
Fig. 71. Uacinapliysalis megaspinosa, nymph.
104
BiucHAM YouNC University Science Bulletin
Fk;. 72. Ilacmaphiisuli'i iiiei^titipiuosa, larva.
BiDi.otiicAL Series, Vol. 15, No. 1
Ticks
105
Hoog.straal and Yamaguti,
M.\p 20. Known distribution of Hucmupht/sidis lucg^ii- spinosa.
Laboratory life cycle of Haeinaphysalis megaspinosa and Haeniaphijsalls flava
Period in Days Stage Phase megaspinosa flava
|
Adult |
Oviposition |
23-26 |
8-27 |
|
Egg |
Incubation |
33-37 |
23-34 |
|
Larva |
Feeding |
4-9 |
3-5 |
|
Larva |
Postparasitic |
17-18 |
13-19 |
|
\yinph \vmph |
(Premolting) Feeding Postparasitic ( Premolting) |
4-7 20- |
3-6 17-20 |
Eggs: megaspinosa .368-2,077 (by 5 females) flava 1,219-2,564 (by 3 females)
At approximately 25 C, reared on laboratory white rabbits.
Disease Relationship: Unknown.
IIarniaj)ht/salis pcntalagi Pospclova-Shtrom (Fig. 73-76) UacnuiphtjsaJis pcntalagi Pospelova - Shtrom, 1935:205-217. Fig. 9-12; Keegan and Toshi-
oka, 1957:28; 1970:367-374.
Discussion:
The description of this species was based on a single male specimen taken from a young Ryukyu black rabbit from Liu-Kiu, Japan, in 1927. Pospelova-Shtroin (1935) did not mention the specific locality, but Liu-Kiu is presumably Amami Oshima because the Ryukyu rabbit, Pen- talagus furnessi, is found only on Amami Oshima and Toku no Shima, and the latter may not have been easily accessible in 1927. Since 1927 no specimens resembling pentalagi have been col- lected by Japanese workers. However, personnel of the 406th Medical Laboratory succeeded in collecting H. pentalagi from the Ryukyu rabbit when the capture of this protected host was per- mitted in April 196S bv the Japanese Govern- ment.
Diagnosis:
This smallest haemaphysalid tick in Japan slightly resembles H. campanulata. but may be separated from that species as follows: coxa I has no spur, coxae II-IV have weak, blunt, and
Mai- 21. Known distribution ot Hiicniiiphi/sdli', jii'u- tiihitli.
106
Bbigham Younc University Science Bulletin
77 A^<U,
Fk;. 73. IhKiniiphij.salix pctitiiltif:,!, female.
Biological Series, Vol. 15, No. 1
Ticks
107
r A^4o
Fic. 74. Haemaphijsalis pentalagi, male.
108
Brigham Young University Science Bulletin
Fig. 75. lUwmaphysalis pcntahigi, nymph.
Biological Series, Vol. 15, No. 1
Ticks
109
-^. 7K,.*<Ju.
0.2 m m
Fig. 76. llaenuiphysulis i)entaUigi, larva.
110
BniGHAM VouNc University Science Bulletin
wide spurs which arc hardly discernible; there is no dorsal spur on palpal article III; palpal article II has a pointed, externally directed pos- teroexternal juncture; in the female, the capitu- luiii and scutum are very large in relation to the small hodv size (engorged, scutal length covers approximately half the body length); the genital aperture is on a level with coxae III; the tarsi arc not humped.
Distribution and Hosts:
Known distribution is Amami Oshima in the Uvukvu Islands. Pospclova-Shtrom's record may be trom the same island. The only known host is the Rvukvu black rabbit, Pcntahi<iu.s furnessi.
Biology:
Hearing experiments were conducted at the 4()6th Medical Laboratory.
Generation and Stage
Phase
Laboratory life cycle of UdemaphijSdlis pentalagi (reared on laboratory white rabbit)
Period in Days
Remarks
P Adult
Feeding
|
p |
Adult |
Post parasitic ( Preoviposition ) |
|
p |
Adult |
Oviposition |
|
F, |
Egg |
Incubation |
|
F, |
Larva |
Feeding |
|
F, |
Larva |
Postparasitic (Premolting) |
|
F, |
Nymph |
Feeding |
|
F, |
Nymph |
Postparasitic |
5-7
7-10
15-17
27-28
5-9
14 +
6-7
2 unengorged females (6S-J-0311) were placed on host on 16 April 1968.
Egg number: .386-523 492 larvae from 909 eggs. 50 live larvae were placed on host on 31 May. S nymphs emerged.
Nymphs died \sithout molting.
The rearing experiment was done from 16 April 1968 to July, and room conditions varied according to outside temperature and humidity.
Disease Relationship: LTnknown.
Ilacmuphi/salis ivellingtoni Nuttall and Warburton (Fig. 77) HaemaphiimUs uellingtoni Nuttall and Warbur- ton, 1908:397-398; Asanuma and Kosaka, 19.54:104-107, Fig. 1-2; Keegan and Toshi- oka, 19.57:4; Asanuma, 1965a^ 111-112.
Discussion:
Occurrence of H. iccUingtoni on the Japa- nese gray thrush, Turdtis cordis cardi.s, was re- ported by Asanuma and Kosaka (19.54). As Turdus rardis cordis is a migratory bird which spends tlu> colder months of the year in countries of Southeast Asia, Asanuma and Kosaka believe that the ticks found on Honshu probably had been carried by birds which had recently arrived from wanner areas. This bird is a summer resi- dent in Japan and breeds from April through August. Its distribution includes Hainan, Indo- china, and South China. Although specimens were not seen, figures published in the paper by Asanuma and his co-worker are excellent and leave no doubt that the specimens collected were actually uellingtoni. Since this species is
parasitic on migratory birds, it cannot be re- garded as a species which has become estab- lished in this area. Diagnosis:
The male of this tick is distinct from other haemaphvsalids in Japan in that palpal article III has a distinct retrograde spur internodorsally and a long ventral spur; palpal article II has S-9 internoventral setae (Anastos, 1950; Sumatra specimen has 7 setae); the cornua is short and blunt. According to Asanuma and Kosaka ( 19.54 ) it is close to H. pavlovsktj Pospelova- Shtrom, 19.35. but may be distinguished from that species by the longer ventral spur of palpal article III, absence of a small blunt projection on the supraintemal margin of palpal article II, and the weaker cornua. Disthibution and Hosts:
Anastos (1950) gave the distribution of this tick as Borneo, Malaya, Siam ( = Thailand), Andaman Islands, India, New Guinea, Bunna. Indonesia, and Indochina. Hosts recorded are goose, turkey, wild fowl, domestic fowl (GaUus gollu-s), buffalo (Buhidufi hrdiohs), ;Hid dog. Biology:
Unknown.
Biological Series, Vol. 15, No. 1
Ticks
111
Fig. 77. Haemuphysalis wetlingtoni, male.
112 BiiK.iiAM VouNG University Science Bulletin
Disease Relationship: groove curved around tlie anus anteriorly. Spi-
IJnknovvn racular plate round or oval. Sexual dimorphism
marked. Venter of male eoveretl with seven Genus Ixodes Latreille, 1795 plates.
Inornate. Eyes and festoons absent. Anal Type Speciks: Ixodes riciniis (Linnaeus, 1758).
Key to the Males of the Genus Ixodes'
1. Coxae without internal spur '2
Some coxae with internal spur 4
2. Legs distinctly longer than body; hypostome pointed at tip; on bats (Fig. 116)
vespertilionis
Legs not distinctly longer than body; hypostome blunt or notched at tip - 3
3. Anal grooves convergent; on sea birds (Fig. 105) ^ signatus
Anal grooves divergent or parallel; on swallows (Fig. 87) Uvidus
4. Coxa I with one spur 5
Coxa I with two spurs , 6
5. Coxa I with a long internal spur; on badger (Fig. Ill) tanuki
Coxa I with a short, blunt internal spur (Fig. 96) ovatus
6. Coxa I witli a thick, long external spur acutitarsus
Coxa I with a blunt, short external spur 7
7. Hypostome with moderate denticles; internal and external spurs of coxa I short, sub-
ecjual (Fig. 83) '^ranuJatus
Hypostome with large, strong denticles laterally; internal spur of coxa I longer than
external spur . 8
8. Spiracular plate ellipsoidal, much longer than wide, usually anglelike contour antero-
ventrallv; mostlv from western part of Japan (Fig. 92) nipponensh
Spiracular plate oval, longer than wide; mostly from northern part of Japan (Fig. 100)
persulcatus
^Ixodes philipi is not included in this key.
Key to the Females of the Genus Ixodes
\. Coxae without internal spur - 2
Some coxae with distinct internal spur - 3
2. Anal groove pointed in front; posterior amis of anal grooves divergent; on bats ( Fig.
108) simplex simplex
Anal groove rounded in front; posterior arms of anal grooves convergent or subparallel 3
3. Legs distinetlv longer than body; hypostome lanceolate; scutum widest behind middle;
on bats (Fig. 115) vespertilionis
Legs not distinetlv longer than body; hypostome blunt at tip; scutum widest at middle
or in front of middle; on birds 4
4. Auriculae well developed; on sea birds 13
-Auriculae a1)sent; on swallows (Fig. 86) Uvidus
•5. Coxa I with one spur 6
Coxa I with two spurs 8
6. Coxa H with a short external spur; scutum with posterolateral margin slightly concave
(Fig. 90) monos])inosus
Coxa H without an external spur; scutum with posterolateral margin convex 7
7. Coxa I with a long internal spur; on badger (Fig. 110) — tantiki
Coxa I with a short internal spur (Fig. 95) ovatus
8. Auriculae well developed - 9
Auriculae absent or poorlv developed 11
9. Internal spur on coxa I much longer than external spur; auriculae nonnal 10
Internal spur on coxa I nearly e(]ual to external spur; auriculae well developed laterally
(Fig. 112) ' ' turdus
10. Internal spur on coxa I sharply pointed, long, iisuallv ovcrlipping on approximately /a of coxa II; postscutal marginal bodv setae pointed, length approximately 0.08 mm;
HioLOCMCAL Series, Vol. 15, No. 1
Ticks
113
inastly from northern part.s of Japan (Fig. 99) persulcatus
Internal .spur on co.xa 1 long, u.sually overlapping anterior margin of coxa II (Fig. 80); postscutal marginal body setae scaled, forked, length approximately 0.14 mm; mostly from western parts of Japan (Fig. 91) nipponensis
11. Coxa 1 with a thick, long external spur; scutum cordate, nearly as wide as long (Fig.
78) — acutitarsus
Coxa I with a blunt, short external spur; scutum distinctly longer than wide 12
12. Anal groove slightly pointed in front; genital aperture on level with coxa III (Fig.
79) __ angustus
Anal groove round in front; genital aperture on level with coxa IV (Fig. 82) granulattts
13. Dorsum except scutum hirsute, uniformly covered with white hairs; basis capituli sub-
triangular dorsally; porous areas well separated (Fig. 10.3) philipi
Dorsum except scutimi not hirsute; basis capituli rectangular dorsally, much wider than
long; porous areas very close to each other (Fig. 104) signatwi
gan and Toshioka, 1957:18-19, PI. 29; Ohara and Tamura, 1958:2.3-24.
EscJiatocephalus acutitarsus: Neimiann, 1901: 290.
Ixodes happinus: Keegan and Toshioka, 1957:31.
Discussion:
Nuttall and Warburton (1911) discussed the synonymy of this species and mentioned that Karschs type came from Japan. These workers also stated that they had seen two females from Japan and Formosa. Sugimoto (1936b, 1936c) merely cited records of Nuttall and Warburton (1911). Kishida (1930a) reported occurrence of this species on the serow, Capricornis crispus, in Gifu, Nagano, and Toyama prefectures, Japan, and later ( 1935 ) from cattle on Formosa. Asa- numa cited the records given above and added the dog and the cow to the host list in the 1947 edition of the Illustrated Encijclopedia of the Fauna of Japan. Keegan and Toshioka (1957) examined two loaned specimens taken from man, one from Wakayama Prefecture and the other from Saitama Prefecture, and identified them as acutitarsus. Yajima's (1955) record was from a cow in Iwate Prefecture, and he reported this tick merely as n. sp. and did not provide a specific name. This specimen was identified by Mr. G, M. Kohls as acutitarsus. Keegan and Toshioka (1957) referred to this specimen as Ixodes happinus, but this is an error since Dr. Yajima had not given a name in his original description, nor used this name in any publica- tion since that time. Ohara and Tamura (1958) reported this tick on man.
DiACNOSIS:
This tick is said to be the largest in the genus Ixodes. Asanuma ( 1947b ) stated that even unengorged females measure as large as 8 mm in length. It mav be distinguished from other Ixodes spp. by the following characteris-
.M.\p 22. Known distribution of UaciiKijihi/.'idlis wcll- in^toni.
Ixodes acutitarsus (Karsch) (Fig. 78)
Haemalastor acutitarsus Karsch, 1880:141-142; Neumann, 1899:180.
Ixodes acutitarsus: Neumann, 1901:286; Nuttall and Warburton, 1911:202-203, Fig. 195; Schulze, 1935:2.34; Kishida, 19.30a:2, 19.3.5: 137; Sugimoto, 1936b: 1-20, 1936c:585; Asa- numa, 1947b:972, Fig. 2766. 1965a: 113- 116, 1965b:399, Fig. 220; Yajima, 1955:52-53; Kee-
114
BniGHAM Young University Science Bulletin
Fig. 78. Ixodes acutittnstis, female.
Biological Series, Vol. 15, No. 1
Ticks
115
tics: coxa I i.s deeply inci.sed witli two thick, long spurs; there are neither conuia or auriculae on the basis capituli; the anal and genital grooves extend dorsallv to reach the distinct lateral groove of the dorsum, thus giving a iestoonlike appearance; the legs are slender, the tarsi gradu- ally taper distally, especially in the teinale.
DiSTniBUTION AND HoSTS:
Nuttall and Warburton (1911) recorded col- lections from Japan, Formosa, and Sikkim from Japanese serow, dog, cow, ;ind human. All specimens were adults; innnature forms have never been collected.
Ixodes acutitarsus
9 Collected and examined
O Collection records from
literature (^ Combination of above
Map 2.3. Known distribution of Ixodes uciititursut;.
Biology: Unknown.
DisE.\SE Relationship: Unknown.
Ixodes angtisfus Neumann (Fig. 79-81) Ixodes anoti.stus Neumann, 1899:136; Asanuma, 1947b:97I, Fig. 276.3, 1951:1-4, 1957:2:34, 1965a:113-116," 1965b:.398, Fig. 219; Ono, 1966:62-68. Ixodes sp. 3 and sp. 61 Asanuma and SekikawM,
1952:111-116; Asanuma, 1955:1240-1242.
Discussion:
In the 1947 edition of the Illustrated Ency- clopedia of the Fauna of Japan, Asanuma (1947b) recorded the occurrence of this tick in Sakhalin and in 1951 gave a description of fe- males taken from the red-backed mouse, Cleth- rionomys rufocantts (Sundevall, 1846), in Sak- halin. Asanuma and Sekikawa (1952) and Asa- numa ( 1955) reported immature forms of Ixodes species on rodents from Hokkaido, and they temporarily numbered these forms as sp. 3 for the larvae and sp. 61 for the nymphs. Later, Asa- numa ( 1957 ) again reported these forms from Hokkaido and stated that the previously report- ed sp. 3 and sp. 61 were the immature stages of Ixodes angiistus.
Diagnosis:
The female of this species is close to /. granu- latus in the key but may be distinguished from that species in that the anal groove is slightly ogival in front; the genital aperture is on a level with coxae III; the hypostome is shorter than the palps and sharply pointed at the tip, the denticles of the inner file are subequal to those of the outer file and occur on almost the entire dentate length except two or three rows near the tip, while in grantdatus the denticles of the inner file are much smaller than those of the outer file and do not occur on the proximal two- fifths of the dentate length; the scutum has well- defined, straight carinae, and the lateral margin of the scutum has a distinct angle which sepa- rates the anterolateral and posterolateral margins.
According to Asanuma (1951), this species is also allied to /. soriciis, I. muris, I. jellisoni, and /. minor but may be distinguished from them by the smaller size and wrinkled integu- ment of the postscutal area, the dentition of the hypostome, the absence of comuae, the larger porous areas, and the strong lateral angle of the scutum.
Distribution and Hosts:
Known distribution of this tick includes North America, Canada, Alaska, Sakhalin, and Argen- tina. Hosts recorded are rodents, other small mammals, and sometimes humans.
Biology:
Little is known about the biology of this tick. Males are rarelv found on host animals, and copulation probably occurs in the nest of the host.
Disease Relationship:
Cooley (1946) reported three positive rec-
116
Brigham Young University Science Bulletin
/ \
• A.
N
Rvjuihi
Fig. 79. Ixodes aniiufihis, female.
BionK-.icAL Series, Vol. 15, No. 1
TlCK.S
117
7n.. TKic^xM,
0.2 m m
Fig. 80. Ixodes angnsUis, nymph.
118
Brigham Young University Science Bulletin
0,2 m m
0.1mm
^. oAt.
Xoo-A-
Fic. 81. Ixodes tmnustus, larva.
Biological Series, Vol. 15, No. 1
Ticks
119
Ixodes angushis
^ Collected and examined
O Collection records from
literature ^ Combination of above
Map 24. Known dLstribution of Ixodes angiistus.
ords of tlii.s tick biting humans in the USA and stated there was no suggestion of disease trans- mission in these instances.
Ixodes granulaius Supino (Fig. 82-85)
Ixodes gramddtus Supino, 1897:230-238; Asa- numa and Kosaka. 1955: 195-196; Keegan and Toshioka, 1957:19, PI. 19, 30; Arthur, 1957:683-694, Fig. 12-17, 19-24; Kawashima, 1963:103; Asanuma, 1965a: 11.3-120, 1965b: 398, Fig. 218.
Ixodes sp. 4, 51 and sp. 101 Asanuma and Seki- kawa, 1952:107-116, Fig. 1, 2, 4, 6, 8, 1953: 99-112, Fig. 9, 10, 1.5, 16\ 17.
Discussion:
Asanuma and Sekikawa (1952, 1953) report- ed the occurrence of three unnamed ticks of the genus Ixodes, including larval and nymphal fonns on rodents on Honshu, Kyushu, Hokkaido, and Shikoku. These unnamed ticks were ten- tatively numbered as Ixodes sp. 4 for larvae, Ixodes sp. 51 for nymphs, and Ixodes sp. 101 for adults. Keegan and Toshioka ( 1957) stated that Asanuma (1956a) indicated that these speci-
mens were actually grcinuhitus. Arthur (1957) discussed Korean specimens.
Diagnosis:
/. gramdattis is easily distinguished from other Ixodes species by the following combina- tion of characters: coxa I has two short spurs, the internal spur is slightly longer than the ex- ternal spur; the cornua is very short in the male and absent in the female; the auriculae are poor- ly developed; the scutum of the female is finely granulated, longer than wide, covering more than half the length of the dorsum and the scapulae are pointed.
Distribution and Hosts:
Anastos (1950) cited distribution records from India, Burma, and the East Indies. Kohls ( 1950 ) gave a collection record from Luzon Island in the Philippines and (1957b) from Ma- laya. Collection records indicate that this tick is primarily a parasite of rodents. Asanuma and Kosaka ( 1955) took a female of this species from the thrush, Turdits celaenopus celaenojnis, on Miyake (Is.), Tokyo-to. Prior to our survey, material in the 406th Medical Laboratory col- lections was from the Ryukyu Islands.
|
Ixodes granulatus |
1 1 |
|
|
^ Collected and examined |
*' // |
|
|
O Collection records from literature 3 Combination OkinawaC |
'^7«ftv |
|
|
« ^ |
' / '^^ ' |
|
|
Irioniote " ^^i;;:^^^ |
||
|
^-^^''^^ |
||
|
g*^ft *^ ^QKochi |
KanagawaQf/y |
Chiba yake |
|
HachijokoOpj Hachijo |
/[x |
|
|
'qO o .00 ;oo.™ |
S. Korea 0\ |
|
|
0 JOO 40C . |
V^ |
|
Map 2.5. Known distrihiition of Ixodes jlrdnulatus.
120
Bhigham Young University Science Bulletin
S.SHIBATA
KiG. 82. Ixodes- finmuhitiis. female
Biological Series, Vol. 15, No. 1
Ticks
121
Fic. 8.3. Ixodes nramdattis, male.
122
Bricham Young University Science Bulletin
Fig. 84. Ixodes g,r(inulnlits. nvmpli
Biological Series, Vol. 15, No. 1
Ticks
123
Fig. 85. Ixodes granulatus, larva.
124
Bhicham Young University Science Bulletin
Biology:
The following life-cycle data is taken from Nadchatram (1960:220).
The life cycle of the progeny of 3 female
Ixodes graiuthitiis
reared in the laboratory on white rats
Period in Days
Females detached from host 0
Females oviposited after 12-14
Females continued oviposition for 16-20
Larvae hatched out after 28-32
Larvae were fed after 15-25
Larvae fed for 4- 6
Engorged larvae quiescent after 6- 8
Nymphs emerged after 14-18
Nymphs were fed after 14-42
Nymphs fed for 6-10
Engorged nvmphs (juiescent after 5- 7
Adults emerged after 16-20
Adults fed after 28-56
Adults fed for 10-14
Total duration to complete life cycle 174-272
Disease Relationship: Unknown.
Ixodes lividtis C. L. Koch (Fig. 86-89)
Ixodes lividus C. L. Koch, 1844:234; Ono, 1967: 217; Uchikawa and Sato, 1969:95-97.
Discussion:
This tick had not been reported from Japan until recently, when Ono (1967) reported it from the nest of the swallow, Riparia riporia ijimae, from Hokkaido, and Uchikawa and Sato (1969) collected it from the nest of Delichon urbica dasijpus in Nagano Prefecture. Three other lots of specimens were collected bv 406th Medical Laboratory personnel in Shimane and ( aimma prefectures.
Ardiur (1963) discussed synonymy and dis- tribution, and he considers 7. pUnnheus, I. cani- •vugfl, and two subspecies, I. I. havaricus and /. /. ohotrUicus, both described by Schulze and Schlottke, to be synonyms of lividus.
DlACXOSIS:
This is the only ixodid tick found on swal- lows in Japan, and while it h;is no unusual mor- phological characters, the following points mav serve to distinguish it from other species: body oval; the coxae have no internal spurs; in front of the anus the anal groove is round; the posterior arms are almost parallel or slightly
divergent; palps broad; the suture between articles II and III sometimes indistinct; cornu;i absent; auriculae ;il)sent. In the female the scutum is cordate, longer than wide, widest an- terior to the middle; hypostome blunt at tip with 6-7 distinct lateral denticles. In the male the palps are broiid ;iiid oval; the hypostome is dis- tinctly notched ;it the tip uith l;iint indications ot scalelike denticles.
DiSTIUBUTION AND HoSTS:
Arthur (1963) states tliat this species has ;in extensive distribution, including Britain, Ger- many (as two .subspecies), France, Kazak SSR, Guryev Province. Arthur suggests this tick may occur wherever sand martins nest. Hosts re- corded are swallows, Riparia sp., and the Japa- nese house nuutin, Delichon urbica dasijpus.
Ixodes Itvidiis
^ Collected and examined
O Collection records from
literature ^ Combination ot above
M.\i> 26. Known distrihiitidn of Ixodes lividus.
BlOLOOY: j
Little is known of tlie biology of this species. .\rtliur ( 1963) st;ites that the males do not occur on the host, but their presence in the nests sug- gests that copulation occurs off the host. In Russia various biological aspects of this species j have been studied by Glashchinska\a-Babenko (1956) (see Babcnko', 1956). Xuttall and War-
Biological Series, Vol. 15, No. 1
Ticks
125
|
r |
ly |
|
|
i I M |
||
|
■■( ' |
^ |
|
|
M |
||
|
A/ > |
\ 1 |
' M |
|
'4 |
||
|
rr |
\x 1 |
Sonobe
Fig. 86. Ixodes Uvkhis, female.
126
Brigham Young University Science Bulletin
sonobe
Fig. 87. Ixiidcw tiiidus, male
Biological Series, Vol. 15, No. 1
Ticks
127
^'
Fic. 88. Ixodes liiidus, nymph.
128
Bhicham Young University Science Bulletin
0,1mm
-TTA-vfJ-o,
Kic;. 89. Ixodes litUlus. l,ir\a.
BioLOciCAL Sehies, Vol. 15, No. 1
Ticks
129
burtons (1911) comineiits on the biology ot /. canhu^ii apply to /. Uvidus. In the 406th Medical Laboratory eight engorged females laid 257-810 (average 5.58) eggs,"which hatched 25- 28 days after the beginning of oviposition. Larvae were placed on rabbits" ears, and 13-17 days later they molted to nymphs.
Disease Relationship: UnknowTi.
Ixodes monospinosus Saito
(Fig. 90)
Ixodes monosiiinostis Saito, 1967:107-112, Fig. 2-9.
Discussion:
Saito's ( 1967 ) description was based on a single female removed from a mountain climber in Niigata Prefecture. Saito illustrated (Fig. 2-9) the dorsal aspect, capituluni, genital and anal regions, spiracular plate, tarsi and coxae (palpi may have been missing). The holotype was not available for study, but the following diagnosis was provided by Saito.
Diagnosis:
The female of /. monospinosus is easily dif- ferentiated from known ixodids from Japan and other countries bv the following series of fea- tures: hvpostome long, with four dental files on each side; auriculae poorly developed; coxa I with onlv a spinelike, stout, inner spur, and the posterior margins of coxa I are thickened transversely; coxae II-IV each have only an ex- ternal spur and a moderately convex ventral sur- face; scutum and carinae well defined, lateral grooves distinct, large punctations on median- posterior areas, and posterolateral margins slight- Iv concave.
Distribution and Hosts:
Saito lists man as the host and the specific locality as Mt. lide, Mitakambara-gun, Niigata Prefecture, Japan. A single female specimen collected by the flagging method in Mie Pre- fecture was loaned through the courtesy of Dr. Kitaoka and illustrated in this paper.
Biology: UnknowTi.
Diskase Rel.\tionship: Unknown.
Ixodes nipponensis Kitaoka and Saito (Fig. 91-94)
Ixodes nipponensis Kitaoka and Saito, 1967:74- 83, Fig. 1-19.
|
Ixodes monospinosus . ,' •0 e |
[ \ |
||
|
^ Collected and examined |
/ |
||
|
O Collection records from k<^0 literature tj^ , |
/ |
||
|
Kf\ |
|||
|
^ Combination of above A ll |
V^ |
n |
|
|
tf // |
')}\V |
'*^" |
|
|
' '^/'\ |
M |
^ |
|
|
? ? '°?"" / \ |
% |
, |
|
|
/7 |
|||
|
o fy-^ ^^ j |
tV |
||
|
^J^--^ ^ A/r |
■ |
||
|
c2 ^'^^ |
gatal |
||
|
# /^W.V^ |
^ |
- |
|
|
* |
X |
i\ |
|
|
.J „ ,» =„.„ |
^ |
. \ |
|
|
' •' |
|||
Map 27. Known distribution of Ixodes monospinosus.
Ixodes persideatus persulcatus: Keegan and
Toshioka, 1957:21-22, PI. 34 (in part). Ixodes sp. LA and NA Ono, 1962a :24-29, Fig. 1-
3, 7, 9, 11, 14, 1962b:155, 1966:62-68, Fig. 1,
B. ?Ixodes ricinus: Neumann, 1904:452, 1911:12-13;
Nuttall and Warburton, 1911:156, 1915:4.33. ?Ixodes ovatus Neumann, 1899:116-118 (in
part ) . ?Ixodes persulcatus: Morel, 1963:925-927. ?Ixodes ])ersulcatus persidcahis: Saito, 1958:70,
1959b: 199-20.3, 1960a :.30.3-.321, 1962a: 127-146,
1962b: 147-1.59; Saito and Ohara, 1961:1-32;
Saito, Ohara, and Unagami, 1960:,323-.329.
Discussion:
Til is species was previously referred to as /. ricinus in Japan, and there is evidence that it has been reported as /. persulcatus. Kitaoka and Saito ( 1967 ) compared a Japanese population with Russian ricinus and persulcatus, and also with ricinus from Scotland and Egypt. They found several morphological differences among these populations and concluded that both per- sulcatus and a species they described, nipponen- sis, occur in Japan. Ono ( 1962ii ) described im-
130
Brigham Young University Science Bulletin
I.O m m
0.5 m m
Kic. 9(). Ixodes monospitwsus, feni;de.
Biological Series, Vol. 15, No. 1
Ticks
131
^,i}hTauJ(i
Fic. 91. Ixodes iiippuriensis, fem;Ue.
132
BiiiGHAM Young University Science Bulletin
'*^ <^^«i<.»«^<»-
Fit:. 92. Ixodes iii/j/JDiicii.vi.s, male
Bi()loc:k;al Series, Vol. 15, No. 1
Ticks
133
Fio. 93. Ixodes ni/j/ioiicruvw, nvmpli.
134
Bhigham Young University Science Bulletin
Fic;. 94. Ixodes nipponcnfii.s, huM
Biological Skries, Vol. 15, No. 1
Ticks
135
mature tick.s, Ixodes .sp. LA (larva) and Ixodes ■sp. NA (nymph), reared from egg.s laid by an undetermined .species of Ixodes. These immature ticks were similar to Ixodes sp. 2 and Ixodes sp. 54 of Asanuma and Sekikawa (1952, 1953), hut separable from these by a combination of sev- eral morphological characters. Ono ( 1962a ) be- lieved that Ixodes sp. 2 and sp. 54 were the im- mature fonns of /. persidcatus and also expressed doubt about the identitv of /. /). perstdcatus, which had been used bv Saito, Ohara, and Una- gami (1960).
Kitaoka and Saito ( 1967 ) showed that Ixodes sp. LA and Ixodes sp. NA were larval and inmphal fonns of Ixodes nip])onensis. Though the adults of nipponensis are very similar to those of persidcatus, the immature forms are separable from each other. Consequently, rec- ords of ricinus reported bv earlier |apanese workers might possiblv include nipponensis.
The material recorded in Keegan and Toshi- oka (1957) included onlv persidcatus, but these specimens were reexamined bv Yamaguti with the help of Dr. Kitaoka, and they concluded that several lots previouslv identified as persidcatus actuallv included nipponensis. Kitaoka pointed out that the specimen on which the illustration of persidcatus persidcatus (PI. 34) in Keegan and Toshioka ( 1957 ) is based, is nipponensis.
DiAGXOSis:
This species is quite similar to /. persidcatus but mav be distinguished from that species in that the bodv is smaller and is covered a little more denselv with thick setae; coxa I has a slightlv shorter spur than persidcatus: in the male, the internal spur on coxa I is shorter and reaches the anterior margin of coxa II, while it is long and partially covers coxa II in perstdcatus; the spiracular plate is much elongated longitu- dinallv and ellipsoidal; in the female, the internal spur on coxa I is shorter and does not cover K of coxa II; the marginal dorsal setae are forked, thicker and approximately 1.7 times longer than the pointed, simple setae in per.sulcatus. More- over, Kitaoka and Saito (1967) and Ono (1962a, 1966) compared the differences between im- mature stages and concluded thev were on a specific level. See also diagnosis of per.sulcatus.
Distribution axd Hosts:
/. nipponensis has been recorded by Kitaoka and Saito (1967) from Honshu, Kyushu, Shiko- ku, and Sado Island, and from a few localities in Hokkaido by Ono ( 1966). Records of the 406th Medical Laboratory include Shiga, Shimane, Mie, Hiroshima, Tokyo, Kagoshima, and Kochi pre- fectures. Hosts recorded are cattle, horse, dog,
badger, hare, weasel, and man. Immature stages are found on small rodents and birds. In a forth- coming publication by Kitaoka detailed informa- tion will be given.
Biology:
An engorged female (67-J-0341) laid eggs in the 406th laboratory and the progeny were used for life cycle studies.
M.\p 28. Known dLstrihution of Ixodes nipjioneruis.
Disease Relationship: Unknown.
Ixodes ovatus Neumann (Fig. 95-98)
Ixodes ovatus Neumann, 1899:116-118, Fig. 2, 3 (in part), 1904:452-453, 1911:18; Donitz, 1905:132; Kishida, 1930a:3; Nakamura and Yajima, 1937:141; Keegan and Toshioka, 1957:29-30; Morel, 1963:925-928.
Ixodes ricinus var. ovatus: Nuttall and Warbur- ton, 1911:158.
Ixodes japonensis Neumann, 1904:458-459, 1911: 22; Nuttall and Warburton, 1911:208-209, Fig. 200, 201; Kishida, 1930a: 3; Nakamura and Yajima, 1937:145-146, PI. Ill; A.sanuma, 1947b: 971, Fig. 2765; Keegan and Toshioka,
136
BmcHAM Young University Science Bulletin
Generation
and Stai^c
F, Larva F, Larva
F, Nynipli F, Nymph
Phase
Laboratory life cvcle of Ixodes nipponensis (reared on rabbits)
Period in Days
Remarks
Feeding
I'ostpaiasitic
(Preniolting)
Feeding
Postparasitic
(Preniolting)
5 30
4
33
Larvae were placed on host on 5 July 1967.
Only 2 nymphs molted to females.
July 1967 to October and the air-conditioned room tem-
199-203, 147-159,
and Yainaguchi, 1944: PI. 2. 1930a: 4-5; Nakamura
The rearing experiment extended from perature did not exceed 25 C.
1957:20-21, PI. 31, 32; Saito, 1959b
1960a; 303-321, 1962a: 127-146, 1962b
1964:59-66; Saito and Ohara, 1961:l-.32; Saito,
Ohara, and Unagami. 1960:323-329; Saito
et al„ 1965:143-159; Asanuma, 1956a:91-96,
1957:2.34, 1965a:] 13-1 16, Fig. 7, S, 1965b:
398, Fig. 217; Ono, 1966:62-68. Ixodes frecjttens Ogura and Takada, 1927:201-
202, PI. Ill; Schulze, 19.30:294-303, Fig. 10;
Kishida, 1930a:3; Nakamura and Yajima,
1937:146-148, PI. IV; Yajima, 1942:507-509,
Fig. 6; Itagaki, Noda,
1-97, PI. 2, 1959:1-118. Ixodes carinatus Kishida
and Yajima, 19.37:140; Keegan and Toshioka,
1957:26, PI. 41. Ixodes sp. 6 and sp. .52 Asanuma and Sekikavva,
1952:107-116, Fig. .3, 9, 1953:99-112, Fig. 11,
12; Asanuma, 19.55:1240-1242; Ono, 1962a:24-
29, Fig. L5, 1962b; 155, 1966:62-68. ?Ixodes shinchikuensis Sugiinoto, 1937:330-331,
PI. VI and IX. ?Ixodes taiwanetiiis Sugimoto, 19.36:336-.346, PI.
II, 1937a :331-333, PI. VII and VIII.
Discussion:
In the Japanese literature this tick has been reported as /. joponensis, I. frecpicns. and some- times /. carinatus.
Ixodes ovatus was originally described bv Neumann (1899) on the basis of material from Amur and Japan. The Japanese material includ- ed four female specimens, one from a horse at Akita collected by Taguchi (No. 702), the others from a Lepus sp., a horse, and a dog at Saga collected by Yainaguchi (No. 703, 704, 705 re- spectively, in the Neumann Collection in the School of Veterinary Medicine in Toulouse). According to NuttaJl and Warburton (1911), these Japanese specimens contained typical ricimis," and subsetiuently Neumann (1904) re-
'Nlorvl (196}) slimvcd llinl tlicsc riciiius I No r02 iind 703) wcro iidiwllv /. pcrsulcnlin. I>nl llicsi' uiiKlU or ]iii(;lu mil iiuUiilo / ponrnsii Kitnnkn and Saito.
described ovatus based on two females from a horse and a dog at Saga (No. 704 and 705) as separate from the other t\\'o. Nuttall and War- burton ( 191 1 ) cited Neumann's description ( 1904 ) , and in ;iccordaiice with Donitz's opinion they noted that the ditferences between the two species were only varietal. It is apparent that Nuttall and Warburton did not see the type specimens. Therefore, they placed ovatus as a variety of ricinus and treated jaj)onensis as a valid species separate from ovatus.
Neumann's original description of /. jajioncn- ■SK ( 1904 ) did not include illustrations. Later, Nuttall and Warburton (1911) provided illus- trations (their Fig. 200 and 201) and a rede- scription of jajwnen.sis based on Neumann's type specimen collected in the vicinity of Tokyo.
Pomerantzev ( 1950 ) considered ovatus as a valid species occurring in Japan, but this appears to be merely a citation from Neumann's original record.
Ogura and Takada (1927) described /. fre- quem\ but Keegan and Toshioka (19.57) sug- gested, "After a study of hosts and localities, it seems clear that frequens is a synonym" (of jafwuensis-ovatus). Thev also examined the holotvpe specimen of /. carinatus Kishida, 19.30 and stated that "the holotype is very much like an example of japonensis, although the basis capituli mav be relatively wider than in that species." More recently Morel (1963) showed that /. jajwnrnsi.s: I. fre(juens, I. carinatus. I. shincliikuensis, and /. taixcanensis were all syno- nyms of /. ovatus. The latter two species were described bv Sugimoto on the basis of Taiwan specimens, ;ind Morel (1963) expressed uncer- tainty regarding their status, probably because of lack of specimens. Although the holotype specimen of carinatus was not available for this study, the original illustration by Kishida (1930a) fits the characteristics of ovaltis and
Biological Series, Vol. 15, No. 1
Ticks
137
•^ -;£^^-i*t-<5-c^<t.
Fig. 95. Ixodes ov(du.s, female.
p
M
136
Bnir.HANf VouNC Univehsity Science Bulletin
»v
w
Laboratory life cycle of Ixodes nipponensis ( reared on rabbits )
|
Generation |
|||||
|
and Stage |
Phase |
Period |
in Days |
Remarks |
|
|
F, Larva |
Feeding |
5 |
Larvae were placed on host on |
||
|
Fi Larva |
Postparasitic (Preinolting) |
30 |
5 July 1967. |
||
|
F, Nymph |
Feeding |
4 |
1 |
||
|
F, Nymph |
Postparasitic |
33 |
Only 2 nvmphs molted to . |
||
|
(Premolting) |
females. ( |
The rearing experiment extended from 5 July 1967 to October and the air-conditioned room tem- perature did not exceed 25 C.
1957:20-21, PL 31, 32; Saito, 1959b: 199-203,
1960a:,303-321, 1962a: 127-146, 1962b: 147-159,
1964:59-66; Saito and Ohara, 1961:1-32; Saito,
Ohara, and Unagami, 1960:323-329; Saito
et al, 1965:143-159; Asanuma, 1956a:91-96,
1957:234, 1965a: 113-116, Fig. 7. 8, 1965b:
398, Fig. 217; Ono, 1966:62-68. Ixodes frcquens Ogura and Takada, 1927:201-
202, PI. Ill; Schulze, 19.30:294-303, Fig. 10;
Kishida, 1930a: 3; Nakamura and Yajinia,
1937:146-148, PI. IV; Yajima, 1942:507-509,
Fig. 6; Itagaki, Noda, and Yamaguchi, 1944:
1-97, PI. 2,'l959:l-118, PI. 2. Ixodes carinattis Kishida, 19.30a :4-5; Nakamura
and Yajima, 1937:140; Keegan and Toshioka,
1957:26, PI. 41. Ixodes sp. 6 and sp. 52 Asanuma and Sekikavva,
1952:107-116, Fig. .3, 9, 1953:99-112, Fig. 11,
12; Asanuma, 1955:1240-1242; Ono, 1962a:24-
29, Fig. 15, 1962b: 155, 1966:62-68. ?Ixodes shinchikuensis Sugimoto, 1937:. 330-331,
PI. VI and IX. Ylxodes taiwanemls Sugimoto, 19.36 :336-.346, PI.
II, 1937a :331-333, PI. VII and VIII.
Discussion:
In the Japanese literature this tick has been reported as /. joponensis, I. jre(jticns, and some- times /. carinatus.
Ixodes ovatus was originally described by Neumann (1899) on the basis of material from Amur and Japan. The Japanese material includ- ed four Icmale specimens, one from a horse at Akita collected by Taguehi (No. 702), the others from a Lepus sp., a horse, and a dog at Saga collected by Yamaguchi (No. 70.3, 704, 705 re- spectively, in the Neumann Collection in the School of Veterinary Medicine in Toulouse). .According to NuttaJl and Warburton (1911), these Japanese specimens contained typical Ticinus,^ and subsequently Neumann ( 1904 ) re-
"MnnU (19(ii) slinwcd th,il ihosc' riciniis iNo. 702 ami 703) ncrc arUi.illy /. iifniilniliis. Iiiil the poncnsis Ktt.ink.-i and Saitti.
described ovatus based on two females from a horse and a dog at Saga (No. 704 and 705) as separate from the other two. Nuttall and War- burton (1911) cited Neumanns description ( 1904 ) , and in accordance with Dbnitz's opinion they noted that the differences between the two species were only varietal. It is apparent that Nuttall and Warburton did not see the type specimens. Therefore, they placed ovatus as a variety of ricinus and treated japo)iensu- as a valid species separate from ovatus.
Neumann's original description of /. japonen- sis ( 1904 ) did not include illustrations. Later, Nuttall and Warburton (1911) provided illus- trations (their Fig. 200 and 201) and a rede- scription ot jai)oneiisis based on Neumann's type specimen collected in the vicinity of Tokyo.
Pomerantzev (1950) considered ovatus as a valid species occurring in Japan, but this appears to be merely a citation from Neumann's original record.
Ogura and Takada (1927) described /. fre- (piens. but Keegan and Toshioka (1957) sug- iiested, "After a studv of hosts and localities, it seems clear that frequens is a synonym (of jajwnemis-ovatus). They also examined the holotvpe specimen of I. carinatus Kishida, 1930 and stated that "the holotvpe is ver\' much like an example of japonensis. although the basis capituli may be relatively wider than in that species." More recentlv Morel (1963) showed that /. jaf>oncmis. I. frecpiens, I. cariiwtus, I. shincliikuensis. and /. tahranensis were all syno- nyms of /. ovatus. The latter two species were described by Sugimoto on the basis of Taiwan specimens, and Morel (1963) expressed uncer- taint\' regarding their status, probablv becau.se of lack of specimens. .Although th(^ holotvpe specimen of carinattis was not available for this studv, the original illustration by Kishida ( 19.30a ) fits the characteristics of ovatus and
mipht iir niij;ht not iiulmlc / '"P"
# A 1%^^
F^m
»;».v
* 1^
^^^^^^^v
iBiUETR Biological Series, Vol. 15, No. 1
Ticks
137
' ')!*
/"^ <:£«^'•~^^x^«,
Fig. 95. Ixodes ovatus, female.
138
Bkioham Young University Science Bulletin
/O' ^-^^r^^Otf^o^
Fit:. 96. Ixodes onitim. male
Biological Series, Vol. 15, No. 1
Ticks
139
'/^^A<
Fig. 97. Ixodes ovatu.s-, iijmph.
140
Bhicham Young University Science Bulletin
0.1 m m
BuucKi
Biological Slries, V'ol. 15, No. 1
Ticks
141
supports Morel's opinion.
The only report of this species as ovatiis was given by Kishida (1930a), but he also reported ja))onensis on the same page. Asanuma and Sekikawa (1952, 1953) reported and described larval and nyniphal ticks parasitic on rodents, giving them temporarv numbers such as Ixodes sp. 6 or Ixodes sp. 52, because the immature stages had not been properly associated with adult fonns. Later Asanuma (1957) showed that these numbered ticks, Ixodes sp. 6 and Ixodes sp. 52, were larval and nvmphal forms of Ixodes japonensis (= /. ovatus) respectively. Ono (1962a, 1962b. 1966) reported immature ticks from rodents in Hokkaido and gave them the numerical designations used by Asanuma and Sekikawa.
DiAGXOSIS:
The dorsal aspect of the male of this species is similar to /. nipponensis and /. perstdcatus, but from a ventral view it may be easilv distin- guished from those two species in that coxae II and III are lacking spurs, and the posterior halves are covered with milky white, membra- nous, eavelike elevations ( = carinatus Kishida, 1930), and consequentlv the posterior margins are smoothly convex; coxa I has a short, blunt internal spur; the spiracular plate is elongate, the L/W ratio is more than 2.0 (approximately 2.3); the dentition of the hvpostome occurs on the apical half, the posterior end is on the same level as the suture between palpal articles II and III. In the female, the basis capituli has well- defined cornua and round porous areas; coxa I has a short, blunt internal spur; on coxa II, the spur is absent and the posterior margin is convex with the elevation similar to that of the male.
DlSTRIBlTIOX .\ND HoSTS ;
Morel ( 1963) states that this species occurs in oriental regions, including China, Formosa, Bunna, Nepal, Sikkim, and Japan, and that hosts of the adults arc large mammals. .'Kccording to Asanuma (1965a), adults are found on hares in great numbers, especially in spring, but imma- ture forms are not found on hares. Asanuma and Sekikawa (1953) and Saito (1959b) state that immature fomis are found on small rodents. Detailed infonnation on the distribution and hosts of this species in Japan is given in Appendix 2.
Biology:
Saito (1960a) studied the life cycle of Ixodes ovatus (as /. japonetviis) with laboratory animals as hosts (Angora rabbits and mice). Eggs hatched in .32-38 days at .30 C, and the
Ixodes ovatus
^ Collected and examined
O Collection records from literature
^nr Shirrbeshifc^fc^O/Kushir Is / -^vTlburi^idaka
Oshima I
ToyamaQ^-r-
^ . FukuiQj ^^»"°, A°jrK'KvSto^nianash
'Okayama"^'^ Hyogo
jKumamoto I Kagoshima
CigataOr OFukushima C'umma .■^_u.„.
^
Tochigi Tokyo Chiba Shizuoka -' Kanagawa
Map 29. Kno
di.stribiition of Ixodes ovatus.
preparasitic larval period was about 40 days. Feeding time of larvae varied according to tem- perature and was from 2-6 days in summer and from 7-10 days in late autumn on Angora rab- bits, and from 4-7 days on laboratory mice. The postparasitic period was 24-.32 days at 30 C, and newly hatched nymphs attached to the host 14 days later. Nymphs fed on the host for about the same period as, or one or two days longer than, larvae, and they molted into adults 30-.37 days after they dropped off the host. The pre- parasitic period for adults was 20-30 days, and the unfed males began mating immediately after they were put together with females on the host. Females fed on the host for 9-13 days in spring and autumn, 7-10 days in summer, and 12-16 days in winter. The engorged females laid eggs in .5-7 days in summer and in 10- L5 days in spring, and the oviposition period varied from 10-40 days, depending on the room temperature. Saito also observed the behavior of mating and oviposition. He stated that summer seemed to be an unfavorable time for feeding of adults in the laboratory.
Disease Relationship:
The relationship of this species to human dis-
142
BiiiciiAM VouNC Univebsitv Science Bulletin
ease has not been established. Saito ( 19621) ) reported the result of experimental transmission of Piisteurclld lularen.sis with /. idjxmciisi.s. The adults of 7. japonensus ( = /. ovatus), detached from hares dead of tularemia, were found to be infected and shown to be capable of transmitting P. tularcn.si.s to Angora rabbits. Attempt.s to demonstrate transovarian transmission were neg- ative.
Ixodes persulcatus Schulze (Fig. 99-102)
Ixodes persulcatus Schulze, 1930:294-303, Fig. 3, 5, 6, 8, 9. Morel, 1963:925-928"; Ono, 1966: 62-68, Fig. 1; Kitaoka and Saito, 1967:82.
Ixodes perstdcdtu.^ i)ersidcatus Schulze, 1930: 294-303, Fig. 3, 5, 6; Asanuma, 1947b :971, Fig. 2764, 1965a: 113-116, I965b:397, Fig. 215; Keegan and Toshioka, 1957:21-22 (in part).
Ixodes sp. 2 and sp. 54 Asanmiia and Sekikawa, 1952:107-116, 1953:99-112, Fig. 13 and 14; Asanuma, 1955:1240-1242; Ono, 1962a:24-29, Fig. 4-6, 8, 10, 12, 1,3, 1962b: 155, 1966:62-68, Fig. 1.
?Ixodes ricinus: Neumann, 1899:166 (in part), 1904:452, 1911:12-13; Nuttall and Warbur- ton, 1911:1.56, 1915:433; Ogura and Takada, 1927:199-201, PI. XI; Kishida, 1930a:2, 1936: 142; Nakamura and Yajima, 1937:142-145, PI. 11; Yajima, 1942:504-508, Fig. 5; Itagaki, Noda, and Yamaguchi. 1944:1-97, PI. 1, 1959: 1-118, PI. 1.
?Ixodes persulcatus ])ers-idcatus: Saito, 1958:70, 1959b: 199-20,3, 1960a:. 303-321, 1962a: 127-146, 1962b: 147-149; Saito and Ohara, 1961:l-.32; Saito, Ohara, and Unagami, 1960:32.3-329.
?Ixodes ricinus var. miyazakiensis Sugimoto, 1937:603-604, PI. 1.
Discussion:
This tick had been widely reported in the Japanese literature as Ixodes ricinus ( Linnaeus, 1785). Schulze (19.30), in his original descrip- tion of /. persidcatus, examined two female speci- mens from Hokkaido and stated that these speci- mens appeared to be conspeeifie with a small Amur specimen ( one of the type scries ) show- ing all the characteristics of /. persidcatus. He also stated that specimens described bv Ogura ;uid Takada (1927) ;is ricinus from Hokkaido apparently belonged to his new subspecies, I. p. persulcatus. Keegan and Toshioka (19.57) re- ported that all the material available at that time was reexamined and identified ;is Ixodes persid- catus j>rrstdcatus. Since then, the ]apanese popu- lation had been thought to include onl\' junstd- catus persulcatus. Later Asanuma (1 96.5a) ex- pressed doubt that onlv persidcatus persidcatus
occurred in Japan and suggested that there mav be two or three separate species contained in the Japanese population of the ricinus group. Asa- numa and Sekikawa ( 1952, 19.53) described im- mature ticks parasitic on small rodents and gave them temporary numbers, i.e., Ixodes sp. 2 for kirvae and Ixodes sp. .54 for nymphs. They pointed out that these numbered ticks were very similar to immature forms of ricinus, but avoided a definitive assignment. Ono (1962a) examined a large number of reared larvae of Ixodes persulcatus and considered that Ixodes sp. 2 and sp. 54 reported by Asanuma and Seki- kawa were the immature forms of Ixodes per- sulcatus.
Keegan and Toshioka (19.57) state that Ixodes ricinus var. niiiiazakicnsis Sugimoto, 19.37 appears to be persulcatus persulcatus. But in the light of current knowledge, the written descrip- tion and the illustrations are too poor for taxo- noinic discrimination, and this material could be l)er.sidcatus or nipponensis. Since the adults of persulcatus are very close to nipponensis and the specimens reported as ricinus by earlier Japa- nese workers are difficult to obtain, there are only a few reliable collection records of /. per- sulcatus.
Diagnosis:
This species is closest to /. nipponensis in the Japanese fauna and also to ricinus in other regions. Kitaoka and Saito ( 1967) gave a critical comparison of these three species and indicated that adults of persulcatus may be distinguished from nij)j)onensis bv the larger bodv size as well as by a combin;ition of characters used in the key. In addition, the following characters of immature stages given bv Kitaoka and Saito (1967) and Ono (1962a, i962b, 1966) are u.se- ful in separating the three species. In the larvae, the scutal setae are of medium length, while they are apparently shorter th;m the post- scutal setae in both nipjwnensis and ricinus; the livpostomc of the nvmph is blunt at the tip rather than pointed as in nipjwnensis: the post- scutal setae are not forked; the auriculae and spurs on the coxae are more distinct in persul- catus than in nipponensis both in the larval and the nvmph;il stages. See diagnosis of nip- jionensis.
Distribution and Hosts:
The tvpe material was collected at .\inur. Schulze mentions other material from Hokkaido which had been described ;is ricinus by Ogura and Takadii (1927). According to Pomerantzev (19.50), the distribution of this .species is from the Pacific Ocean to the Furopean p;irt of the
Biological Series, Vol. 15, No. 1
Ticks
143
/ okCXuiA,
Fig. 99. Ixodes persulcatu.s, It'inalf .
144
Bricham Younc University Science Bulletin
^.oAv^ujd^
Ki<;. 1(K). Ixodes persulcatus, male.
BioLocicAi. Sehies, Vol. 15, No. 1
Ticks
145
-^. 'TriAjiu'Ju-
Fic. 101. Ixodes persulcatus, nymph.
146
Bricham Younc; IInivehsitv Science Bulletin
j^m^Ti/)
Fit:. 102. Ixodes persulcntus. Iar\;
Biological Series, Vol. 15, No. I
Ticks
147
USSR, and al.so in northern Germany. The dis- tribution in Japan i.s given in Appendix 2. The only Korean record is from a horse at Kankyo- lioku, now known as Hamgvong-pukto, by Ita- gaki et al. (1944, 1959). Kishida (1936) men- tions that /. ricintis was recorded from North Korea bv Akasawa in 192S. We have been un- able to confinn this record or to find any paper by Akasawa written in 1928. Asanuma and Sekikawa (1953) listed the hosts of ricimis and j)ersulcatus\ citing the records by Kishida ( 193()a ) and Nakamura and Yajima ( 1937 ) . They are human, cow, horse, wild rabbit, deer, dog, fox. leopard, weasel, martin, hedgehog, sheep, goat, and rat. Pomerantzev (1950) stated that the main hosts of adults in the USSR are wild ungulates such as deer, elk, roebuck, and the Siberian stag. Hosts of larvae and nymphs are small mammals ( Rodentia, Insectivora, Car- nivora) and birds. Nymphs are also parasitic on the Amur hedgehog, dormouse, scjuirrel, hare, and wood-hen. Both adults and nymphs attack man.
hodes persulcanjs and/ nanus
1 ^ Collected and examined
2 Q Collection records from
literature
3 3 Combination of 1 and 2
4 A Collection records of
/ ncinus
5 <$ Combination of 2 and 4
6 O Combination of 1, 2 and 4
" StiiribestiiftnCy''i^'>'^"*"° ^^ThIa^,-^'T°''0':l'i
OshimaT^
Aomofi I •/ ®'wate
kMiyagi
Yama9ata( SadoC^ 1} ^Niigata^-r *A Fukusfiima
— ^ Naganoflt)^ J.Saitama
-O Kyoto . ? "Jr<j2ii»« Tottori —>llS'^"tV\ fQ.^® Tokyo Shimanejn^Hyogd^t^_X '/K'iO' '
Yamanaslii , Karwgawa
:Mie"~^ " /
,Ehime5%'f*\-^^
Kuniamoto Miyazaki
Map .30. Known distriUntion of /ayx/cs prrsulrcilus and /. ricniuv.
Biology:
The biology of /. pcrsulcutm has been
studied by Russian workers who were concerned with the transmission of spring-summer encepha- litis to man. The life cycle was studied in nature by Serdukova, and the infomiation is given by Pomerantzev ( 1950 ) . The length of the develop- mental cycle depends up(m environmental fac- tors and is said to take three years in the boreal taiga in the USSR in warmer regions; and, in the presence of abundant host animals for all the three active stages, the tick develops in a re- duced two-year cycle. Ixodes j)ersulc(itus is a typical forest inhabitant and is most abundant in mixed deciduous-coniferous forests.
The seasonal occurrence of persulcatus in Niigata Prefecture, Japan, was studied by Saito (1959b) through collections from wild animals or from vegetation using the flagging method. The following biological information was iiuoted from Saito's extensive studies, but as noted in the discussion, this might be applicable to per- sulcatus. provided Saito's material definitely did not include nipponensls. Both immatures and adults were found together on hares. The adults were generally found active on the host through- out the year, showing a slight increase in num- bers in spring and late autumn. Both nymphs and larvae were most abundant on the host from April through July. Larvae were not collected on hares in winter and midsummer. The imma- ture fomis were also rare on Apodemtis specio- stis sadoensis in summer. Using the flagging method, all stages were collected from vegeta- tion, and the peak abundance was from summer to early autumn. Saito ( 1960a ) studied the labo- ratory life cycle and mating behavior, using chiefly the Angora rabbit as the host animal. Females fed for 13-18 days in spring and autumn and 7-14 days in summer. He found that if fe- males leave the host before becoming fully en- gorged they weigh less than 70 mg, and they rarely feed on another host. Females laid eggs 5-8 days after engorgement in summer and 9-14 days in spring. Oviposition lasted 10-14 days un- der various environmental conditions, and larvae hatched out in 30-36 days at 30 C. For larvae the preparasitic period was approximately 30 days, and they fed for periods of from 3-13 days. Nymphs emerged in 17-19 days after larvae had dropped off the host. Ten days later they fed on the host for periods which varied from 4-15 days. Adults emerged in 25-30 days at 30 C. The minimum time for the total life cycle in the laboratory was approximately 4.5 months.
An interesting peculiarity of the behavior of persulcatus was observed by Balashov (1954). The dropping of engorged female ticks from the host coincides with the time when the cows were
148
BiuGHAM Young University Science Bulletin
in the pasture. If cows were pastured during tile davtinie, tlien tlie ticks dropped during the daytime, hut if cows were pastured during the night, then the ticks dropped at night.
Disease Relationship:
Ixodes persulcaius transmits the virus of spring-summer encephalitis to man in the USSR. Ponierantzev (1950) states that the causal agent ot cattle piroplasniosis can he transmitted experimentally. Saito ( 1962b) studied the trans- mission of PasteurelUi tularensis by /. perstilca- tus and found that adult female specimens of persulcatus, collected from wild hares which had died of tularemia, were infected with P. tularen- sis and that nymphs and larvae that were ex- perimentally infected with the causal agent of tularemia transmitted it from host to host. He also demonstrated transstadial transmission, both from larvae to nymphs and from nymphs to adults, but he was unable to demonstrate trans- ovarian transmission. The role of /. persulcatus as a vector of P. tukirensis from hares to man has not been established.
Ixodes philipi Keirans and Kohls (Fig. 103)
Ixodes philipi Keirans and Kohls, 1970:725-72e, Fig. 1-8.
Dl.SCUSSION:
The description of this species was based on two partially replete females collected by H. E. McClure in 1951 at Sanganjima, Iwate Prefec- ture, Japan. Keirans and Kohls (1970) (juoted from a personal communication from Dr. Mc- Clure in which he described the collecting area as ". . . high islands of volcanic ash covered with pin(> and deciduous forests. In the loose soil beneath the trees the Streaked Sheanvater, Pttffimis leucomelas, and the Madeiran Storm Petrel, Oceanodroma castro, dig nesting tunnels about arm-length deep." Ticks were collected during an avian blood survev but it isn't certain which of the two species of birds was the host from which the ticks were collected. Japanese ornithologists now refer to the Streaked Shear- waster as Calonectris leucomelas and the Ma- deiran Storm Petrel is sometimes called the Ma- deiran Fork-tailed Petrel. Keirans and Kohls (1970) indicated that the female characters ful- fill the criteria for placement of /. philipi in the subgenus Scaphixodes Schul/.e, 1941 but because of the lack of males and immature fomis thev are hesitant to do so. More extensive tick collec- tions from sea birds are essential for ecological and taxonomic studies.
Diagnosis:
We (juote directly from Keirans and Kohls (1970): "Ixodes ))hilipi n. sp. superficially re- sembles /. kohlsi described by Arthur (1955) from Eudtjpttda minor or from its nest, and keys to /. kohlsi in Roberts (1960), However, /. philipi is much more hirsute dorsally and has a less elongate scutum. Ventrallv /. ))hilipi has a smaller, less distinct external spur on coxa IV, and spur on trochanter I\' obsolete."
Of the Japanese Hcks associated with sea birds I. philipi most closely resembles /. signatus and kevs out to that species. However, females may be distinguished from /. sigmitus by the following diagnostic characters: Dorsum pos- terior to scutum uniformly covered with white hairs; coxae I-III anned with a more pronounced external spur; anal groove markediv constricted and subparallcl posterior to anus giving "key- hole" shape; basis capituli ventrally with broadly rounded posterior margin, with straight lateral margin anterior to auriculae; auriculae broadly triangular or nearlv right-angled ridges; porous areas well separated by about half the width of one of them; hypostome untoothed at base, about
M.AP 31. KniiNMi ilistrihiitioii i>t Ixo(lc.\ philipi.
Biological Series, Vol. 15, No. 1
TlCK.S
149
Fig. 10.3. Ixodes philijn, female. ( .After Kierans and Kohls 1970)
150
BHif;iiAM VouNG University Science Bulletin
one fourth as long as the wliolc length, with more prominent teeth on inner file.
Distribution and Hosts:
Known only from Ptiffiniis Icucomchis (Tem- minch, 1S35) or Oceanodwiiia castro (Harcourt, 1851), Sanganjima Island, Iwate Prefecture, Japan.
Biology; Unknown.
Disease Relationship : Unknown.
Ixodes signatiis Binila (Fig. 104-107)
Ixodes signatus Rirula, 1895:. 357-358, PI. 1, Fig. 10-13; Nuttali and Warburton, 1911:261-264; Kishida, 19.30a:2; Znmpt, 1952:18; Asanuma, Okubo, and Fnkuda, 1955:85-86; Asanuma and Fukuda, 1957:147-159; Keegan and To- shioka, 1957:22, PI. .35 and 36; Asanuma, 1965a: 11.3-116, 1965b:398, Fig. 216.
Ixodes parvirostris Neumiinn, 1901:284.
Ixodes eudijptidis var. signata: Neumann, 1904: 451.
Ixodes eudi/ptidis signatus: Neumann, 1911:21.
Discussion:
This tick was first reported from Japan by Neumann ( 1901 ) as Ixodes parvirostris, which he described from eight female specimens taken from the connorant, PIxdacrocorax pelagicus, on Yezo, Japan. Yezo is the archaic term for Hok- kaido. Nuttali and Warburton made parviros- tris a synonym of sigiiatus in 1911. Kishida (1930a) and Znmpt (1952) cited Neumann's collection record, but gave no new information. Asanuma, Okubo, and Fukuda (1955) reported an infestation of the black-tailed gull. Lams crassirostris, by this tick at Kabujima, Aomori Prefecture, Honshu. Additional specimens from the same locality were collected by 406th Medi- cal Laboratory personnel.
Diagnosis:
This species is distinct from other Ixodes sp. and may be easily identified using the key. Description of the male by Keegan and Toshi- oka (1957) is based on material collected from Lariis crassirostris at Kabujima, Japan:
Body widens abruptlv from scapular region; with nearly parallel lateral margins; posterior body margin bluntly rounded; lat- eral groove distinct, (^apituhii)i relatively short and broad, but without the lateral
saliences present in the female; cornua lack- ing; palp article III slightly longer than Article II, both articles II and III relatively shorter and broader than in the female. liijpostomc about three-fourths as long as palpi, relatively shorter than in female; den- ticles indistinct. Scutum: Cervical grooves distinct, nearlv straight, diverging; two longitudinal, nearly straight, slightly diverg- ing grooves posterior to cervical grooves. Between these is a single, indistinct, longi- tudinal groove with faint cross-bars which extend to the paired longitudinal grooves. A dorso-lateral sclerotized patch on either side of the body at a level between coxae II and III. Posterior margin of scutum ir- regular in outline, and tenninating in a point in most specimens examined. The scutimi does not cover the entire dorsal sur- face within the lateral groove. This is par- ticularly true on the posterior fourth of the bodv. Palpi more noticeably haired than in the female; especially distinctive are a group of short, stout setae on the lateral, dorsal aspect of articles II and III; terminal setae also are more conspicuous. Legs and coxae as in female. Coxal spurs on legs III and IV may be relatively larger than in female. Spiracular plate oval rather than circular; macula displaced anteriorly; gob- lets as in female. Genital aperture between coxae II. Genital grooves start to converge at level of anterior margin of anal groove. Median and adanal plates not distinct; a slight trace of sclerotization and pigmenta- tion in the areas usually occupied by the adanal shields, and a more extensive pig- mented area in the usual location of the ventral shield; this pigmented area has ir- regular lateral and posterior margins. Aiud groove rounded anterior to anus; the arms of the groove are curved throughout, and converge posterior to the anus; distance be- tween these arms is greatest a short dis- tance posterior to the anus.
Distribution and Hosts:
Znmpt (1952) has reviewed the distribution of signatus and other species of ticks infesting sea birds. This species has been collected on only a few occasions in other areas. It has been found on the connorant on Hokkaido, Japan, the Aleutian Islands, and from t\vo locations in California. It was taken once from a seal in the Aleutian Islands and on several occasions from the black-tailed gull in its nesting areas at Kabu- jima, Aomori Prefecture, Japan.
Biological Series, Vol. 15, No. 1
Ticks
151
i JHIBATA
Fig. 1()4. Ixodea signatus, female.
152
BiiiGHAM VouNC University Science Bulletin
S.SHIBATA
Fig. 105. Ixodes ni^iuittia, male.
Biological Series, Vol. 15, No. 1
Ticks
153
RuucKi Fic. 106. Ixodes signatus, nymph.
1.54
Bhigham Young Univebsity Science Bulletin
W t'ia'iin\un\ii.
Fio. 107. Ixodes sisiuitiis, larva.
Biological Series, Vol. 15, No. 1
Ticks
155
|
Ixodes signaws |
1 ' |
—I |
|
^ Collected and examined |
' / |
|
|
O Collection records from |
ft / V?\ Abashin |
|
|
literature O Combmation of above |
||
|
i 0 ff" |
^ T^Kabujima |
|
|
■ ^ V^ |
W J \ |
|
|
= ^^'^.^ |
'h\J |
|
|
^JlXr |
||
|
(i |
^^^lXj |
|
|
<^^ |
||
|
■^ j;^^^^^ |
■~-^X^M^ J Kanagawa * s |
|
|
, |
||
|
■• |
j{ |
|
|
w ( |
4^ |
|
|
*0O ? "» -?"■"' |
l3i |
|
|
V>^ |
||
|
^^ |
Map 32. Kntwn distribution of Ixodes si^uatus.
Biology:
Interesting information on the biology of thi.s specie.s has been reported by Asanuma and Fukuda ( 1957 ) . In each of their field observa- tions they found the ticks parasitic on the host only in May. In 1951, larvae constituted the dominant stage on the host; in 1955 it was nviiiphs; in 1956, females. No males were found on hosts; and, in addition, no males which had fed on blood were found even in crevices of rocks near the birds' nests. They speculated that the life cycle of this species at Kabujima likely recjuires 3 v<?iirs, because the host birds stay and breed there during a limited period of approxi- matelv 5 months, from April to early August. Field obser\ations and the duration of each stage in the laboratory strengthen his theory. They also found males copulating with newly emerged females in the crevices of the rocks.
Disease Relationship:
The relationship of /. sig,nafus to human dis- ease is unknown. Asanuma and Fukuda ( 1957 ) stated that they found a dead black-tailed gull with a heavv infestation of /. sif^natus, but the cause of death was not established.
Ixodes simplex siinj)Iex Neumann (Fig. 108-109)
Ixodes simplex Neumann, 1906:197-198; Kishida, 1930a: 3.
Ixodes simplex simplex Neumann, 1906; Arthur, 1956:180, Fig. 24-26, 28-30, 33-43; Keegan and Toshioka, 1957:23-24, PI. 37; Asanuma, 1965a: 113- 116.
Discussion:
Arthur (1956) divided Ixodes simplex into two subspecies, /. simplex simplex and /. simplex africanus. The latter has been found in the Rift Valley, Kenya. Kishida (1930a) reported occurrence of this tick on bats in Nagano Pre- fecture, Honshu, Japan. He did not give the specific names of the bats but gave the common names in Japanese. They are Kikugashira- Koomori and Yama-Koomori, now known by the specific names Rhinoloplnis fernim-equimim and Nyct(diis lasiopterus.
Arthur (1956) examined two collections of simplex from Japan. Both of these are in the collection of the Rocky Mountain Laboratory. One of these, RML No. 30943, consisted of one female and three larvae from Mijotis macrodac- tijlus taken 30 September 1952, near Kameaka, Japan. The other, RML No. 22366, consisted of one nymph from Miniopteris schreihersi, Kama- kura, 24 December 1945. There is a tjuestion concerning the location of "Kameaka," given as the locality for collection RML No. .30943. There seems to be no "Kameaka" on Honshu, although there is a Kameoka located in Kyota Prefecture. Keegan and Toshioka ( 1957 ) examined one fe- male specimen loaned by Dr. Asanuma which was taken from a bat in Shiga Prefecture.
Diagnosis:
The female is easily distinguishable from the only other bat tick, /. vespertilionis, in that the legs nomially are long; the anal groove is point- ed in front of the anus, with posterior arms di- verging; palps relatively short, broad and blunt, broadest near distal end, narrow proximally; without comua; auriculae as small, blunt ridges. This species is also distinct from other Ixodes spp. in that all coxae are without internal or ex- ternal spurs. Males are unknowTi.
DlSTIUBUTION AND HoSTS:
As reported by Arthur (1956), this tick has been taken from bats in Palestine, Greece, France, Japan, China, and Africa. The distribu- tion and hosts in Japan are given in Appendix 2. Larvae and nymphs are also found on bats.
156
Bricham Young University Science Bulletin
RTSUTSUMI
l-'ic. 1()8. Ixodes ximplcx Kimplrx, female.
Biological Series, Vol. 15, No. 1
Ticks
157
Fic;. 1()9. Ixodes .simplex simplex, nymph.
158
BiuGHAM Young University Sc:ience Bulletin-
Map 33. Known distribution of Ixodes simplex sim- plex.
Biology :
Nothing is known concerning the biology. Males probably do not feed, or they may feed very rapidly and then quickly hide themselves.
Disease Relationship: Unknown.
Ixodes tamtki Saito (Fig. 110-111) Ixodes tanuki Saito, 1964:59-66, Fig. 1-9.
Discussion:
Ticks taken from a badger, Nyctereutes pro- cijonoides viverrinus, m Niigata Prefecture, Ja- pan, were described liy Saito (1964). He col- lected one male and three females in 1959 and an additional female in 1963. The specific name "tanuki" was derived from the Japanese common name for the host. Since the specimens were not available for study, the following infonnation is taken from Saito's paper.
Diagnosis:
Saito states thai /. taiuiki mav be easilv dif-
ferentiated from other Japanese ix(Klids by the following characteristics: "In /. tanuki, with a long sharp internal spur on coxa I, external spur is absent on coxa I and II each. These morpho- logical features are not observed in the other ticks. Ixodes ozarkus Cooley, 1944, of North America, is thought to be resembling /. tanuki, but measurements of all parts of ozarkus are generally larger than those of tanuki. The for- mer has an external spur on coxa I and II each, but these are lacking in the latter. The spiracu- lar plate of ozarkus is elongated oval in shape, but oval in /. tanuki."
Distribution and Hosts:
Distribution and hosts of the species are at present limited to the badger in Niigata Prefec- ture, and there are no reports from other areas. Dogs have been suggested as possible hosts.
|
Ixodes tanuki . , ■? 1 |
— ) — |
|
0 Collected and examined * // |
|
|
O Collection records from tf^ *9 literature £^ ^ ^tsC^ |
|
|
3 Combination of above ft I/ ) -^^Sv |
|
|
tf / ' j?^z^*^'*r?^ "' |
|
|
^A / C^l^^^i?^ |
|
|
'iy* A >/ XJ'*-' ■ X*^ |
|
|
' / 'Y^^ ' |
|
|
■ ^ y^ W\ |
|
|
= t,^ ^^ 'hM |
|
|
===--=^""''^ ^JU\ |
|
|
r-^ ^ONiigatal |
|
|
yfCrw-v/v vV |
|
|
^ _^^ -^^ |
|
|
r:^^C^w^-i.'^^r |
|
|
- ^ s^^^^Mr'^ '' |
|
|
■mm^ |
|
|
'■%/ |
Xx c |
|
^^B 0 1«0 JW..-T, |
\y |
|
• |
^ |
Map 34. Known distriliiitioM of Ixoilcs tanuki.
Biology: Unknown.
Disease Relationship: Unknown.
Biological Series, Vol. 15, No. 1
Ticks
159
1 .0 mm
A
0.2 mm
0.5 mm
Fic. 110. Ixodes tdiiuki. ffiiiale.
160
Bhicham Vounc University Science Bulletin
1 .0 mm
Fic;. 111. Ixodes tanuki, male.
Biological Series, Vol. 15, No. 1
TiCK.s
161
Ixodes turdtts Nakatsiiji (Fig. 112-114) Ixodes tardus Nakat.suji. 1942:292-294, Fig. 4; A.sanuma and Ko.saka, 1955: 192-194, Fig. 1-4; Asanuma and Nakagawa, 1955:549; Kee- gan and Toshioka, 1957^24, PI. .38; Asanuma, 1965a: 113-1 16.
Ixodes sp. 1 Asanuma and Sekikawa, 1952:107-
116. Discu.ssiON:
The description of /. tiinltis Nakatsuji, 1942 was based on seven females taken from the thrush, Turdus celacnopus ceJaetwpus, on Ha- chijo Island. .\dditionaI infomiation concerning distribution, hosts, and morphology of the spe- cies was given by Asanuma and Kosaka ( 1955 ) .
Diagnosis:
Asanuma and Kosaka ( 1955 ) commented on the close similarity between I. turdus and I. brunneus Koch. 1844 and were of the opinion that the following characters may serve to dis- tinguish the two species: the scutum is large, long and rhombic, with a fairly distinct lateral angle at the widest part, the postlateral margin is slightly concave (distinct from brunneus in this); the auriculae are rectangular, well de- veloped, and project externally ( the tip is visible from the dorsal side); the hypostome is 4/4; the anal groove converges posteriorly; the coxae have well-developed external spurs.
Distribution and Hosts:
This species has been collected only in Japan. Asanuma and Kosaka ( 1955 ) have collected this species from a variety of birds, and on a few occasions from Apodemus speciosus. The majori- t\' of these collections were made in the Izu- Schichito Islands ( Seven Islands of Izu ) soutli of Tokvo Bav, and other specimens were taken in Chiba, .Shizuoka, and Tochigi prefectures, all on Honshu. Hosts listed by them are given in .Appendix 2.
Biology:
Unknown. Disease Relationship:
Unknown. Remarks:
During the course of this studv, material in
the 406th Medical Laboratory Collection (No.
148) was again sent to Mr. G. M. Kohls, who
gave the following opinion:
Very similar to /. brunneus Koch; see
Cooley and Kohls 1945, p. 205. Differs
from brunneus in characters of the scutum,
the punctations being more numerous in
the Japanese specimen, the median posterior
M.\p 35. Known distribution of Ixodes turdus.
area is flattened rather than convex, and the hairs are not quite as heavy as in brunneus. Quite likely this is Nakatsuji's (1942) species and if so the description is poor and the figures are inaccurate, espe- cially with regard to hypostome, auriculae, and the comua. I should like to mention too that the 2 nymphs accompanying the fe- males are very similar to those of /. cali- f amicus Banks, which is based on nymphs only, and which is definitely brunneus (Cooley and Kohls, 1945:215).' Since validity of this taxon remains in (juestion, the name /. turdus used in this paper is tenta- tive.
Ixodes iiriac White Ixodes uriae White, 1852:10; Keegan and Toshi- oka, 1957:.30; Asanuma 1961:181, 1965a: 116. Ixodes putus: Kishida, 19.30a: 2. Discussion:
Keegan and Toshioka (1957) listed /. uriae in the section, Questionable Distribution Rec- ords. As noted by them, I. fuitus. which has been synonymizcd by Cooley and Kohls (1945) under uriae. is recorded by Kishida (1930a) as
162
Hni<;nAM VouNC University Science Bulletin
/«^ Jy£fc«.<-ai»_
I'"k:. 112. Ixodes turdus; feinak-.
Biological Series, Vol. 15, No. 1
Ticks
163
Tn. 7nu^i-Ji4^
Fic. ll'J. Ixoilen turdu.s. nymph.
164
Bricham Voung University Science Bulletin
Ruuchi
Vic 111. /.vfi(/cv tunliis. larva.
Biological Series, Vol. 15, No. 1
Ticks
165
occurring on a bat, Ntjctahis sp., in Niigata and Nagano prefectures. Kishida's report of putus appears to be highly doubtful in view of the fact that I. iirUic is primarily a parasite of marine birds. Keegan and Toshioka ( 1957 ) expressed doubt about his identification.
Neumann (1902) created the genus Cera- tixodes for putus on the basis of the structure of tlie capitulum and the peculiar character of Haller's organ. Since then, the genus Ceratixodes has been either suppressed or used, depending on the workers. Zumpt ( 1952), having discussed the status of the genus, considered that the characteristics of iiriac were sufficient to sepa- rate it from Ixodes. However, the taxonomic problem related to Ceratixodes does not appear to be settled; moreover, Japanese specimens were not available for examination. Therefore, the species is here treated as belonging to the genus Ixodes, .sen.m lata.
Illustrations of this species were included in papers by Nuttall and Warburton (1911) and Pomerantzev (1950). The following informa- tion was taken from the literature.
Dl\gxosis:
This "puffin-tick" has very unique character- istics. The size is very large in both sexes. The male has five distinct tufts of long hairs at the posterior margin of the body; the 3rd palpal seg- ment is conicallv shaped. In the female the body is covered thicklv with white hairs; the 3rd pal- pal segment is much broader and shorter than the 2nd, and the suture between them is poorly defined, the 1st palpal segment is dorsally prominent.
Distribution and Hosts:
The tick has a wide geographical distribu- tion. The recorded localities for this species comprise many islands or seashores facing oceans encircling both poles. The species is therefore called a "bipolarlv distributed" tick.
Zumpt ( 1952 ) discussed the distribution, cit- ing the opinions expressed by Schulze ( 19.38 ) . Detailed localities for the tick are given bv Nuttall and Warburton (1911), Schulze (19.38), f:oolcy and Kohls (1945), and Zumpt (1952).
The hosts are mainlv sea birds inhabiting the areas mentioned above. They are gulls, al- batrosses, petrels, shearwaters, and penguins. Pomerantzev (1950) and Nuttall and Warbur- ton ( 1911 ) state that it occasionally attacks man. The sole reliable record in the Japanese litera- ture is Asanuma's report (1961, 1965a) which contains the record of a nymph and a larva from a puffin. Ltinda cirrhata. on Hokkaido.
Biology:
This species is said to be a three-host tick. Males do not feed and are assumed to copulate with the females after they have dropped off from the host. Zumpt (1952) considers that the wide distribution for this species can be at- tributed to the sticky eggs, which have a long incubation period and can be easily transported from one place to another on the feet of birds. Other biological infonnation can be obtained from Nuttalfand Warburton (1911).
Disease Relationship: Unknown.
Ixodes vespertilionis C. L. Koch
(Fig. 115-118)
Ixodes vespertilionis C. L. Koch, 1844:217-2.30, Fig. 9; Arthur, 1956:180-184, Fig. 1-13; Hoog- straal, 1956:567-573, PL 67;' Keegan and Toshioka, 1957:25, PI. 39-40; Asanuma, 1965a: 113-116, 1965b:399, Fig. 221.
Discussion:
Arthur (1956) and Hoogstraal (1956) both refer to a single collection (RML No. .32112) of /. vespertilionis from RhinoIoj)hus ferrum- eqtiinum Sawada, "Myiagi," Honshu, 13 May 1952 (record was given in personal correspon- dence from Kohls to Hoogstraal). The error in spelling (should be "Miyagi") is perpetuated in Arthur (1956) and Hoogstraal (1956). Kee- gan and Toshioka (1957) examined this single female specimen and gave several records of vespertilionis from bats collected on Honshu and Shikoku. Specimens from Hokkaido were borrowed from Mr. K. Hattori, and other study material was collected in Iwate Prefecture and in Korea bv 406th Medical Laboratory personnel.
Diagnosis:
This bat tick is very distinctive and may be easily distinguished from other Ixodes spp. The legs are thin, much longer than the body, giving a "spidery" effect; all coxae lack spurs; the anal groove is round in front of the anus and the pos- terior arms are nearly straight or slightly con- verging. In the male, the palps are clavate and have numerous long setae; the hypostome is pointed at the tip with faint indications of small, scalelike teeth. In the female, the scutum is nar- row and much longer than wide, with the widest point just behind the middle (in Nuttall and Warburton, 1911, broadest in the middle; in Pomerantzev, 1950, widest in front of the mid- dle), the basis capituli is dorsally triangular and has a flange externally directed at the postero-
166
Bbicham Vounc University Science Bulletin
jy/f///y.l yx^
Fig. 1 15. Ixodes tcspcriilioni^, lemale.
Biological Series, Vol. 15, No. 1
Ticks
167
Fig. 1 16. Ixodes vexpertilionui, male.
168
Bricham Young Univehsity Science Bulletin
1.0 mm
Fk;. 117. Ixdilcs vc\i)crtilioru.s. nymph.
Biological Series, Vol. 15, No. 1
TiCK.S
169
■^.77U*<r^
0 2 mm
Fir.. 118. Ixodes vesjierlilionis, larva
170
Hricham Young University Science Bulletin
external juncture; the liypostomc is lanceolate and has long denticles.
DiSTIUBUTION AND HOSTS:
The distribution and hosts of this species in areas other tlian Japan are presented by Arthur (1956) and Iloogstraal (1956). This species oc- curs on a variety of bat hosts in Europe, Africa, the Near East, and Japan. Russian workers re- port the tick from Russia, northern Iran, and western Siberia. Hosts from which Ixodes ves- jwrtilionis has been taken in japan are given in Appendix 2.
Map .36. Known distriliiitid
)f Ixodes
spvrtiUou
Biology:
Nothing is known of the biology of the Japa- nese population. Excellent reviews of biology, morphology, etc., are given by Arthur (1956) and Iloogstraal (1956): "No males have been found on bats, but thev wander over the walls of the caves, partieularlv where the rocks arc creviced. Unfertilized females and unfed im- mature forms occupy similar niches. It seems likely that the fully fed stages secrete them- selves between the stones of the caves to digest blood meals" (Arthur 1956).
Arthur's (1956a) comparison of the data from Switzerland and from Macedonia leads him to believe that, because there is a high reasonable catch of partiallv and full en- gorged ticks between October and January and a number of unfed nymphs and females during the summer, feeding is accomplished mainly during the winter months. This pic- ture, possibly modified bv the host's sea- sonal breeding cvclc and activity, re(|uires further observation. Nuttall and Warburton (I91I) postulated that males may either feed very rapidly and then leave the host, or that they may not feed at all. Neumann (I9I6) believed that the various degrees of engorgement in which male specimens are found might not necessarily prove that males do feed but rather mav be an indica- tion of degree of nvmphal feeding. This conclusion is based on the atrophy of the male hypostome in comparison with its ro- bust development in females and in imma- ture stages (Hoogstraal, 1956).
Disease Relationship:
Unknown.
Genus Rhiplcephalus C. L. Koch. 1844
Usuallv ornate. Palpi short. BasLs capituli usually hexagonal dorsallv. Eyes and festoons present. The males with a pair of adanal and accessory shields. Coxa T bifid. Spiracular plate comma-shaped.
Rhipiccphahts sanguineus Latreille, 1806 (Fig. 119-122)
Ixodes sanguineus Latreille, 1806:157
Rliipiccplialus sanguinctts: Koch. 1844:2.3.S-2.39; Sugimoto, 1937b:610-612; Keegan and To- shioka, 1957:26-27, pi. 42.
Discussion:
Although this tick, known as the "kennel tick," "brown dog tick," or "tropical brown dog tick," is one of the most widely distributed ixo- did ticks, present infonnation indicates that it is very rare, at least in Japan, Korea, and the Ryukyu Islands. Sugimoto ( 1937b) reported that R. sanguineus had been taken in Kumamoto Prefecture, Kvushu, but it has not been found b\- other collectors in that area.
During the course of this stud\-, threi' lots were examined and identified as R- sanguineus. judging from the 406th Medical Laboratory col- lec-tion n>cords, it is difficult to detennine whether this species has been established in
Biological Series, Vol. 15, No. 1
Ticks
171
Fig. 119. Rhipucpluihi.s sanguineus, female.
172
Brigham Young Univebsitv Science Bulletin
Sonobe
iMG. 120. Rhipiiciihcilus sunj^uiiinis, male
Bi()LO(;icAL Sehies, Vol. 15, No. 1
Ticks
173
TAmIo
Fig. 121. RhipirrphnJtis sdUf^tiiurtis. iivmpli.
174
Bhigham Young University Science Bulletin
-A^<U
Fig. 122. Rltipiccphalus- suni^uinvus, lana.
Biological Series, Vol. 15, \o. 1
TicK.s
175
Japan. However, it is readily siippcsed that it inav have been introduced into Japan on pets of U.S. Forces personnel or in household effects.
Feldniann-Muhsani ( 1952) has suggested that the tick commonlv known as R. sangttineus is actually not one species, but two. She pro- posed the specific name secundus for the new species, which is distinguished from sanguineus s. str. by the nature of the female genital aper- ture, and in immature stages by the shape of the capitulum. This distinction has not been fullv accepted bv Hoogstraal (1956).
DlAGNO-SIS :
Since there is only one species of the genus Rhipiccpltalus that occurs in this area, it is easil\- identified.
Distribution and Hosts:
Leeson ( 1951 ) did not include Japan, Korea, and the R^■ukvu Islands as part of the distribu- tional area of this species, but the special map published by the American Geographic Societ}' ( 1954 ) shows Korea and the Rvukyu Islands as areas where sanguineus occurs. Anastos (1950) gave its distribution in the East Indies, and Kohls (1950) discussed its occurrence in the Philippines. As reported by Anastos (19.50) and Hoogstraal (19.56), sanguineus is primarily a parasite of dogs but also attacks a great variety of domestic and wild animals. It has been taken from l)irds as well as mammals, and in some areas has been known to attack man.
Biology:
Hoogstraal ( 1956 ) has reviewed the litera- ture on the biology and medical importance of this tick. According to Hoogstraal's review, it is said to be a three-host tick, and the life cycle mav be completed in as little as 6.3 days under favorable conditions. If conditions are unfavor- able, it may be prolonged for many months. "In Nuttall's laboratorv experiments, larvae fed on dogs and rabbits, nvniphs on dogs, jackals, and hedgehogs, and adults on dogs and jackals" (Hoogstraal, 1956). He also cited several reports on parasites and predators of this tick. Hunter- elhis hookcri Howard, 1907 ( =IxocUphagus cau- curtci du Buvsson, 1912), a wasp, has been oc- casionallv reported as a parasite of R. sanguineus in .\frica, Brazil, USA, and other areas of the world. The eggs of this wasp are laid in the body of the nvmph and the developing hymenopter- ous larvae feed on the contents of the engorged nvmph and pupate in the body. A predaceous spider, Teutena triangulosa Wick, has been ob- served feeding on both immature and adult forms
|
' 1 ■ ' •- ,- . . , Rhipicephalus sanguineus , ^ 0 / |
T |
|
|
^ Collected and examined // |
||
|
Q Collection records from ,/*CPa 11 literature £? // ^fst\ (^ Combination of above A // \C\)^ |
||
|
Okinawa ^T ^ ^^S^^y\ / |
||
|
/\JjA^ ^ TOLrj^-J^^ > J^J^ Kanagawa |
||
|
**' «^33l Kumamoto |
4i |
|
|
.J . ,^ ,^.„ |
■ n^ |
|
|
■ •' |
\f^ |
|
Mai' 37. Known distribution of Rhijnceplmlu.s san- guineus.
of this kennel tick in a Corsican house (Sautet, 19.'36).
Disease Relationship:
The medical importance of this tick has been thoroughly reviewed by Hoogstraal ( 1956). This species is known to transmit boutonneuse fever, Indian tick typhus, and Rocky Mountain spotted fever in areas where these diseases occur. In addition, it is of importance to veterinary medi- cine as a vector of canine rickettsiosis and ca- nine piroplasmosis. The relapsing fever spirocha- ete, Borrelia theileri, is transmitted to domestic ruminants by R. sanguineus. Opinions differ re- garding this tick as a parasite of man.
DOUBTFUL RECORDS
Amhhjomma cordijerum Neumann, 1899:218-
219; Kishida, 1930b: 134.
Keegan and Toshioka (1957) stated that the specimen was not in Dr. Kishida's collection and a misidentification might have been made. Ac- cording to Kishida (19.30b), one female speci- men (lacking capitulum) was collected from
176
Cijclemijs flavomarginatd by N. Nakayama. He did not indicate the date and locality, but the host turtle is found on Ishigaki (Is.) and Irio- mote (Is.) in the Hyukyus.
Ambhjonima <^eoei)it/(lae (Cantor, 1847) is recorded from Ctjcleimjs fkwoimirginata on Irio- mote (Is.).
Ambh/oynina ciiprhim Neumann, 1(S99: 2 19-221;
Nakatsuji, 19-13: lcS2-lS4, Fig. 1.
Nakatsuji (1943) reported the collection of one female tick of this species from a turtle, Geoeimjda spengleri, on Okinawa. The figures published with this report are not adecjuate for discrimination.
Anastos ( 1950 ) stated that there were no au- thentic records of parasitism of reptiles by A. ciiprhim cijprium; it may be that the specimen reported bv Nakatsuji was actually '^eoemijdae, v\'hich is known to occur on turtles in the Ryu- kyu Islands (Keegan and Toshioka, 1957).
Haemaphiisttlis cinmiharina C. L. Koch, 1S44:
337; Kishida, 1922a ;846-855.
Kishida (1922a) listed both sexes of this species in his keys to the species which occur in Japan. Keegan and Toshioka ( 1957 ) stated that the occurrence of this species in Japan is very unlikely. Yajima (1956) listed it a,s cK-curring in Aomori Prefecture, but his record may have been derived from Kishida ( 1922a ) ( see Ap- pendix 1). Keegan and Toshioka (1957) also ({noted Kohls"s 1957 personal communication noting that cinmiharina, described from Brazil, is a valid name for chordeilis, which occurs in North America.
Haema]>htjsalis jxipuana Thorell, 1882:62-66;
Nakamura and Y;ijima, 1937:158-159, Fig.
5-9 in PI. VIII.
Nakamura and Yajima ( 1937 ) reported two female specimens collected from a dog at Hamg- yong-pukto. North Korea, 26 Julv 1935. Accord- ing to Anastos (1950) this is essentially a tropi- cal species, and Kohls (1957b) expressed doubt that this species occurs in Korea and other non- tropical regions. The illustrations provided by Nakamura and Yajima (1937) do not look like those given by Anastos (1950) but appear to be similar to those of //. japonica.
Haemaplnjsalis punctata Canestriui and Fanzago, 1878:189.
llaemaplujsaliH cinnahaiiud jiuiuialir. Neumann, 1907:215-2.32, Fig. 1-4; Kishida, !922a:852; Pomerantzev, 1950:92-93. Kishida (1922;i) listed l[. c. punctata in his
key. which seems to be derived from Neumann
BiiK.iiAM VouNC Univebsity Science Bulletin
(1907). Although Pomerantzev (1950) stated that the species had been recorded from Japan, such records do not (K-cur in the Japanese litera- ture. He pn)b;ibl\- cited Neumaim's record. Nutt- all and \\';irl)urton (1915) expressed doubt as to the correct identification of Neumann's ( 1907 ) specimen from a horse in Aomori Prefecture, Japan. There are no specimens in Japanese col- lections at the present time, and the species has i not been collected by Japanese acarologists.
Ilaeniaphysalis shibutamiensis Yajima, 1955.
This is apparently a nomen nudum. Although Keegan and Toshioka (1957) state that Yajima described the species in the ]aj)anesc Jattrnal of Sanitary 7.oolofiy 6( 1 ) :52-53, the specific name was neither given in the paper nor referred to in later papers bv Yajima. Tvpe specimens were not available for examination.
Haemapliysalis u(itamil>ci Yajima, 1942. (See under //. flava. ) '
llaemaplii/salis hirudo L. Koch, 1877.
There were no figures with the original des- cription, and the description itself was not ade- (juate for detennination. See under //. concinna.
Hyalomnm ac<^ii])ticum (Linnaeus, 1785); Kish- ida, 19.36:139.
Kishida (1936) reported this species from a cow on Quelpart Island ( = Chejudo Island, Korea), and it is the onlv record of this species from this area. Hoogstraal (1956) indicates that j //. aegypticum occurs in the Mediterranean area | and the Near East. It has been recorded from Russia, many parts of Asia Minor, and Afghan- istan. This species is primarily a p;uasite of tor- toises but may attach to other reptiles as well , as mammals and birds. Pomerantzev ( 1950) i gave infonnation concerning the distribution and biology of this species. On the basis of this in- fonnation, there is some doubt concerning the ] validity of the record from Quelpart Island (Kee- gan and Toshioka, 1957).
\xodcs acumimilus \vumd\m, 1901:287; Ki.shida, j
19.%: 142.
"Kishida ( 1936) stated th;it Mr. Ak:isawa had , collected Ixodes acuminutus from central Korea in 1928. No specimens have been reported from Korea, Japan, or the Ryukyu Islands since that date, and the species is not represented in exist- ing collections from these areas" (Keegan and To.shioka, 1957).
Ixodes affiuis Neuiiumn, 1899:120; Kishida, 1930a:3; Suzuki, 1930:90-92. Fig. 1. Kishid;i ( 1930a) listed /. affinis from ;i rabbit,
Biological Series, \'oi.. 15. \o. 1
Ticks
177
Lepus hmclii/urus an^tistidens. at Fukushinia City. 15 Deccniher 1929. collected bv Dr. Ohara. Suzuki (1930) described and illu.strated the species using a male specimen which had been taken from a rabbit of the same species. These specimens were not a\ailable for study, and unfortunatc]\' the reproduction of Suzuki's illus- tration of the male is not satisfactory for deter- mination of the species.
Ixodes corcensis Kishida, 193.3; Kishida, 1936:139. "Efforts to locate the original description of this species were not successful. Kishida (1936) referred to this tick, giving the tvpe locality as suburbs of Keijo ( = Seoul), and the host as Nyctereutes koreensis. The species is not repre- sented in any of the collections in Japan, and Dr. Kishida has infonned the author that the holo- type was lost during the war" ( Keegan and To- s'hioka, 1957).
Ixodes fossulattis Neumann, 1899:120; Kishida,
1930a: 2.
Kishida ( 19.30a ) listed this species as occur- ring on cattle and dogs at Sefuri-muri. Kanzaki- gun. Saga Prefecture. Keegan and Toshioka (1957) state that the specimens are not in the collection of Dr. Kishida and no additional specimens have been collected by Japanese workers, /. fossulattis is originally an Ecuadorian species.
Ixodes Juiiij)imis Yajima, 19.55; Keegan and To- shioka, 1957:31. This is a nomcn nudum. The specific name, 7.
hapjiinus. is not given in any published papers
by Yajima. See under /. acutitarsus.
Ixodes haijashii Nakatsuji, 1942:287-328. Fig. 3.
"This species description was based upon a single female tick taken from a dog on Miyake Island. 26 July 19.37, by K. Hayashi. Figures given of the female include dorsal and ventral aspects of the tick (showing only the posterior portion of the capitulum), hypostome, and tar- sus I. Unfortunately these are not adequate for identification of the species. The hypostome, as figured, is unlike that of any of the Japanese Ixodes. No specimens of this tick were collected by 406th MGL personnel and the species is not represented in an\- of the Japanese collections" (Keegan and Toshioka, 1957).
Ixodes lisphilus Kishida, 19.30; 1922b: 962-964,
Fig. 1-2, 19.30a:. 3-4.
This species was described by Kishida (1922b) using nvmphal specimens taken from a squirrel, Sciunis lis. Nagano Prefecture, Japan, 1 June 1918, by Uchida. However, he (1922b) did not give the specific n;une but designated
it later as a new species (1930a). According to Keegan and Toshioka ( 19.57), the type specimens have been lost and no other material was avail- able for study. Judging from the host and the figures given by Kishida (1922b), the species appears to be similar to Ixodes persulcatus.
Ixodes putus (Pickard-Cambridge, 1878). See under /. uriae.
Xenoacarus Kishida, 1925.
Palpi short; scutum not heavily sclerotized; postanal median groove extends to posterior body margin; festoons and eyes absent.
Xenoacarus ut'ikinoi Kishida, 1925:31.
The original description was not avail- able for translation. Dr. Kishida infonned Dr. Seiichi Toshioka that the original des- cription consisted of only a few lines and no illustrations. No specimens were available for study. The generic diagnosis given above and the following species description were taken from a discussion of the species in Essential Para.ntologii of the Domesticated Animals. April 1926," p. 199-200, by S. Yo- shida and T. Makino. A figure with this discussion showed a dorsal view of the scutum and capitulum of the female of makinoi.
"Male: Unknown.
"Female: Postanal median groove ex- tends to posterior body margin; dorsal sur- face of basis capituli rectangular and bow- ed forward anteriorly. Dorsal scutimi not heavily sclerotized and monochromatic. Pal- pi short as in Haemaplnjsalis. Coxa I with a large spur. [One unintelligible phrase pre- ceded this statement.] Festoons and eyes absent. Legs and scutum yellowish in color, even in engorged specimens." Tliis species has been found on leaves or stalks of plants. It has been collected from horses, etc. Para- type collected from Japan: Honshu, Aomori Prefecture, Kamikita-gun, Misawa-mura, Hirosawa pasture, August, 192-5 (Keegan and Toshioka, 1957).
ACKNOWLEDGMENTS
We are happy to express our appreciation to Mr. Glen M. Kohls, Rocky Mountain Labora- tory, Hamilton, Montana; Dr. Harry Hoogstraal, U.S. Naval Medical Research Unit No. 3, Cairo, Egypt; Dr. Shigeo Kitaoka, National Institute of Animal Health, Kodaira, Tokyo, who have been very generous in confinning our identifica-
178
Bhigham Vounc University Science Bulletin
tions or in loanins^ valuable specimens and ofter- iu'Z advice on taxa, the taxonoinie status ot whicli was in (juestion.
We wish to express our appreciation not only to the above but also to the following persons who have cooperated in various ways to make possibU; the completion of the paper:
Dr. Zen-emon Ono, Hokkaido Institute of Public Health, Sapporo; Dr. Kiyoshi Asanuma, Research Institute for Natural Resources, Tokyo; Mr, Keisaku Tattori, Hokkaido Institute of Public Health, Sapporo; Mr. Kimio Endo, Iwaizumi-cho, Iwate Prefecture; Mr. Kimito Uchikawa, Nagoya Institute of Public Health, Nagoya, for the kind loan of specimens.
Dr. Hiroshi Takahashi, Medical School, Tok- yo, Japan Ground Self-Defense Force; Dr. Kazu- yuki Uchinomi, Hachijo Station of Tokyo Cent- ral Health Center; Dr. Naoki Namba, Hokkaido National Argicultural Experiment Station, Sap- poro; Dr. Ryuichi Taniguchi, Shintoku Experi- mental Station of Domestic Animals, Shintoku, Hokkaido; Mr. Koichi Ono, Mita Pasture, Koz- awa, Hokkaido; Mr. Toshihide Takeda, principal of Yamato Middle School, Aniaini Oshima, Ka- goshima Prefecture; Mr. Zentoki Hajime, Yuwan, Amami Oshima, Kagoshima Prefecture; Dr. Shoi- chiro Ohara, Ohara Hospital, Fukushima; Mr. Susumu Yamamoto, Kagoshima Institute of Pub- lic Health, Kagoshima; Dr. Kunitada Sato, Shiz- unai Pasture of Hokkaido University, Shizunai, Hokkaido; Dr. Yasusuke Nagayoshi, Fukushima Hospital, Kushima, Miyazaki Prefecture; Dr. Me- guinu Hasegawa, Hokkaido Institute of Public Health, Sapporo; LTC Yasuo Nakamiya, Camp Obihiro, Japan Self-Defense Force, Obihiro Hok- kaido; Dr. Hajime Kanio, Department of Parasit- ology, Tottori University, School of Medicine; Dr. Atsuo Sato, Department of Parasitology, Ka- goshima University; Dr. Asahiko Yajima, Kitama- chi. Kichijoji, Musashinoshi, Tokvo; Dr. Yutaka Saito, Department of Medical Zoology, Niigata University of Medicine, Niigata; B. M. Mutsuko
Okumuni, Toyonaka-shi.
We are also grateful to the following persons. Department of Kntomologv, 406th Medical Lab- oratory, USAMCJJ, who have prepared illustra- tions for this publication:
Mr. Saburo Shibata, Mr. Takashi Ando, Mr. Shozo Ohtawa. Mr. Masao Ilasunuma, Mr. Yii- saku Sonobe, Mr. Yoshitsune S;isaki, Miss Mut- suko Misaki, and Mrs. Atsumi Ryuchi. In addi- tion, we express appreciation to Miss Yoshiko Yoshida for her careful recording and filing of the original illustrations made by the above art- ists.
We express thanks to Mr. Hiroshi Suzuki and Mr. Shogo Sasagawa for their cooperation and devotion to dut\' in the rearing of field material for laboratory life-cycle studies and to Dr. Kazuo Tanaka for hLs continuous encouragement and timely advice on taxonomic problems.
Mr. Kiyoyuki Mizusawa and other field col- lectors of the 406th Medical Laboratory provided a large proportion of the material examined in this paper.
We wish to take the opportunity to express thanks to LTC Alexander A. Hubert, chief of the Department of Entomology, and Dr. George M. Davis, Department of Medical Zoology, for their frequent encouragement and for reviewing the manuscript; to Mrs. Anne Lester for her pains- taking tvping and correcting of early drafts of this manuscript; we acknowledge the continuous support and interest of COL J.E. McCarty, Com- manding Officer of the 406th Medical Labora- tory.
We express our thanks to Mr. Robert C. Saunders and Mr. Lynden P. Baum for their review of the manuscript and Miss Phyllis Mad- den and Miss Dianna Rice for typing the final draft of the manuscript. All are students at Brigham Young University.
The authors would like to acknowledge with sincere appreciation all others who may have been omitted by name but who gave assistance.
LITERATURE CITED
American Geocrapiiicai. Society. 1954. .Atlas of di.seases. Plate 12. World distrihiition of rickettsial di.sea.ses - tick and mite vectors. ( New ^'ork ) .
Amerson, a. B.. Jr. U)68. Tick distrihulioii in the Central Pacific as influenced bv sea bird movement. J. Med. Ent. 5(.3) :332-.3.39.
Ana.stos, G. 1950. The scutate ticks, or Ixodidae, of Indonesia. Kntomologica Americana 30( 1-4 ); 1-144.
Anonymous. 1964. Survey on ticks in the pastiire at Sanbe. Sliimane Prefecture. Report of the Institute of Animal Health of Sliimane Prefecture, p. 1-10. (Japanese text, mimeographed.)
Arthur, D. R. 1955. Ixodes kohhi, a new species of
of lick from Austr.ilia. J Parasilol. 41 ( 1 ) : 18-23.
Arthur, 1). R. 1956. The Ixoilcx ticks of Chiroptera (Ixodoidea, Ixodidae). J. Parasit. 42(2) : 180-196.
.Ahihuh. D. R, 19,57. Studies on exotic ticks (Ixodoi- dea. Ixodidae) from United States Navv and Army activities. |. Parasit. 43(6) :681-694.
.\inTiuR, D. R. 1960. Ticks, a monograpli ol the Ixodoidea. Part V. On the genera DcrmiucntOT, Anoretitor, Coxmiommu. Bonphiliif:. and .\/(;rp;r();)i(.s. C;ambr. Univ. Press. 251 p.
Ahthuh, D. R. 1963. British ti<ks. Butlinvorth & Co., Ltd., London, 213 p.
A,s.\NUMA, K. 1941. Hticvuiplnj.stilix coiiciniui. Zool.
Biological Series, Vol. 15, No. 1
Ticks
179
Mag. 53(6):306-307. (Japanese te.\t.)
As.ANUMA, K. 1942. On tlie dog tick Hucmupht/salis campunuliitii. Ciinvciken 11 ( 1 ): 17-26. ( |apanese text. )
.As.\NUMA, K. 194-4a. Some obsenations on the (luan- tity of blood sucked by i.xodids and on their body weight. ( Studies of the egg-laying capacity of Ixodidae. I.) Inst. Re.s. Sci. Re.sour."Proc. (6):35- 40. (Japanese text.)
As.\NUMA, K. 1944b. Minimum quantity of sucked blood capable of inducing egg-deposition in ixodid-s. ( Studies on tlie egg-laving capacity of Ixodidae. 11.) Inst. Res. Sci. Resour. Proc. (6):41-46. (Jap- iuiese text. )
AsANUMA, K. 1944c. Influence of the (juantity of sucked blood on the number of eggs laid by ixodids. ( Studies on the egg-laving capacity of Ixotlidae. III.) Inst. Res. Sci. Resour. Proc. (6):47-54. (Ja- panese text. )
.■\sANUMA, K. 1944d. Egg-laying habits of ticks. Misc. Rep. Res. Inst. Nat. Resour. (19):21-31.
AsANUMA, K. 1947a. Note on the role of blood in ovulation in a tick, Hacmaphysalis hispinosa, to- gether with remarks on the so-called specific dif- ferences in the number of eggs laid by ticks. Studies on the ovipositing ability of ticks, V. Seibutsu 2(1):7-14. (Japanese text.)
AsANUMA, K. 1947b. p. 971-974. In Uchida et. al.. Rev. ed. of illustrated encyclopedia of the fauna of Japan. Hokur^ukan Co., Tokyo. 1898 p. ( Jap- iinese text. )
.\sANUMA, K. 19.51. Notes on the tick, Ixodes angus- tus Neuman, new to Asia. Misc. Rep. Res. Inst. Nat. Resour. 22.1-4. (Japanese te.xt. )
AsANUMA, K. 1954. Studies on ticks and bloodsuck- ing mites infesting rats (genus Rattus) in Japan. Part I. Misc. Rep. Res. Inst. Nat Resour. ( 36); 108- 119. (Japanese text. English translation: NAMRU .3-T48. )
AsANUNLA, K. 1955;l On the i.xodicLs and laeapitids pariisitizing mice of Hokkaido, and mode of trans- mission of pathogens among liost mice bv the acari. Nihon I ji Shinpo (1611):! 240- 1242.' (J apanese text.)
.\SANUMA, K. 1955b. Mode of disseminating the caus- ati\'e agent ;imong the rats with various ticks and mites ;ls vectors. Jap. Med. J. ( 161 1 ) :1240-1242. (Japanese text.)
AsANLMA, K. 19.56a. Tularemia, ticks, and mesostig- matid mites of the Far Eiist. Ann. Prof. Rep., 406th Medical General Laborator)'. p. 91-96.
.\s.\NUMA, K. 1956b. Personal communication.
AsANUMA, K. 1957. Occurrence of the tick, Ixodes angustus Neuman. in Hokkaido, Japan, with re- marks on its larval and n\mphal stages. Jap. |. Saint. Zool. 8(4):2.34. (Japanese text.)
.\sA>aiMA, K. 1958a. New hosts of ixodid ticks. Shawa .3.3rd Monbusho Sogo Kenkvii Hokoku Shuroko (Med. and Pharmacy), p. 279. (Japanese text.)
AsANUMA, K. 19.58b Ecology of Ixochdae parasitic on media animals for yato-b\o. 2. Rep. Ohara Hosp. 2(1):23. (Japanese te.xt. )
.\sANUMA, K. 1958c. Yato-bvo in Sado Island. 2. Rep. Ohara Hosp. 2( 1 ) :.38'-39, (Japanese text)
.\sANV>L'^, K. 1960. Host and distribvition records of the soft tick, Ornithodoros capensis Neumann, in Japan. Jap. J. Sanit. Zool. 11(2):94. (Japanese text.)
.\sANUMA, K. 1961. Blood-sucking acari - 2. Para- sitic ticlts of sea birds in Japan. Showa 36tti Mon- busho Kenkyu Hokoku Shuroku ( Med. and Phar-
macy), p. 181. (Japanese text.)
.\sANUMA, K. 196.5a. Suborder Ixodides, p. 101-128. In Sasa, Mites, an introduction to cliissification, bionomics and control of Acarina. Univ. of Tokyo Press. 486 p. (Japanese text.)
AsANUMA, K. 1965b. p. 397-401. In Okada et al., New illustrated encyclopedia of the fauna of Japan, Vol. II. Hokuryukan Co., Tokyo. 803 p. (Japanese te.xt. )
AsANUMA, K. 1969. Personal communication.
AsANUMA, K., AND S. FuKUUA. 1957. On the life-his- of a tick. Ixodes signatus, infesting a black-tailed gull (Larus crassirostris) in Japan. Jap. J. Sanit. Zool. 8(3):147-1.59. (Japanese text. )
AsANUMA, K.. AND K. KosAKA. 19.54. Notes on the oc- currence in Japan of a tick, Hacmaphi/satis welling- toni, collected from a Japanese gray thrush, Turdus cardis cardis. Misc. Rep. Res. Inst. Nat. Resour. 36:104-107. (Japanese text English translation: NAMRU 3-T35.)
AsANUMA, K., AND K. KosAKA. 1955. Notes on a tick, Ixodes turdus Nakatsuji 1942, found on birds in Japan. Bull. Biogeogr. Soc. Jap. Appendix 16-19: 192-196. (Japanese te.xt. English translation: NAM RU 3-T50.)
AsANUMA, K., AND H. Nakacawa. 1955. Studies on endemic and medically important arthropods on Niijima Island, Tokyo-to. Ochanomizu Igaku Zasshi 3( 6 ) :549. ( Japanese text. EngUsh summary, p. 680. )
Asanuma, K., K. Okubo., and S. Fukuda. 1955. In- festation of a black-tiiiled gull ( Larus crassirostris ) with a tick, Ixodes signatus, in Kabushima, Aomori Prefecture, Japan, Misc. Rep. Res. Inst. Nat. Resour. 38:85-86. (Japanese text.)
Asanuma, K., and N. Sakural 1958. On the seasonal occurrence of the tick, Haemuphijsalvi flava, on wild hares (Lepus timidus bracht/urus) in the en- demic area of yato-byo or tularemia in Chiba Pre- fecture, Japan ( A preliminary report ) . Misc. Rep. Res. In,st. Nat. Resour. (48):28-39. (Japanese text. English summarv, English translation: NAMRU 3-T46. )
Asanuma, K., and N. Sakurai. 1960. Comparison between the infestation of Ixodes japonensis and Haemaphi/salis fliiva on wild rabbits. 7. Rep. Ohara Hosp. 4(2):26. (Japanese text.)
AsANUM.'\, K., N. S.\kuhai, S. Ohara, H. Odajima, K. KosAKA, and H. Nakagawa. 1955. On the host and distribution records of a tick, Haemaphy satis flava, a dominant panisite of wild hares in Japan. Mi.sc. Ref. Res. Inst. Nat. Resour. (.37 ); 127-128. (Japanese te.xt. English translation; NAMRU 3- T49.)
.\sanuma, K. and K. Sekikawa. 1952. Studies on ticks of the genus Ixodes parasitic on small rodents of the superfamily Muroidea in tlie Japanese Islands (a preliminarv note). I. Misc. Rep. Res. Inst. Nat. Resour. ( 28)';107-1 16. (Japanese text.)
Asanuma, K., .'Vnd K. Sekikawa. 1953. Studies on ticks of the genus Ixodes parasitic on small rodents of the superfamilv Muroidea in the Japanese Islands (a prehminary note). II. Misc. Rep. Res. Inst. Nat. Resour. (31):99-112. (Japanese text.)
Asanuma, K., A. Shishido and M. Kitaoka. 1960. The hfe of Rickettsia in the lx)dies of larvae and nymphs of II aenuiphy satis- tiispiiwsa. 7. Rep. Ohara Hosp. 4(2):26. (Japiuicse text.)
.\sANUMA, K., T. Takeda, and H. Nakacawa. 1961. Some observations on ticks found on birds in the Mt. Fuji area. (Japanese abstract of report before
ISO
Bmr.HAM VouNc; University Science Bulletin
13 Ann. Meet. Jap. Sw. Saiiil. Zool., |ulv 5.) Jap. J. Sanit. Zool. 12(2): 139.
Asanuma, Y. 1943. Con.siderations on the amount of 1>1o<kI .sucked h\ two .species of ticks parasitic on dogs. Acta Araciin. 8(4):87-97. (Jap:uiese te.\t. )
AuDY, J. 11, .\I. Nadchatham, and B. Lim. 1960. Host di.strihution of Malayan ticks ( l.xodoidea ) . Malaysian piirasites - XLIX. Stud. Inst. Med. Res. .Malaya 29:225-246.
Badenko, L. V. 1956. Ixodes Inkliis Koch as a rep- resentative of ixodid burrow inhahitiiijT ticks. Mater. Pozn. Fauny Florv SSS R, n. s. Otd. Zool., vypusk ,34 (XLIX); Ektoparazity, Fauna, Biol., Prakt. Znachenie (3):21-105.
Balashov, Yu S. 1954. Pecidiarities of the day and night rliythm of the dropping of engorged females of Ixodes persulcatus from cows. Dokl. Akad. Nauk. USSR, n. s. 98(2):317-319.
BiKULA, A. 1895. Ixodidae novi vel parm c-ogniti Musei ZtK)logici Academiae Caesareae Scientianim Petropolitinae. Bull. Acad. Sci. St. Petersb. Ser. 5, 2(4):.35.3-.364.
Bi.ANCHAHD, R. 1909. L'in.secte et I'infection, histoire naturelle et medicaUe des arthropodes pathogenes. Fa.sc. 1. Aczriens. Libraire Scientifique et Litter- aire; Paris. 160 p.
Bhemner, K. C. 1959. Observations on the biology of Hacmaphijsalis hispinosa Neumann (Acarina: Ixo- didae) with particular reference to its mode of reproduction by parthogenesis. Aust. J. Zool. 7:7-12.
Canestbini, G. 1887. Intomo ad alcuni acari ed Opilionidi dell'America. Atti. Soc. Veneto-Trenf. Sci. Nat. 11:104-105.
Canesthini, G., and F. Fanzaco. 1878. Intorno agli acari italiani. Atti R. 1st. Veneto Sci. Lett. Arti (1877-1878), 5. s, 4(l):69-208.
Cantor, T. 1847. Catalogue of reptiles inhabiting the Malay Peninsula and Islands. J. Asiatic Soc. Bengal 16:608.
Cerny, v., and E. Khatochvilova-Kkalova. 1963. Development of the tick Hiiemapfn/fialis concintiu in natural conditions of southern Moravia ( Czechos- lovakia). Zool. Listy 12(3):259-261.
Chikaki, H., and a. Ot.\ke. 1956. Ecology of ticks attacking cattle in a pa:;ture. J. Jap. Soc. Applied Zoology 21(2):53-62. (Japanese text. EnglLsli summiuy. )
Chikaki, H., A. Otake, and T. Miura. 1956. Con- sideration of cotton-flannel for the investigating method of Haemuplu/sidis hixpinosa Neumann (Acarina: I.vodidae) in the pasture. Jap. |. Ecology 6(l):35-39. (Japanese text. English summary. )
Cooley, R.' a. 1946. Note on the tick Ixodes angustiis Neumann. J. Parasit. 32(2):210.
CooLEY, R. A., and G. M. Kohls, 1945. The genus Ixodes in North America. Nat. Inst. Hlth Bull. 184:1-246.
Dhanda, v., and U. R. Biiai. 1968. Ilacinuphi/sidi.s- (Haemap]ujsulis)indoflmii sp. n. (Acarina: Ixodi- dae) from the Himala\an region of Uttar Pradesh and Madras. India. J. Parasit. .54(5) : 1063-1067.
Donit-z, \V. 1905. Die Zecken des Rindes als Kran- kheitsubertrager. S. B. Ges. Naturf Fr. Berlin 4:105-134.
Elbl, A., and G. Anastos. 1966a. Ixodid ticks (Acarina, Ixodidae) of central Africa. Volume I. General introduction. CJenus Andili/omma Koch, 1844. Ann. Mus. R. Afr. Cent., Ser. .Svo. s. .Sci. Z<x)l. (145). 275 p.
Elbl, A., and G. Anastos. 1966b. Ixodid ticks (Acarina, Ixodidae) of central Africa. Vol. II. Cienus Ixodes Latreille. 1795. Ann. Mus. R. Afr. Cent., Ser. Suo, s. Sci. Zool. (146). 205 p.
Elbl, A., and G. Ana.stos. 1966c. Ixodid ticks ( Aca- rina, Ixodidae) of central Africa. Vol. III. C.enus RhipU-epludus K(X-h, 1844. Ann. Mus. R. Afr. Cent., Ser, 8vo. s. Sci. Zool. (147). 555 p.
Elbl, A., and G. Anastos. 1966d. Ixodid ticks (Aca- rina, Ixodidae) of central .Africa. Vol. IV. Genera Aponommu Neumann, 1899, Boophilus Curtice, 1891, Dermaceiiliir K(x-h, 1844. Hiieni/ipluiscdis Koch, 1844. Ilt/cdommu Koch, 1844 and Hhipi- ceiitor Nuttall and Warburton, 1908. Lists and bibliography. Ann. Mus. R. Air. Cent.. Ser. Svo. s. Sci. Zool. (148^ 412 p.
Feldman-Muhsam, B. 1952. On the identity of Rhi- piccpludus sanfiuitieus. Lat. Bull. Res. Council Israel 2( 2 ): 187-194.
Fuller, C. 1899. Notes on the Queensland cattle tick and its relationship to the Texas fever tick and the blue tick of Cape Colony, South Africa. Qd. Agric. J. 4:389-394.
Glaschenskaya - Babanko. 19,56. (See Babenko, 1956.)
Han, T. W., and Y. Jeon. 1967. Studies on ticks. III. Observations on the life cycle of Haetiuipht/scdis Imiiiuosa. Res. Rep. Off. Rural Dev. 10( 5) :. 3.5-42. ( Korean text. English suniniarx'. )
Han. T. W., and S. K. Kim. 1965. Survey and arti- ficial infection of bovine PiropUisartia in Korea. Res. Rep. Off. Rural Dev. 8(3 ): 109-120. (Korean text. English summarx'. )
Han, T. \V., Y. Jeon, and S. K. Kim. 1967. Studies on ticks. II. The monthly life flow of ticks in Korean cattle. Res. Rep. Off. Rural De\ . 10(5): 2.5-33. ( Korean text. English summar\ . )
Han, T. W., S. K. Kim, and Y. Jeon. 1966. The classification and distribution of Korean cattle tick. Res. Rep. Off. Rural Dev. 9(3):91-103. (Korean text. English summary. )
Hara, J. 1963. A case report of the fowl tick, Ar^as persicus (Oken, 1818) which invaded residences (Acarina: Argasidae). Bull. School. Physi. Edu., Jnntendo Uni\-. 6:123-125. (Japanese text. English summar\'. )
Hirst, S., and L. F. Hirst. 1910. Description of five new species of ticks (Ixodidae). Ann. Magaz. Nat. Hist. 8(6):299-.308.
Honma, T., .\nd S. Ohara. 1961. Morphology of yato-byo ticks. II. Internal stnicture of Hacma- phi/scdix fldvd Neumann 1897. Male. .\nn. Report - Ohara Hosp.. No. 8. .5(I):l-7. (Japanese text. English summary'. )
Hoocstraal. H. 19,56. .■\frican Ixodoidea. I Ticks of the Sudan (with special reference to Ef|uatoria Province and with preliminary reviews of the genera Boophilus, Margarojtus, and Ht/idonima). Bureau of Med. and Surgers . Department of the Navy. 1101 p.
HooosTR^Ai.. H. 1961. Rediscriplion of Ilnermiplii/- s(dis ( Alloccriiru) incrmis avouommoides War- burton. 1913 (Ixodoidea, Ixodidae). J. Parasit. 47 (2):317-318.
H<)Oc.sTR\Ai., H. 1962. Haemaphijsahs ncpidensis sp. n. from a Himakuan rodent and man and de- scription of the male of H. aponommo'des War- burton (n. comb.) (Ixodoidea Ixodidae). J. Para- sit. 48(2): 19.5-203.
Hoocstraal, H. 1966. Ticks in relation to human diseases caused b\ viruses. .\nn. Rev. Ent. 11:261-
BiOLOOiCAL Sehies, Vol. 15, No. 1
Ticks
181
3()S.
HooGSTK.\AL, H. 1967a. Tick - Iwme hemorrliagic fevers, encephiUitis, and typhus in USSR and southern Asia. Exper. Parasit. 21( 1 ) :98-lll.
Ht)OGSTR.\-\L, H. 1967b. Ticks in rehition to liuman diseases caused bv rickettsia species. Ann. Rev. Ent. 12:377-420.
Hi)(k;sth.\al, H. 1968. Personal communication.
HoocsTR.\.4.L, H. 19e9a. Letter No. 1013. 28 Mav 1969. N.WIRU 3.
HoocsTRA.^L, H. 19(39h. Hacmaphiisulis (Allocenwd) kitaokai sp. n. of |ap;ui, and keys to species in the stnicturidly primiti\e subgenu* Alloceraea Schulze of Eurasia ( l.xodoidea, I.xodidae). J. Parasit, 55(1): 211-221.
HoocsTKAAL, H., AND M. N. Kaisek. 1960. Booplii- liis kohlsi n. sp. (,'Vcarina: Ixodidae) from slieep and goats in |ord;ui. J. Parasit. 46(4 ) :441-448.
HoocsTK.\.\L. H., F. H. S. Roberts, G. M. Kohls, and V. J. Tipton. U)68. Review of Hciemaplii/sali.i ( Kaiserianu ) loiif:,con>h Neumann ( resurrected ) of Australia, New Zealand, New Caledonia, Fiji, Japan, Korea, northeastern China, and USSR, and its parthenogenetic and bisexual pop ulations (Ixodoidea, I.xodidae). J. Parasit. 54(6) : 1197-1213.
HocxJSTHAAL, H., AND H. Trapido. 1966. Redescrip- tion of the type materials of Hacmaplnjsalis ( Kai- seriana) bispinosa Neumann (India), H. (K. ) neumanni Donitz (Japan). H. (K.) higrangei Lar- rousse (Vietniun), and H. (K.) i/eni Toumanoff (Vietnam) (Ixodoidea, l.xdidae). J. P;\rasit. .52(6): 1188-1198.
HoocsTRAAL, H., H. Trapido, and C. M. Kohls. 1965. Studies on soutlieast Asian Hacmaphifsalis ticks (Ixodoidea, Ixodidae). The indentity, distribution, and hosts of //. (Kaiscriami) lu/stricis Supino. J. Par:isit. 51(3):467-480.
HoocsTRAAL, H., AND N. Vamac.uti. 1970. llacma- physalis ( H. ) pentalagi Pospelova-Shtrom, a para- site of the (apiuiese black rabbit; rediscription of the male and description of the femiile, nymph, and larva (Ixodoidea, Lxodidae). J. Parasit. 56(2):367- 374.
Inatomi, S., and N. Yamaguti. 1960. On an Ar- gasid tick found at Niimi Cit\ . Bull. West Br. Jap. Soc. Sanit. Zool. 15:17-18, (Japanese text.)
IsHiHARA, T., AND S. IsHii. 19.56. Studies on piro- pla.smosLs bv Theileria mutans in Japan. Abst. Proc. Mon. Meet. Staff Natl. Inst. Anim. Hlth. 3:41. (Japanese text.)
Itagaki, S., R. Noda, and T. Vamaguchi. 1944. Ticks parasitic on domestic animals of Asia. Jap. Horse-Racing Assn. 97 p. (Japanese text.)
Itagaki, S., R. Noda, and T. Yamaguchi. 1959. Studies on the ticks of the domestic animals in the Far East. Jap. Soc. Promotion Sci. 118 p., 27 pi. Ito, S. 190.3. Ixodes riciniis. Chugai Iji Shinjjo (561), 23(15):ia32-10,34 (p. 24-26).
Ito, Y.. T. Komatsu, and A. Suzuki. 19.59. Chronic tularemia and the epidemicological mass survev in Takada Citv, Iwate Prefecture. 4. Rep. Ohara Hosp. 3(l):25-29. (Japane.se text. English sum- mary.)
Iro, T., and Y. Saito. 19.54. On the rickettsioid micrtxjrganlsms foimd in Steatoniissus sp. and Arnii.s sp. parasitic on PinistrcUus ahramus. Tokvo IJLshinishi 71(10):.3-4. (Japanese text.)
Ka.mo, H. 1962. Parasitic acari in western part of Japan. Showa .37th Monbu.sho Kenkvu Hokoku Shnroku (Med. and Pharmacv). p. 173. (Japanese text. )
Karsgh, F. 1880. Vier neue Lxodiden des Berliner Museums. Mittheil. d. Munchener Entomol. Ver- eine. Jahrg. 4:141-142.
Katsura, S. 1957-1959. Studies on the bacterial and infectious diseases transmitted by ticks in Japan. Showa 32iul, 33rd, and 34th Monbusho Sogo Kenkyu Hokoku Shuroku. p. 265-274 (1957); p. 277-288 (1958); p. 327-336 (1959). (Japanese text. )
Kaw.^vshima, K. 1963. Studies on the acarine fauna of the Ryukyu Islands ( 1 ) . Some records of ticks. Jap. J. S;uiit. Zool. 14(2): 103. (Jap;inese text.)
K.\WASH1MA, K., H. KaMO, and I. MiYAZAKI. 1960. A
case of human infestation with a hard tick, Am- bhjomma tcstudinarium Koch, in Japan. Kyushu J. Med. Sci. ll(2):77-80.
Keegan, H. L., and S. Toshioka. 1957. Ixodid ticks of Japan, Korea, and the Rvukyu Islands. Report — 406th Medical General Laboratory. 37 p., 42 pi.
Kierans, J. E., and G. M. Kohls. 1970. Ixodes philipi n. sp. (Acarina: Ixodidae), a sea bird parasite in Japiui. J. Med. Ent. 7(6) :725-726.
KisHiDA, K. 1922a. Notes on the ticks of the famiK' Ixodidae parasitic on Japanese mammals (Artiixlactvla: Bunodontia ) . Zool. Mag. 34(408): 846-857. (Japanese text.)
KisHiDA, K. 1922b. Tick from a squirrel. Zool. Mag. 34(410):962-964. (Japanese text.)
KisHiDA, K. 1925. Abstracts of the Seventh meeting of the Tokvo Parasitological Society, No. 2459:31.
KisHnJA, K. 1927. Acarina, p. 985-986. lit Oka et al.. Illustrated enc\clopedia of the fauna of Japan. Fig. 1897-1899. Hokuryukan Co., Tokyo. (Japan- ese text. )
KisHiDA, K. 1929. Supplements to the knowledge of Japanese cattle tick, Boophilus annulatus cau- datus. Lansania 1(2): 19-22. ( Japiuie.se text. )
KisHiD.^, K. 1930a. Notes on Japanese ticks of the genus Ixodes Latreille, 1795. Lansania 2(ll):l-5. ( Japanese text. )
KisHiDA. K. 1930b. Occurrence of a turtle tick, Artd)h/omnm eordiferuin, in Loo-choo. Lansania 2(19): 134. (Japanese te.xt.)
KisHiDA. K. 1935. Notes on the Acarina-mites and ticks, from the islands of Formosa collected in August 1935. Lansania 7(69) : 129-144. (Japanese text.)
KisHiDA, K. 1936. Notes on the Acarina-mites and ticks, from the peninsula of Korea and the island of Quelpart collected in August 1936. Lansania 8(79): 131-144. (Japanese text.)
KisHiDA, K. 1939a. i3n the cattle ticks of the Boophyleae from Japan. Zool. Mag. 51(7) :. 538-552. (Japanese text.)
KisHiDA, K. 1939b. Arachnida of Jehol: Order Aca- rina. Rep. 1. Sci. Exped. Manchoukuo, Sect. 5, Div. 1, Pt. 4, Art. 13. p. 1-49. PI. 1-.5.
KtsHiDA. K. 1947. Acarina, p. 972-975. 7;i Uchida et al.. Rev. ed. of Illustrated encvclopedia of the fauna of Japan. Fig. 2767, 2774, 2775. Hokuryii- kan Co., Tokvo. 1868 p. (Japanese text.)
KisHiDA, K., AND K. AsANUMA. 1965. Acarina, p. 397. In Okada et al.. New illustrated encyclopedia of the fauna of Japan. Hokiirvukan Co.. Tokyo. 803 p. (Japanese text.)
KiTAOKA, S, 1961a. Physiological and ecological studies on some ticks. IV. Physiological stage and lipid deposit during the blood-sucking process in the tick. Nat. Inst. Anim. Hlth. Quart. l(2):8.5-95.
KiTAOKA, S. 1961b. Physiological and ecological studies on some ticks, V. Nitrogen and iron
182
BmcHANr '^oUNG Univebsity Science Bulletin
excrt'tion and amount of blood meal ingested durini; tlu' hlood-siitking process in tlie tick. Nat. Inst. Anim. Hltli. yuart. 1(2);96-104.
KiTAOKA, S. 1961c. Physiological and ecological studies on some ticks. V'l. Rate of digestion of l)l(K)d meal and nitrogen, iron, and sterol economy during the oNipositing process in the tick. Nat. Inst. Anim. Hlth. ^uart. 1( 2) :1()5-1 12.
KiTAOKA, S. 1961d. Physiological and ecological studies on some ticks. VII. Parthenogenetic and hise.\ual races of Haetnupht/suli.s hiiintwaa in |apan and experimental crossing between them. Nat. In.st. Anim. Hlth. Vuart. 1 ( 3) ;142-149.
KiTAOKA, S. 1962. Physiological and ecological stud- ies on .some ticks. \'III. Diunial and nocturnal changes in feeding activity during the blood- sucking process of Hcicmaplu/suli.'i hixpiiiosti. Nat. Inst. Anim. Hlth. Viiart. 2( 2 ) : 106-1 1 1.
KriAOKA. S. 1967. Some characteristic features of feeding in ticks. Jap. Agric. Res. Quart. 2(4): 18-21.
KiTAOKA, S. 1970. llcicmapht/suli-s (llaemaphyjsalis) fiifi.saria sp. n. ( I.xodoidea, I.xodidae), a cattle |xxrasite in Japan. Nat. Inst. Anim. Hlth. Quart. 10(2):25-33.
KiTAOKA, S.. AND T. MoRii. 1963. Supplementary tests on the effects of new organophosphonis and other compounds ;i,s tickicides against Bo- ophilus nticniptus: Nat. Inst. Anim. Hlth Quart. 3(l):32-.35.
KiTAOKA, S., AND T. MoHii. 1966. The l)iologv of Hacniaplu/salis irwrniis Binila and //. cnncinna Koch with particular reference to the mode of blood-sucking process. (Abstract 18. Ann. Meet. Jap. Soc. Sanit. Zool. April 29-30, 1966). Jap. J. Sanit. Zool. 17(2) : 141-142. (Japanese te.xt. )
KiTAOKA, S., AND T. MoRii. 1967a. The biology of Hacinaphi/xali.'i ( Alloccraea ) amhigua Neumann, 1901, with description of immature stages. ( I.xo- doidea. I.xodidae). Nat. Inst. .-^nim. Hlth. Quart. 7(3): 145-152.
KiTAOKA, S., AND T. MoRii. 1967b. Effects of gamma radiation and chemosterilants on the cattle tick, Boophilus microplus. Jap. I. Sanit. Zool. 18(2-3): 126-129. (Japanese text.)'
KiTAOKA S., AND Y. Saito. 1967. Ixodes nipponetvtis n. sp. (Ixodoidea. Ixodidae). \ common cattle tick in Japan. Nat. Inst. Anim. Hlth. Quart. 7(2): 74-83.
KiTAOKA. S.. AND A. Vajima. 19.58a. Ph\siological and ecological stiulies on some ticks. I. Process of growth bv blood-sucking. Bull. N:U. Inst. .iKnim. Hlth. 34:1.35-147.
KiTAOKA, S.. AND A. Yajinia. 1958b. Physiological and ecological studies on some ticks. II. Phase change of ovi[x>siting ability with blood-sucking quantity. Bull. Nat. Inst. Anim. Hlth. .34:149-162.
Kitaoka. S.. and a. Yajima. 19.58c. Physiological and ecological studies on some ticks. III. Para- sitic duration of Ritophilus ritudatus. Bull Nat. Inst. Anim. Hlth. 35:179-188.
Kitaoka, S„ and A. Yajima. 1 961 a. Comparison of effectiveness between pesticides :igainst Bo- ovhilus microplus bv topical application and spray- ing. Nat. last. Anim. Hlth. Quart. I(I):4I-52. '
Kitaoka, S.. and A. Yajima. 1961b. Bisexual par- thenogenic races of Hiirmaphi/salis hisninosa. (Japanese abstriict of report before 13. ,\nn. Meet.) Jap. J. .Sanit. Zool. 12(2) : 141-142. (lapauese text.)
Koch. C. L. 1844. Svstematische Ulx-rsicht liber die Ordnung der Zecken. .Arch. Naturgesch. 1:217-239.
Koch, C. L. 1887. Japanesische Araclmiden und Myriapoden. Verh. k. k. zoolog-botan. Ges. Wien. 27:786.
Kohls, C. M. 1950. Ticks (Ixodoidea) of the Philippines. Nat. Inst. HUh. Bull. 192:1-28.
Kohls, G. M. 1957a. Acarina: Ixodoidea. In.sects of Micronesia 3( 3) :8.5-I04. Bemice P. Bishop Museum, Honolulu, Hawaii.
Koiiis, G. M. 19.57b. Ticks (Ixodoidea) of Borneo and Malaya. Malaysian Parasites — XVIII. Stud. Inst. Med. Hes. Malaya 28:65-94.
KoHi.s, (;. M. 1967. Personal communication.
Khijcsman, B. J., AND S. A. S. PoNTo. 1932. De teken van den Oost-Indischen .\rchipel. Veeart- senijk, .Meded. Batavia 79:62 p., .5 maps.
I.ATREILLE, P. A. 1802. Histoire naturelle, generale et particuliere des crustaces et des insectes ( etc. ) . 3:468 p. Paris ( Cf . p. 67-68).
L.\THEiLLE, P. A. 1806. Genera cnistaceonim et insectonim. I. p. 1.57.
Leeson, H. S. 19.51. The recorded distribution of the tick Rhipiccplialus sanguineus (Latreille). Bull. Ent. Res. 42( 1 ): 123-124.
Linnaeus, C. 1758. Systema Naturae. Regnuni Ani- mals, 10th ed.
LouNsiiURY, C. P. 1905. Habits and peculiarities of some South African ticks. Brit. Assn. Advanc. Sci. Sec. D. S. Afric. 10 p.
Mei.'nikova, T. G. 19.58. On the development of UaenHiphijsalis concinna ticks under natural con- ditions of the Crimean State game preser\'e. Zool. Zh. .37(2):297-300. (Ru.ssian text. English translation: NAMRU e-T207.)
Minning, W. 1934. Beitrage zur Systematik und Morphologic der Zeckengattung Boophilus Curtice. Z. Parasitenk. 7(1): 1-43.
MiNNiNC, W. 1935. Zur Kenntnis des genus Bo- ophilus Curtice. I. Z. Parasitenk. 7(6) :719-724.
MiNNiNC, W. 1936. Zur Kenntnis des genus Bo- ophilus Curtice, II. Z. Parasitenk. 8(3) :365-370.
Morel, P. C. 1963. Identite d'lxodes ovatus Neu- mann, 1899 (Acariens, Ixodidae). Ann. Parasit. Hum. Com. ,38(6) :92.5-928.
NADCHATRAXf. M. I960. Notcs on rearing Malayan ixodid ticks ( Acarina. Ixodidae ) with special ref- erence to Ixodes firunukitus Supino and Amhli/om- ma neoevufdac (Cantor). Malaysian Parasites — .XLVIII: Stud. Inst. Med. Res. Malaya 28:217-224.
N.\CAHANA, M., AND K. Matsuo. 1962. Two species of ticks parasitic on the human body. Med. and Bio. 62(5):1I9-12I. (Japanese text.)
Nakamura. T.. AND A. Yajima. 19.37. On moipho- logy and distribution of the family Ixodidae in Japan, particularly species parasitizing cattle and horses. Jeuki Chosasho Kenk\ u Hokoku ( Rep. of Gov. Exp. Sta. for Anim. Hyg. ) 17:1.3.3-184. ( Japanese text. )
Nak\mura, T.. AND A. Yajima. 1942a. On the life history of Haemaphi/salis hi.ipinosa Neumann, 1897. Rep. of Gov. Exp. St. for Anim. Hyg. 19:21-23. ( Japanese text. )
Nakamura, T., and A. Yajima. 1942b On tlic l:ir\al stages of four species of ticks of the family Ixodidae. Rep. of Gov. Exp. Sta. for Anim. Hvg. 19:.34-,39. (Japanese text.)
Nakatsuji, K. 1942. Arachnida from Izu-.shifito. No- gaku Shuho. Tokyo Univ. Agric. I (4) :2«7-328. (Japanese text.)
Nakatsuji, K. 1943. Some Arachnida from Is. Oki- nawa and Is. Amimi Oshima. Nogaku Shuho. To- kyo Univ. Agric. 2( I ): 182-184. (Japanese text.)
HioLocicAi, Sehies, Vol. 15, No. 1
Ticks
183
Namb.\, N. 1953. On the effect of temperature and
humidit)' upon the embnonic de\elopnient of
ticks in pasturage. Res. Bull. Hokkaido Nat. Agric.
Exp. Sta. M: 130-135. (Japanese text. Enghsh
summary. ) Namb.\, N. 1954. On the behavior of hirvae of
Hacniaphysalis hispiiiosa on pLmts. Hokkaido Nat.
Agric. Exp. Sta. Rep. 66:49-55. ( Japanese text.
Enghsh summar)'. ) Namba, N. 1958. Ecological studies on Haemaphy-
salis hispinosa. a liarmful tick in the pasture of
northern Japan. Hokkaido Nat. Agric. Exp. Sta.
Rep. 50:1-99. (Japanese text. English summary.) N'.AMBA, N. 1963a. On methods of collecting unfed
ticks on vegetation in grazing pastures. Jap. Soc.
Grassland Sci. J. 9:21-28. (Japanese text. English
summ;u-y. ) Namba, N. 1963b. Researches on the distribution
of ticks and their habitat in pastures of Hokkaido.
Res. Bull. Hokkiiido Nat. Agric. Exp. Sta. 80:103-
114. (Japanese text. English summiin. ) Neum.\nn, L. G. 1897. Revision de la f;unille des
i.xodides. 2e memoire. Mem. Soo. Zool. Fr. 10(3-4):
324-420. Neumann, L. G. 1899. Revision de la famille de.s
ixodides. 3e memoire. Mem. Soc. Zool. Fr. 12:107-
294. Neumann, L. G. 1901. Revision de la famille des
Lxodides. 4e memoire. Mem. Soc. Zool. Fr. 14
(2-3): 249-372. Neumann, L. G. 1902. Notes sur les ixodides. I.
Arch. Parasit. 6( 1 ) :109-128. .Neumann, L. G. 1904. Notes sur les i.xodides. U.
Arch. Parasit. 8(3) :444-464. Neumann, L. G. 1905. Notes sur les ixodides. III.
Arch. Parasit. 9( 2) :225-241. Neumann. L. G. 1906. Notes sur les ixodides. IV.
Arch. Parasit. 10(2 ): 195-219. Neumann, L. G. 1907. Notes sur les i.xodides. V.
Arch. Para.sit. 11(2) :21. 5-2.32. Neumann, L. G. 1908. Notes sur les ixodides. VI.
Arch. P;irasit. 12(l):.5-27. Neumann, L. G. 1911. Ixodidae. Das Tierreich, 26.
Berlin. 169 p. Neumann, L. G. 1913. H. Sauter's Formosa Aus-
beute. Pediculidae. Siphonaptera, Ixodidae. Sup-
plementa Entomologica herausg. vom Deutschen
Entomol. Museum No. 2:1.34-137. Berlin Dahlem. Neumann, L. G. 1916. Ixodidei (Acariens): pre- miere serie. Arch. Zool. exp. gen. 55( 12) :517-527. Nuttall, G. H. F., and C. Warburton. 19t)8. On a
new genus of the Ixodoidea together with a
description of eleven new species of ticks. Proc.
Cambr. Philos. Soc-. 14(4 ) :.392-416. Nuttall, C;. H. F., and C. Warburton. 1911. Ticks.
a monograph of the Ixodoidea. Part II. Ixodidae.
Section I. Classification. Section II. Tlie genus
Ixodes, p. I0.5-.34S. Cambr. Univ. Press. Nuttall, G. H. F., and G. Warburton. 1915. Ticks.
a monograph of the Ixodoidea. Part III. The genu.s
Haemaphiisahs. p. 349-550. Cambr. Univ. Press. OoURA, K. 19.36. The ticks parasitic on the principle
domestic anim ds in Formosa, Japan. Mem. Fac.
Sci. Taihoku Imp. Univ. Formosa 19(2) :75-8.5. Ogura, K.. AND K. Takada. 1927. The ticks para- sitic on cattle and horses in Hokkaido Japan.
J. CoU. Agric. Hokkaido Imp. Univ. I8( 4 ): 199-206. Ohara, S. 1954. Studies on vato-byo (Ohara's dis-
ea.se, tularemia in Japan). Rep. I. [ap. J. Exp.
Med. 24:69-79. Ohara, S. 19.5.5. Studies on vato-b\o (Oh:ira's dis-
ease, tularemia in Japan). Rep. II. Jap. |. Exp. Med. 25:7-14.
Ohara, S., and K. Uhikawa. 1962. The distribution of yato-byo and its clinical observations. Annual Report of' Ohara Hospital No. 11, p. 1-2.3. (Jap;ui- ese text. English summary. )
Ohara, S., T. Noguchi, T. Honma .and M. Kato. 1959. Morphology of yato-byo ticks I. Internal structure of Haemaphysulis flava Neumann, 1897, Female. Ann. Rep. — Ohara Hospital, No. 6. 3(1):8-18. (Japanese text. Engli.sh summary.)
Ohara, S., and M. Sato. 19.53. Tick-borne infection of yato-byo. Rinshoigaku 8:87.
Ohara, S. and S. Tamura. 1958. A case report of a tick infesting die hirnian anus. Bull. West Br. Jap. Sanit. Zool. 14:23-24. (Japanese text, mimeo- graphed.)
Oken, L. 1818. Sogenannte giftige Wanze in Per- sien. Isis. p. 1567-1570, PI. 19, Fig. lA.
Oliver, J. H. Jr., and C. Bremner. 1968. Cyto- genetics of ticks (Acari: Ixodoidea) 3. Chromo- somes and sex determination in some Australian hard ticks (I.xodidae). Ann. Ent. Soc. Amer. 61 (4):8.37-844.
Oliver, J. H., and H. Tanaka. 1968. Personal com- munication.
Olsufjev, N. G., and V. G. Petrov. 1960. Discovery of HacmuphysuUs concinnn Koch ticks naturally infected by the tularemia agent. Trud. Inst. Zool. Akad. Nauk. Kazakh. USSR. (12):54:56.
Ono, Z. 1962a. Descriptions on the immature forms of Ixodes sp. reared in the laboratorv of Ohara General Hospital, Fukushima, Japan. Ann. Rep. Ohara Gen. Hospital, No. 11, 6(2):24-29. ( Japanese text. Engh.sh summary. )
Ono, Z. 1962b. Some notes on the ectoparasites found on small rodents :uid shrews in Maru- vama district, Sapporo. Hokkaido. Jap. J. Sanit. Zool. 13(2):1.5.5. (Japanese text.)
Ono, Z. 1966. Studies on ticks parasitic on small rodents of the family Muridae in Hokkaido, Rep. Hokkaido Inst. Publ. HIth. 16:62-68. (Japanese text. English summary. )
Ono, Z. 1967. Ixodes lividus (C. L. Koch, 1844). A bird tick new to Japan. Jap. J. Sanit. Zool. 18 (4):217. (Japanese text.) "
Pavlovsky, E. N., and V. D. Soloviyev. 1963. Hae- maphysalis concinna ( circulation of the vinis of spring-summer encephalitis in the organism of tick-vector Haemaphysahs concinna). R;xbot. Ek- sper. Parazit. p. 197-204. lUus. (Russian text. Eng- lish summary.)
PicKARD - Cambridge, O. 1878. An ac-count of the collections made in Kerguelen Land during the transit of Venus Expedition (1874-7.5). 2^1ogy: .Arachnida. In Philos. Trans. Rov. Soc. London 168:222. PI. 13. Fig. 4.
Pomerantsev, B. I. 1950. Fauna of the USSR. Arachnida. 4(2). I.xodid Ticks (Ixodidae). Zool. Inst. Acad. Sci. USSR. New Series No. 41, Mos- cow. 224 p. ( English translation published by Amer. Inst. Biol. Sci.. Washington.)
Pospelova-Shtrom, M. V. 19.35. On the svstematics of ticks of the genus Hacmapliy.satii C. L. Koch. Trav. Fil. Acad.' Sci. USSR, Tadzhik. 5:20.5-217.
Robinson, L. E. 1926. The Genus Amhhjomma. Ticks, a monograph of the Ixodoidea. Part IV. p. 1-302. Cambr. Univ. Press.
Saito, K., T. Iijima and ^L Minai. 1969. Studies on the effect of several insecticides against Argas jiiponiru.t. Jap. ]. Sanit. Zool. 20(1):39-41. (Japan-
184
Bkk;ham Young University Science Bulletin
ese text. English summary. )
Saito, Y. 1955. On iive species of ectoparasites on bats {Piptsiri'llux ahnwnis) collec-tcd in Niigata City, Japan. Niigata Igakkai Zasshi 69(1):7-12. ( Japanese te.xt. )
Saito, Y. 1958. On (he intenial organs ol lanal, nymphal, and adult stages of the tick, Ixodes persulcatus perxulcutu.'i. Jap. J. Sanit. Zool. 9:70. ( Japanese te.xt. )
Saito, Y. 1959a. Studies on the transovarial trans- mission of Pasteurclhi ttilurcnsis in the three tick species ( Ixodes pcr\ulviitu\ persulcatus, Ixodes japotierisis. and Uaemaplujstdis jluvti). Jap. J. Bad. 14:565. (Japanese te.vt.)
Saito, Y. 1959b. Studies on ixodid ticks. I. On ecology, with reference to distril:>ution and seasonal occurrence of ixodid ticks in Niigata Prefecture, Japan. Acta. Med. Biol. 7( 3 ): 193-209.
Saito, Y. 1960a. Studies on ixodid ticks, II. On the rearing and life history of three tick species ( Hac- ni/ipliiisalis flaia. Ixodes persulcutus persulcatus, and Ixodes japoncitMs) in Japan (Acarina: Lxodi- dae). Acta. Med. Biol. 7(4):303-321.
Saito, Y. 1960b. Studies on Lxodid ticks, IV. The internal anatomv in each stage of Hdemapht/salis flava Neumann', 1897. Acta. Med. Biol. 8(3): 189-293.
Saito, Y. 1962a. Studies on ixodid ticks. VI. Addi- tional report of tick rearing, with a presentation of a new tick container. Acta. Med. Biol. 10(2): 127-146. Ilhis.
Saito, Y. 19621). Isolation of Pasteurclhi tulurensis from ticks and chiggers panusitic on tularemia- wild hares, with some tularemia transmission ex- periment. Acta. Med. Biol. 10(2) : 147-159. Illus.
Saito, Y. 1964. Studies on ixodid ticks, VII. Notes on the ticks infesting badgers in Japan with a description of Ixodes tauuki n. sp. Acta. Med. Biol. 12(l):59-66.
Saito, Y. 1967. Studies on ixodid ticks, IX. Ixodes inotuispiuosus u. sp. from man in Niigata Pre- fecture, Japan ( Ixodoidea, I.xodidae). Acta. Med. Biol. 15(2): 107-112, Fig. 9.
Saito. \. 1969. Studies on ixodid ticks, .\. Ilacnui- pht/salis me^aspinosa n. sp. ( Ix(xloidea, Ixodidae) from Kanagawa Prefecture, |apan. Acta. Med. Biol. 17( 1 )':87-96.
Saito, Y., M. Kuhota, A. Y.^jima, T. Wat.^nahe. .\nd K. Kamino. 1965. Studies on ixodid ticks, VIII. On Haemaphijsalis hi.spinosa Neumann, 1897 in Niigata Prefecture. Japan, with some supplemen- tary observation on bovine piroplasmosis. Acta. Med. Biol. 13(2): 143-159. Illus.
Saito, Y., and S. Ohara. U)61. Studies ou ixodid ticks. V. Further studies on the reactions of the skin of laboratorx animals to the bites of immature ticks. Acta. Med. Biol. 9(l):l-32.
Saito, Y. S. Ohara, and T. Unac.-vmi. 1960. Studies on ixodid ticks. III. Comparative observation on the histological changes of host tissue caiised b\ tick bite. Acta. Bed. Biol. 7(4)323-329.
Sautet, J. 19.36. Invasion domicilaire de Ehipiceph- alus saufiuineus et de Teutana truin<iulosa. Role ixodiphage des araignees. hxm. Parasif. hum. comp. 14(2): 126-129.
ScHULZE, P. 1930. t)ber einige Verw.uidte \nn Ixodes ricinus L. aus Osta.sien. Z(X)1. Anzeiger. 90:294-303.
.ScHULZE, P. 19,33. Die Arten der' Zeikengattinig Derrnacentor s. 1. aus Europa, .Asieii. und Ncu- Cuinea. Z. Parasitenk. 6(3) :416-431 .
ScHiL/.E. P. 1935. Zur Zeckenf:uina Formosas. Zool.
Anzeiger. 112:233-237.
ScHULZE, P. 1938. Ober die "bipolar" Zecke Cera- tlxodes uriae ( White) =;)iif».i ( Pick.-Cambr. ). Zool. .\nzeiger 12,3( 1-2) : 12-17.
Sharif, M. 1928. A revision of the Indian I.xodidae with special reference to the collection in the Indimi Museum. Rec. Indian Mus. .30( 3) :217-334.
Sharif', M. 1938. Diseases transmitted by the Indian species of ticks and the possibility of their preven- tion through biological control. Ind. J. Vet. Sci. 8(4):3.53-.366. (reprinted 1939.)
Shioamori, M., K. Aso. and A. Yajima. 1953. On the Ixodidae in Kumamoto Prefecture, Kvushu, Japan. Nihon Juigakkai Zasshi (Jap. |. \'et. Sci.) 6:290-293. (Japanese text.)
SoMov, G. P., ;uid V. I. She-stakov. 1963. On spon- taneous infection of Ilactnaplu/salis japonica doup,- lusi Nuttal and Warburton ticks with D. sibiricus rickettsiae in Primorsk region. J. Microbiol., Mos- cow 40(12):51-56.
SuGiMOTo, M. 1935. Note on the Formosan ticks,
1. Taiwan no Chikusan 3(9):Suppl. 1-33, 3 PLs. (Japanese text.)
SuGiMOTO, M. 1936a. Note on the Formosan ticks, II. Taiwan no Chikusan 4(1):1-11. (Japanese text. )
SuciMOTO, M. 1936b. Note on the Formosan ticks, IV. Taiwan no Chikusan 4(.3):l-20. (Japanese text. )
SuGiMOTO, M. 1936c. Note on the Formosan ticks. J. Cent. Soc. Vet. Med. 49(7 ) :577-.590. (Japan- ese text. )
SuGiMOTO. M. I9.36d. On a new v.irietx ( Haema- phi/salis cornif^eru \ar. taiwana n. var. ) and a new species ( Ixodes taiwanensis n. sp. ) of ticks from the Formosan domesticated animals. J. Soc. Trop. Agric. Taiwan 8(4) ;336-346. ( [ap.mese text. )
SuciMOTO, M. 1937a. Notes on the ticks in the Formosan mountain reservation for the Aborigines. J. Cent. Soc. Vet. Med. .50(5) :.30.3-,3.3.5. (Japanese text. )
SuciMOTO, M. 1937b. Notes on the ticks in Kvushu and Okinawa islands. J. Cent. Soc. Vet. Med. 50 (9):597-615. (Japanese text.)
SuciMOTO, M. 19.37e. On the uvmph and larxa of And>lijomma testudinuriuin Koch. 1844. Trans. Nat. Hist. Soc. Formosa 27:160.
SuciMOTO, M. 19.37d. Studies on the Formosan ticks (9th Report). (On the tick, Haeniaphi/salis uishi- ijatnai Sugimoto, 1935). J. Soc. Trop. Agric. Tai- wan 9:287-290. (Japanese text. English summary.)
Sui'iNO, F. 1897. Nuovi Ixodes della Birmania. (Nota Preventiva). Atti. Soc. Veneto-Trent. di Sci. Nat.
2. Ill, f:i.sc. 1:2,30-2.38.
Suzuki, T. 1930 Notes on the male of a tick, Ixo- des affiuis. I.ansani.i 2(16):90-92. (Japanese text. )
Tanaka. H., K. Sato, Y. Sugita, J. Okabe, Y. Saka-
GAMI. T. CiIIKATSUNE, AND T. YoSHIDA. 1960.
On a tick para.sitizing the human perianal region. J. Osaka City Med. Cen. 9(3) :743-744. (Japanese text. English summarw )
Thorell, T. 1883. Descrizione di alcuni .\r.niiidi inferiori deirArci|ielago Malese. .Ann. Mus. Civ. Storia Nat. Genoa (1882-a3) 18:21-69 (not pub- lished in 1882).
Tokishige, H. 1911. Report of in\estigation on en- demic diseases of cattle in Kumamoto Prefecture. J. Cent. Soc. Vet. Med. 24:6-12, 25(3). (Japanese text.)
Biological Series, Vol. 15, No. 1
Ticks
185
TouM.\NOFF. C. 1944. Les Tujue.s ( I.xodoidea ) de rindtxliine. Institute Pasteur de I'lndochine, Sai- gon, 220 p.
Trapido, H. 196.5. Notes on eritical Asian Hacma- physalis speeimens in Euro{>ean museum collec- tions, with designations of lectotypes and a neotype (Acarina: I.xodidae). Proc. Ent. Soc. W;Lsh. 67: 152-165.
TsucHiE, v., A. IsniK.wvA, AND M. Okui. 1966. Stud- ies on ticks in pasture (II), (III). Report of the Institute of .\ninial Health of Shimane Prefecture. 13:1-15. (Japanese text, mimeographed.)
UcHiKAWA, K., AND A. Sato. 1968. Tarsal cliaetota.w of Argus japomcus Yamaguti, Clifford, and Tipton, 1968. (Ixodoidea: Argasidae). Jap. J. Sanit. Zool. 19(3):157-161.
UcHiK.AWA, K., AND A. S.\TO. 1969. The cK'currence of Argus japonicus and Ixodes lividus in Nagano Pref., Japan. J. Med. Ent. 6(l):9,5-97.
I'cHiKAWA, K . .\. Sato, and M. Kucimoto. 1967. Studies on the argasid infesting the Japanese House Martin, Dclichon iirhica. Med. J. Shinshu Univ. 12(2):141-1.55.
W'ARHrBTON, C. 1908. On some new and obscure species of tlie genus Hacmaphystdis of the Ixodi- dae. Proc. Cambr. Philos. Soc. 14:508-519.
\\'.\rburton, C. 1913. On four new species and two new \arieties of the ixodid genus Hncmaphysalis. Parasit. 6(2) : 121-1.30.
W'ahhurton, C. 1932. On five new species of ticks (Arachnida, Ixodoidea). Parasit. 24:.5.58-,568.
Warburton, C, and G. H. G. Nuttall. 1909. On new species of Ixodidae with a note on abnormali- ties observed in ticks. Parasit. 2:.57-76.
White, A. 1852. Appendix. Journal of voyage in Baffin's Bay, by P. C. Sutherland, 2:210.
Wilson, S. V,. 1946. Sea.sonal occurrence of Lxodi- dae on cattle in Northern Province. Nyasaland. Para-sit. 37:118-125.
Yajlma, a. 1942. On the Ixodidae collected from Iwate and Aomori prefectures. J. Assn. Anim. Hyg. 10(6):499-510. (Japanese te.xt.)
Yajima, a. 1950. On tlie forms of female and each stage of Haenuiplu/S(dis ias Kishida. 1936. Jueki Chosasho Kenkyu Hokoku 25:197-200.
Yajima, A. 1955. On two new species of ticks ( Ixo- didae) from Iwate Prefecture. Jap. J. Sanit. Zool. 6(1) :52-53. ( Japanese text. )
Yajima, A. 1963. On the distribution of the i.xodid ticks on the cattle in the pastures in Japan. Jap. J. Sanit. Zool. 14(2): 103. (Japanese text.)
Yamaguti, N., C. M. Clifford, and V. J. Tipton. 1968. Argas (Argas) japonicus new species, as- sociated with swallows in Japan and Korea ( Ixo- doidea, Arga.sidae). J. Med. Ent. 5(4) :4.53-459.
Yamaguti, N., and S. Inatomi. 1961. On the mor- phologN' of larva and nymph of Argas species col- lected in Okayama Prefecture, and some additional observations on its biology. Jap. J. Sanit. Zool. 12(2): 142. (Japiuie.se text.)
ZuMPT, F. 1940. The Rhipiccphalus species of the USSR. An account of variabilitv in the sanguineus group. Z. Parasitenk. 11:400-409.
ZuMPT. F. 19,50. Preliminary sttidv to a revision of the geniLs Rhipiccphalus Koch. Docum Mocambiciue 60:. 57- 125.
ZuMPT, F. 19.52. The ticks of sea birds. Australian Natl. Antarctic Res. Expedition Rep. Series B. 1:12-20.
APPENDIX 1
Notations on the Literature Cited
Yajima's (1956) list was often referred to as a source of information by Keegan and Toshioka (1957). Since it is a mimeographed note and has not been published yet, it is excluded in this paper.
Serial reports by Sugimoto entitled "'Note on the I ormosan ticks, part 1, 2, 3, 4" were published in Taiwan no ( hikusan .3(9). 1935,4(1), 1936,4(2), 1936, 4(3), 1936, re- spectively, and these four papers were at the same time published in Bull. No. 118, 1936, Dcpt. Agric. Govt. Res. Inst., 1 orinosa, Japan. Likewise, another of his papers < 1936d) appeared both in the Taiwan no Chikusan 4( 1 1 ), pp.
1-11. 1936, and in the J. Jap. Soc. Vet. Sci. 16. pp. 1-10, 1937. 1 iirthermore, the paper of 1937a contains two papers (with the same title) published in Taiwan no Chikusan 5(3):l-25, 1937 and 5(4):1-15, 1937. His paper (1937e) is also found in J. Jap. Soc. Vet. Sci. 16( 1), pp. 11-16.
The paper "Studies on the ticks of the domestic animals in the Far East" by Itagaki. Noda, and Yamaguchi ( 1959) is an English translation of their 1944 paper, and the contents of both are identical, except that H. jezoensis is synonymized under H. japunica and spelling of Korean localities is amended so as to be up to date in the later (1959) publica- tion.
186
Bkigham Young University Science Bulletin
APPENDIX 2 Collection Records '°
Species
Host or llabitjt
Area
Preli'Ltutc Locjluy
Collection or Literature Data
Argas japonicus Di'lichon urhica dasypus (House Martin)
Japan
llirttndo daurua japontca I Striated Swallow!
Nagano Agcmatsu-cho
Matsuinoto-shi
Iwatc
lwai/-umi-cho
Dcliihon urhua dawpus (House Martin)
Tt)chigi Chu/enji
Tokyo lacliikawa M H
Shiniane Iwannod.i-shi
! ukushinij Shinohu- r.ika> u
Oka>ania
Totlori Miyazaki
Gifu
Okayama
Nagano
Nakanosawa, Inawashiro
Yonago-shi
Takachjho-cho
Mitakc, Kani-gun
Nnnii-shi
Agcmatsu-cho
*66-J-0032.Coll. Toshioka. 17 Jul •66(42 I .•»4 M. 227 N, 39 I. 2 F. 2M. 21).
Coll Vamaguli, 7 Jul '66 (5 1 . 5 M. ION. 10 L).
Coll. Sato. 1 3 Sep '66 (5 1 , 5 M, 5 N).
#66-J-0()43.Coll Yaniagud. 7 Aug •66. IWH I , 17H M. 130 N.J/-.
2 M. : 1 1
*66-J-()()47.Coll Hatovanui. 24 Aug '66. (ION).
■C:66-1-II04S. coil, lloluhcc & Sugl- yania. 24 Jun '66. (36 I . :ili M. 153 N. 25 L, J/-'. 2M)
#66-J-O()50. Coll. Holubec & Sugi- yaina. 7 0ct'66. (16 I . 11 M.47 N).
»6)(-J-(l336. Coll, Suzuki & Sasa- gawa. 10 Jul '6S. (700 I . 600 M. 1.100 N),
#6X-J-0337. Coll. Suzuki & Sasa- gawa. 12 Jur6X.(4 N).
#66-J-()(l37. Coll. Mizusawa & llatoyama. 21 Sep ^66. (89 1 . 57 M. 108 N. 34 L.J/-', 2M.2 I.).
Coll, Yainaguti. Sep. ^60 (5 I .
3 M.4 N. 7 L).
#66-J-()()53. Coll, Mizusawa & Haloyama. 1 9 Sep '66. (45 1 . 62 M,89 N.J /-, J.W).
#66-J-()044.Coll, roshioka& Suzuki. 13 Scp^66. (63 V . 30 M. 496 N. 5 L.J/-. J.Ul,
^66-J-()()62. Coll, Tanaka. IK Oct '66. I I Nl.
Inatomi* "iamaguti. 1960:17. Yamaguli & Inalomi. 1961:142 Coll. Yaniagull. Sep "60. (I . M. N. Ll.
Har.i. 196.t. (oil. Ilara i^\ pirsitiis).
Sleeping tpiarters
Korea
Kyongsang- pukto
Matsunioto-shi. Hakuha-mura. Ageinatsii-cho. Akasliin.i-clio. 1 ukusliinia-cho. Hotaka-etio. lida- shi. Nagano-shi. Okaya-shi. Otari- niura. Saku-shi. Sakai-niura & Ueda-shi
Taegu
Uchikawa. 1969:96. Coll. llchikawa. 20 Mar '67 - 7 Apr '68. (A. N. 1 ).
YaniaguO. Clifford & Tipton. 1968: 458. Coll. Tibbets. 20 May '53. (51.1 M. 3 N).
:'(1 iiiituhcrs in iwiientlicscs inilicale numlH-r of spt'< inions sent In Mr (.len .M Knliis fni i.li'iilifu.itiiin
Biological Seuiies, \'ol. 15, No. 1
Ticks
187
Argas japoiticus Swaiinw nt-sl
Argas Vesperiilio \upiraiut^
icsficrfilHinis (Irostcd Bat)
Ornithodoros i-api'iisis
Korea Kyonirvinj;- Shin C lion l)i
pukto
lj|Mn I ukusliiin.i Slidji!, Bango-cho
|
Pipisrrcllus ahramus (Hal) |
" |
1 ukuoka |
" |
|
Pipistrclliis savii vclox 1 Uall |
" |
Hokkaido |
Sapporo-shi |
|
Pipislrclhn ahramus {Hall |
" |
Niipata |
Niigata-shi |
|
Mvotis sp. (Hat) |
Korea |
- |
KunKhon |
|
l:pfcsi(ii\ (orccnsis (7) (Big Brown Hat) |
»• |
- |
Syani. Heijo |
|
Hats |
Japan Korea R\uk\ u Islands |
" |
" |
|
Man |
Japan |
"loiton |
- |
|
Lan4s crassirostris (Black-Tailed Gull) |
Aoniun |
Kabujima, Hachinohe-shi |
Sliinianc Kyojima.
Hinomisaki
Vamaguti. lllllurd & I iplon. I96S: 45K.roll.Tipton. 9 Jul'55. (2 1 . I M, 12 N. 2 L).
#66-J-l)lll2.(oM Kaneko. auliinin '65. ( 171).
#67.J-O215.(oll 'laniaguti, I 1 Aug '67 (23 h. II M, 15 N. / F.
2 M. I N).
27 Jiin '67.(23 I.)
Coll. Hatlori. 21 Aug '63. ( I L).
Saito. 1955:7-12. Toll. .Sailo, 30 Apr-IONov'52. (137).
'I'amaguln.-l al.. 1968:453. Coll. llvong-.Sun All, 17 May "57. (9LI.
Klshlda. 1936 142 (asCam Vf^pirtlllnnis llL-iIo-Pyong>aiig).
KlvhlJa. 1927:9X6.
Kanio. 1962:173.
Coll Tosliioka & Akiyaina. IS Aug
'55.
Coll Akiyaina, 14 Oct '55.
#66-J-0055. Coll. Toshioka & Sasagawa, 15 Aug '66, ( 105 i\ 1 M, 36S N)
#6 7-J-0(193. 12 Mav '67, (242 I. SON)
#55-J-()54, Coll. Tovhioka & Aki- yama, 17 Sop '55, (Approx. 100 A
& Nl
A mhlyomma gcocmydac
CaUmcctris Icucomcias (Streaked Sllearuaten
Crevices of rocks near nest of I. ants crassiro- slris (Black-TailedGull)
l.ants crassirostris (Black-Tailed Gull)
CaUmcctris Icucomrlas (Streaked Shc.irwaterl
Syilthlihoratiiphus antiquus { \n<y^ni Auk)
Crevices of rocks near nests of t.ants crassiro- stris (Black-Tailed CJull)
(U'in-myda spcflglcri japotiica ( lurtle)
Ryukyu Islands
^'ainagata lobishima,
Sakala-shi
Shunanc ()hakajuii,i.
Chibu-gun
Aoinori Kahujima,
Hachinolie-shi
Tokyo ,Aogashuiia
Kanagawa Hayaina-clio
Shiniane Kydjiina
Yona, Okinawa
»66-J-0054, Coll Toshioka & Betchley, 9 Jul '66, (96 1 , 3S M. 129 Nl
#68-J-0332, Coll. Yamaguti & Blccha, 4 Jun '6(i, (51) I , 63 M, 276 N. 2 L).
#68-J-0335. Coll. Suzuki & Sasa- gawa, 7 Jul "68. (I f, I M, 1 N).
Coll Yjiiijguli, I 3 Sep '66. |4 1.1.
Keegan & loshioka, 1957 23. Coll. Toshioka & .Akiy.iiii.i, 18 Aug & 14 Oct '55.
Asanunia, 1960:94 Coll ■\sanuiiia, Jul '55
Asaniuiia, 1960:94. Coll. .'\sanunia. I 5 Jul '58,
Asanunia, 1960:94. Coll. .\sanuina, 14 Mar '58.
Kohls. 1957a:85- 104. Coil, Keegan. 16 Sep '55. (A. N).
:;r66-R-OOI6. Coll, Mi?usawa. I Jun '66. (J.Wl.
188
Brigham Young University Science Bulletin
Amblyomma geoemydae
Cyclemys flavomarginata flavomarginata (Turtle)
Land torioisc
Ryukyu Islands
Gosj-dukc, Iriomolc (Iv)
Okinawa
^66-R-OOI7.('oll. Mizusawa, 19 Jun "66.)/ h'. 10 \. 14 I ).
-ft6-R-U006. Coll. Pennington. Oct 'bt.{4 N. 2S I.).
Gcoeniyda spcngleri japomca (Turtle)
Clemmys japoniva
(Turtle)
Cyclcrriys flavomarginata flavomarginata (Turtle)
Japan
K\ ijk> u Islands
Kagoshinia
^ una. Okinawa
Kunijranii. Tane- gasl ima
■=6N-R-(I()15. Coll Mi/iisawa. Apr "68. (11 N).
Keegan & Toshioka. !957:S. (as A. malavanum) Coll. Toshioka. 19 Jur55 (1 1 . 1 M, I N).
I'utanaka. Ishi- Kecgan & Toshioka. 1957:8. (as
gaki (Is.) .1. malavanum) Coll, Keegan &
loshioka. 6 Nov "55. (I I . 1 M).
Hanoshiro. Ishi- gaki (Is I
Keegan & Toshioka. 1957:8. (as .1 malavanum) ("oil. Keegan & loshioka. 27 0e( *55. ( 1 Nl.
Gosa-dake. Iriu- Kaw.ishima. 1963:103. las,l,
mote (Is.) malavanum) ("oil. Kawushima. 14
Aug'62. (9I-.4 M. 3N. 1 L).
Keegan & Toshioka. 1957:8. (as A. malavanum) Coll. Keegan & Toshioka. Nov '55. 406 MCil -IHI. ll N)
(icocmyda spcuglcn (Turtle)
Yona. Okl^.l\^a K.iwjshniia. 1963:103. (as.-l.
malavanum) Coll. Kawashhiia, 6 Jul "62.(3 N. n L).
Turtle
West ot Arakawa. Okinawa
Keegan & Toshioka. 1957:8. (as^l. malavanum) Coll. Hubert. IS May '55. 406 MtU -25. (6 N).
Amblyomma nitidum
Man
l.aticauda scmifa- sciata (Sea snake)
Mountain near Keegan & Toshmka. 1957:8. (as.4.
Ishigaki ( ity. malavanum) CoW Keegan & Toshioka
Islugaki (Is.). 27 Get "55. ( I N). (Probably a
stray troni the turtle ot 27 Oct '55 eoUection examined the same day).
Khigaki (Is I
-66-R-(XH)8. Coll, Toshioka & Keegan. 10 Nov '62. (/ /■')-
^66-R-nf)09. Jun '57. (/ F)
A mblyomma Sus scrofa leucomysiax
testudmarium (Wild pig)
Japan
loehigi
Ryuk\ u Kagoshima
Islands
.Ashiken. Anianii Oshiina
\'u\van. Aiiuiini Oshinia
=6(( J-045.(oll Aki>ama. Nov '60.1/ h\ n.M)
#67-J-0246. Coll. Mizusawa. 20 Sep '67. (2 I . I Af)-
-=^67-J-0256. Coll. Mi/usjwa. 1 Oct "67.(3 I . / /-". / M).
Domestie pig
Ashiken. Anianii Oshima
Islngaki (Is.j
'I'uvK.in, Am.inii
OshlMKl
^67-J-0272.ColI Mi/usawa, Apr '67. ( 1 M).
#6«-J-()310. Coil Mi/usuwa. 15 Mar 'bH.{7 F).
*68-J-0318. Coll. Mi/usawa, 1 1 Apr '68. (2 I . 2 M, 4N.5 L. 3 N. 5 I. ).
^670-0271. Coll. Mi/usjwa. 19 Sep *67. (/ ;V).
^55-R-()17. Coll. Keegan & loshi- oka. 4 Nov '55.(7 F. 5 /.),
^65-J-()OOI, Coll. Hajime, 13 Dee •65.(J/-", J M)
Biological Series, Vol. 15, No. 1
Ticks
189
Amblyomma Ifstudmanum
DomestiL- pig
Ryukyu Kagoshimj
Isbnds
Cenus nippon (Sika Deer)
Domestic dog
Penialagiis Jurnessi (Ryukyu Rabbit)
Vegetation
Yuwan. .Anianu Oshima
Ukenson. Atiiami Oshima
Ibaraki
Shizuoka Misakubo-cho
Ryiikvu Ragoshiina \'u\van, Amami
Islands Oshima
„ „ Ukenson. .Amami
Oshima
Yuwan . AmamI Oshima
LIkenson, Aniami Oshima
Yuwan. Amami Oshima
Japan
#65-J-0002. Coll. Ila)nne. Nov '65, I / \U.
#67-J-0067. Coll. I ipton & Mizu- sawa, 18 Mai '67.(1 h. 12h L. 5 F. 5M. 19 1.).
#67-J-()073, Coll, Tipton & Mizu- sawa. 19 Mar'67.(/ F. .^M).
#()7-J-0077.C'oll. I ipton & Mi/u- sawa. 1,1 Mar '67. (//•'. / A/).
#67-J-OI83. Coll. Yamaguti. 22 Mur'67,(/Af).
#67-J-0293. Coll, Su/.uki, 4 Dec '67, (/yW).
#67-J-0273. Coll, Mi/usawa. Apr '67.1/ F).
#67-J-00b9. Coll, Ttpton & Mizu- sawa. 24 Mar "67.(1 .16 L. 1(17 I.).
#67-J-0081, Coll. Hajnne. 1(1 F eb '67. (7 95/.).
#68-J-0314. Coll, Mizusawa. 5 Apr '68. ( / An.
#67-J-(J082. Coll, Tipton & Mizu- sawa. 23 Mar '67. (/ /.).
#67-J-0076, Coll. Tipton & Mizu- sawa, 23 Mar '67. (97i).
#67-J-0084. Coll. Tipton & Mizu- sawa, 24 Mar '67. (45 I.).
*68-J-0316. Coll, Mizusawa. 5 Apr '68. (JFl- #68-J■0317.Coll. Mizusawa. 10 Apr '68, 1/ .V. ,5/.).
#66-J-0031.Coll, Ucno. 28 Jul '5 7. (IF).
Man
Horse
Wild pig
Hyogo
Wild pig
Vegetation Domestic pig
Microhyla fissipes (= M. ornata) (Frog)
Wild pig
|
«• |
Kyoto |
- |
|
.. |
Osaka |
- |
|
•• |
Miyazaki |
Kamie. Koy u |
|
Ryukyu |
. |
Inoda. Ishigaki |
|
isiands |
(Is.) |
|
|
" |
- |
Inoda, Ishigaki (Is.) |
|
tt |
- |
Ohara. Iriomotc (is.) |
|
Japan |
Osaka |
Ikcdashi |
Ryukyu Islands
Tokyo
Ishigaki (Is I
Kawashima. Kanio Ai Ml\a/aki. 1960: 77-80.
Nagahana* Matsuo, 1962:119-120.
Janaka. ct al,. 1960 69-70.
Itagaki. Noda & Y'amaguchl. 1944 & 1959,
Kc-cgan & Toshioka. 1957:9- Coll, Keegan & Toshioka. 2 Nov '55. 406 MGL#82.(I M).
Keegan & Toshioka. 1957:9. Coll. Keegan & Toshioka. 12 Nov '55.406 MGL =98. < 2 I I,
Keegan & Toshioka. 1957:9,
Coll. Keegan & Toshioka. 406 M(;L
;i207.(l M),
Keegan & loshioka. 1957:9. Coll. Hiromatsu. 1 I Mar '42.
Kishida. 1922a:S50-851.Coll. Taguma.
Sugimoto. 1 937a: 3 1 7-323. Coll. Iwasaki. ( 1 L).
Sugimoto. 1937b:612-613. Coll, Iwa-aki. 5 leh '37.(1 M).
190
Bhicham Young University Science Bulletin
Amblyomma Wild pip
testudinarium
Green trog
Horse
Boophilus
micro plus
Domestic cattle
Ryukyu Islands
Japan
Ryukyu Islands
Japan
Miyazaki. Hyogo
Miya/aki
Kagushima
Nagasaki
KagitsliiiiKi
Iriomotc (Is.) Ishipaki (Is.)
Takanabc Anuinu Oshniia
CJonoura. Iki (Is.)
Hatsuyama, Iki (Is.)
Iwa/ato. 1 suvhiina
k liinan
Sugimoto, 1937h:612-613. Coll. Ohasi. Mar '37, II M).
Sugimoto. 1937b:6l2-613. Coll. Iwasaki, (1 L).
Nakamura & Yajima. 1937:174.
Robinson. 19:6:257. Jun & Oit '12 IN 2909a).
Robinson, 1926:257. Jun & Oil •|2(N 2919).
#67-J-lllll3. Coll. lipton 4i Min/u- sawa, 2 May '67, 16 1 i.
#67-J-0107, Coll. Tiplon & Mizu- sawa, 4 May "67, {3 M. J .V).
*67-J-0l IS. Coll. Tipton* Mizu- sawa, 6 May '67, 1 /.^ F. 7 M. I J ,V).
#67-J.|li:4. ( .111 1 iploii i Mi/u- sawa. Ill May '67. IK I, J/-, .5 M. 4 ,V).
#67-J-OI 25, Coil, Tiplon & Mizu- sawa. 10 May "67, 1 1 I, 5 F. 4 .V).
#67-J-OI26.Coll hptonii Mi/u- sawa. 10 May "67,17 I , // /■',
: M I .V).
#67-J-l)l 28, Coll. liplon & Mi/u- sawa, 10 Mav "67, I 12 I-, .V/-',
#67-J-l)l2M.( oil. lipton & Mi/.u- sawa, HI Ma\ '67. 14 I . h /■).
W.7-J-III 31, Coll. Tipton & Mizu- sa»a. 10Ma> '67,119 I . 7 /•', 71/. 4 \\
#671111 32, (oil. I ipton & Mizu- sawa, II) Ma\ '67, lA'F, 4 M).
#67J-ni 34, Coll. lipton &. Mi/u- sjw.i. III M.n '67,15 1. :i /■', SM, 4 \)
#67-J-0135, Coll. Tipton* Mizu- sawa, 10 May '67,(2 V.jh: JM. 4N).
#67-J-0 137, Coll. Tipton* Mizu- sawa, 10 May 'bl. iiO l . 25 F. V M. / ,V1.
Sakura Jinia
Ryukyu Islands
,Sunii>o, AiiKimi Oshinia
Burcn, .Amaini Oshiina
Yuwan. .Xniami Osliillla
??67-J-013»(, Coll. Mizusawa, 14 May '67, i \\ . 4 F. I M].
#67-J-014ll. ( oil Mi/iis.iwa, 14 May '67, 1/ F)
#67-J-OI4l,Coll. Mi/usawa, 14 May '67,(/7K 5 M).
J?67-J-014:. Coll Ml/lls.iw,i. 14 Mav '67, |7 I , I M, I N, J F. 1(1 V. I \).
-67-J-()064, Coll. Tiplon & Mizu- sawa, 22 Mar '67,124 I ...'() /•, -V.AH.
=67-J-0252, Coll. Mi/usawa, 2X Sep '67, I 15 I , 3 M. /(;/■'. t, M).
.=67-J-026l, Coil, Mizusawa, 3 Oct '67,(93 1-, 3M, 3N).
Biological Series, Vol. 15, N(
Ticks
191
Hoiiphiiiis tiikroplii\
DonicslK caltic
Inlands
Mivara. Khigaki (Is.)
=h(.-R-(l(l(i:. (nil, HnlulK'c & Don- ccllc, 2 Nov '(i(>. (,V7/-', l,S At).
Huhalus hu halls (.^siatK WaUT Hullalo)
Donicsdi. ^hccp Hoitphillis spp. Cattle
Cattle, water buttalo. horse, deer, pheasant
Cattle
Horse and duniesiic cattle
Horse, cattle, domestic sheep. ,'\ sialic Water Buttalo
Korea
R> uk\ 11 Islands
Korea
Cluingihont;- pukto
Hirai. Ishigaki (Is.)
Kurnik. Cheju Do
Macng-dong Myi>n
Uni-Song Klin
Hir.ii. Ishigaki (is)
Isodong. Cheju Do
=?66-R-()00.1, Coll. Doucette & Su- glyama. 2 Nov '(id. (s'.s /■, _\S A/).
#f.6-R-0004. Coll. Doucette & Ho- luhec, 2 Nov '66, (^.5 F. 4.i AT).
-hV-R-OOl l.Coll Tiplon. K I eh '67. (15 1 . ,s M. 2 N)
#67-R-()IILl.C oil. 1 ipton. (. May "67, (X I . i Ml.
*h7-K-lll)19. Coll. Parsons, ft May '(.7.1/ ,V|.
#h7-K-()029. Coll. Cantello & Town- send, 12 Mav '67. (9 N. / F. / M.
I .V).
#(>7-K-(10.V). Coll. Cantello & I own- send. 12 May '67. I .^ I . 9 N. / M).
#67-K-(X)3 1 . Coll. Cantello & Town- send. 12 Mav '(.7. 1 2 i: 7 N. / K / ,Vi.
#h7-R-(l(ni), C oil. Tipton, 9 I eh '67, ll I 1 . 9 M, 2S N).
#67-R-0(ll2, Coll. Tiplon. S I ch '67.(1 I 1.
#67-K-(l(122. Coll, Unggers, 6 May '67. (/ /■■).
|
Japan |
Saga Kuniamoto Olta |
Tokishige. 1411. {AS Boopfitlus annulatus caudatus) cited by Sugimoto, 1937b. |
||
|
Japan |
- |
Southern |
part o( |
Kishida. 1927:985. {as fi. |
|
& |
Kyushu |
cauiiatus Neumann). |
||
|
Ryukyu |
||||
|
Islands |
||||
|
Korea |
S. Kcisyo |
Nanrai. S |
. Keisyo |
Kishida. 1936: 139-142. (as |
|
(-Kyong- |
and coast |
region ol |
Palpob<>oplnlu\ minmn^). |
|
|
sangnamdo) |
S. Korea . |
& Cheju Do |
||
|
Jjpan |
Kuniamoto |
A so |
Ogura. 1936:75. (as /i. |
|
|
a. |
anmilatus caudatus) cited by |
|||
|
R\ ukyu |
Sugimoto. 1937c. |
|||
|
Islands |
||||
|
Japan |
lukuoka |
. |
Nakamura& Yajima. 1937:133- |
|
|
K |
Kagoshima |
iS4. (as Vroboophilus caudatus. |
||
|
Ryukyu |
Nagasaki |
Urohoophilus sinensis. P. |
||
|
Islands |
Kuniamoto Hyogo Yamaguchi Okmawa |
minmngi) N. and Y. cited Minning's record trom Miyazaki Pret". as known distribution. |
||
|
Japan. |
Hiroshima |
Kishida. 1939a:538-552. (as |
||
|
Rvukyu |
Miya/.aki |
i rohoophdus caudatus, U. |
||
|
Islands |
Kumamoto |
sfianfi. U. sinensis. Palpo- |
||
|
& |
Kagoshima |
hoophihis hrachyurus). |
||
|
Korea |
||||
|
Japan |
Kagoshima |
Tanegashi |
inia |
Kishida. 1929:20. (as fl. |
|
& |
annuiafus caudatus) De*. "24 |
|||
|
Ryukyu |
Okniawj |
|||
|
Islands |
||||
|
Japan |
Nagasaki Kuniamoto Kagoshima |
Itagaki. Noda &. Yainaguclii. 1944: 33-39. 19.S9:4I.(as/;. caudatus). |
||
|
Korea |
Kyongii-do Kyongsang- pukto Kyongsang- namdo |
192
Hhicham Young University Science Bulletin
Boophilus spp. Horse, cattle. domesiiL
sheep. Asiatic Water BuITjIo
Korea Cholla-
nanido
Itapaki. Noda & Yamaguehi. 1944. 33-3'J, 1959:41 (usfi. cauJatus).
Cattle and horse
Cattle, horse and water butTalo
Cattle
Horse
Jupan Kuinainoto
Ryukyu Islands
Korea K\ongii-do
Japan lokyo
Miyazaki
Ishigaki ( Is.), Inomote (Is.), Yonakiini (Is.). Matenima, Kobarna
Seoul
Shigemori. Aso (S: Vajima, 195.V 290-293. (as Vroboophilus cQudatus and Palpohoophilus brachyuhs).
Keegan&'Ioshioka. 1957:10. las Boophilus niicropliis).
Nultall & Warburton. 1915:433. (as Hoopfulus sp.. tound with //. hisponosa = H. longicornis. 16 Oct "11. submitted by Dr. Miyajima).
Neumann. 1897:413. (as Rhipiccphalus atmulatus cauJatus) (23 r, 2 M).
Dermaccntor spp Wild pig
Chiba
Kiyosumi-vama Kishida. 1922a: 847-851. Coll.
Aoki. 30 Oct '09 (as auralus auraiwi) (1-, M).
Domestic pig
Korea
Tokyo
Kishida. 1922a:847-851. (as rclicularus rcticulatus) (I-. M).
Kishida. 1922a:847-85 1. (as rcticulatus reluulatus) (I . M).
Horsi
Japan Iwute
Selcnarctos tiuhctanus
japonicus
(Asiatic Black Bear)
Wild pig
Korea
Japan
Iwatc
Hanig\ *uig-
piiklu
Shi/uoka
Kyoto
Keigen-gun. N. Kankyo ( = Hamgyong- pukto)
Taniura, Tayama- mura, Johoji- mura
Kishida, 1936:142. las/;. vancgaius). .According to Kishi- da. Mr. Asakawa recorded this species.
Nakanuiraii Vajinui. 1937:162-164. {dsreticulalus) li . M).
Terada. h^atc•gun. Yajima. 1942:499-510. (as
Okunakayania. Ninohc- rcticulatus) (1. M).)
gun.
Johoji-mura, Ninohe-
gun.
Arasawa. Ninohc-gun.
I utaniata Pasture.
Sannohe-gun
Kamikita-gun
Shimokita-gun
etc.
Yajima. I 942:499-5 I 0. (as rcticulatus) (W M).
Tcmmabayashi- mura, Kamikita- giin
Takizawa. Iwale- gun
Unggi. Kyonghung
Oshinia-inura, Sunto-gun
Shi/uhara
Itagaki. Noda & Yamaguchi. 1944. 1959. (;is rcticulatus) (28 1-. 16 M) Aug& Sep '41 & Oct '42.
Itagaki, Noda & Yamaguchi. 1944. 1959. ids rcticulatus) (12 l. 10 MlOct '41.
Itagaki. Noda & Yamaguchi. 1944. 1959. as (-0^/^(61 I. 34 M) Jun *41 & 29 May-5 Jun '42. Ori- ginal description oi corcus. no indication ot type.
Keegan. & Toshioka. 1 957 : 1 0- M . Coll. Asanuma.(2 M). (labelled as auratus).
Keegan & Toshioka. 1957:10-11. Coll. Akiyama. 12 Dee '55. 406 MGL -101, closely resembling the above. Illustrated in pi. 14 ot K &. T.. reproduced. (2 M).
Kccgan & Toshioka. 1957:10-11. Coll. Toshioka & Akiyama. 29 I eb '5h. 406 MGL =194.(2 Ml.
Biological Series, Vol, 15, No. 1 Dcrrriaantor sp[v Wild pig Japan
Horse
Hacniaphysalis Man
cairifyantilala
Korea
Japan
Domestic dog
Domestic cattle and horse
Korea Japan
Korea
Japan
Ticks
Kyoto Kyoto-shi
Aoniori
193
Iwate
Uog
Niigata
Hokkaido
Tokyo
Chiba
Niigata
Saitama
Kanagawa
Shizuoka
Mic
Osaka
Hyogo
Kochi
Miyazaki
Kyongii-do
Nagasaki Niigata Ciunima Ishikawa Yam ague hi
Hamgyong- pukto Kangwang- do
Kumainolo
Kanugawa
Okunakavama
Yuki ( = Unggi)
|
Hyogo |
1 akarazuka |
|
Kanagawa |
Miura |
|
Chiba |
- |
|
Tokyo |
. |
Takada-shi
Kitanii. Abashiri
Narita YokcJgoshi-mura
Yokohama
Keegan & Toshioka. 1957:10-1 1. Coll. loshioka, 17 Nov '56, 406 MGL#I94, (1 M).
Keegan & loshioka, 1957:10-1 1. Coll Asaniinia, Jun '42. I I Ml, (labelled as rctiaitalus).
Keegan & Toshioka, 1457:10-1 I. Coll. Asanunia, Jul '4N, (I 11, (labelled as rt'ticutatusi.
Keegan & Toshioka, 1957: 10-1 1. Coll. Yajima, Oct '38. No. 2490 ( 1 I), (labelled dsreticularus).
Keegan & Toshioka, 1957:10-11. Coll. Yajinia, 14 .Sep '19 (1 Ml (labelled as rcriculatus).
Keegan & Toshioka. 1957:10-1 1. Coll. Yiijima. 16 Oct 'iH.iS 1 , 4 Ml (labelled j'. reticulatiis). llliislrated in pis. 12 & 1 3 of K. & T., reproduced.
Keegan & Toshioka. 1957:10-1 1. C oil Asanuma. Jun '42. (4 1 . .^ M) (labelled d^coreus). Illus- trated in pis, 10 & 11 ot'K. &T.. reproduced.
#41-J-002, Coll. Hiromatsu. 15 Jan "41.(33 b. I0M.4 N, / Aft.
*52-J-004, Coll. Asanuma. Jul '52. (2 b, 14 M, :F).
#67-J-01KK. 12 Jun '67.(1 N. / /■').
Keegan & Toshioka. 1957:12. Coll. Yajima, 15 Jur33, (2 M),
Keegan & Toshioka. 1957: 12. Coll, Yajima, 20 Jul '34. (1 1 ),
Keegan & Toshioka, 1957: 12. Coll. Kishida, 20 May '21.11 Ml.
Itagaki. Noda & Yamaguclii. 1944, 1959.
Nakamura& Yajima, 1937:133-184,
Shigemori, Aso & Yajima, 1953: 290-293,
Warhurton, 1908:508-514
Nuttall & Warburton, 1915:491-492, Coll, Owslon. Apr '02, (2 1 , 2 M). submitted by Rothschild.
194
Bhicham ^'(>UNc University Science Bulletin
llafmaphysalis Dog
campamtlata
Japan
lokyo
Nuttall& Warburton, 1915.491-492. Coll. Janson. I X94, ( *2XK6) (I A).
Nuttall & Warburlon. 1915;491-492. 18 Jun "14. submitlfd by Dr. Miyajima(#299l) d A).
Nuttall & Warburton. 19 l.^;49l•492. 1 I Sep '12. subniittfd by Or. Miyajima(*29l7l (I Al.
Hat- maphy sails cominna
Uokkaidu
Donicstii' cattle
DonicstK cattk' and horse
Horse
Dog
Korea
Japan
Ncinuro-shi, Neniuro
Alsukeshi. Kusluro
Tokachi. Kushiro, Ncmuro & Kitami
Hamgyong- pukto
Saga
Unggi
Kitaoka, May '64. {I h. J M). Kitaoka. Jul '6h, (/ h'). Ogura& lakada. I 927;2ll2-2(>4.
|
Korea |
- |
- |
Kisluda. 193h:l42. |
|
Japan |
Aunion |
Mlsawa, Kantiklta- gun |
Yajima. 1942. .SOS. |
Itagaki, Noda & YainagULlu. 1944; 1959.
Keegan & loshioka, 1957: 1 .1, Jul '4 1 (1 M).
Koch, 1S!(7;7S6. las;»TOj<))(l I).
Neumann. 1K97:.141. 1905:2.19. 1911: 111. {AS lunido. cnminna var, kochi and Kfiuinna kochi). Coll. Hilgcndorl iDcrl. Mus.) (3 I ). Coll. YaniagiiLlii. on dog. cited by Nuttall & Warburton. 1915(2 I ».
Hat'tnaphyialis Ooinestic cattle
sp. (//, CDrnigcra group)
Hactnaphysalia
lornificra or Uui'maphysalis lus
Tokyo
Hachijo Ko- shnna
Miyakc (Is.l
Nutlall & Warburlon. 1915:457. 1 xaninicd Neurn.inn's ( .1 1 ) in Berlin Museum, cited Neumann's record, 1905.
#67-J-0206. Coll. Yamaguli. 13 Jur67,(/ F. I M).
Keegan & Tosliioka. 1957:13. Coll. Kishida, 2 Oct '34,(1 M) ( labelled as ia\).
Keegan & Toshioka. 1957: 13. Coll, Yajima, 29 Jun '54, ( I 1 , I Ml (labelled as ias).
Keegan & Toshioka, 1957: 13, Coll. Yajima. I Jul, ( 1 M) (labelled as ias).
Dog Domestic cattle
Uirds
Ilaemaphysalis dociiitzi
Pheas;mt
Kagi>shima
Hachijo (Is.)
Miyake (Is.)
Hachijo (Is.)
.Akune
Keegan & Toshioka, 1957: 1 3. Coil, Asanuma. 29 May '49. (I I ).
Keegan &. Toshioka, 1957: 1 3, Coll. Asanuma, May '42, (2 M) (labelled as ws)-
Nakatsuji, 1942:295, (as ia.v).
Nakamura* Yajima. 1937: 1 33- 1 «4. Coll. Kishida. 2 Oct '35 (as i<j.vl.
Yajima. 1950 1 97-200. Coll. ,Sasa, Apr ■4X, (II) (,isMs).
Asanuma, 19h5a: I 12 (N, Ij,
#56-J-057, Coll. loshioka, I 0 Jan '56. (5 M. / F).
Biological Series, Vol. 15, No. 1
Ticks
195
H ac ma phy sails JoeniCi
Phasianus cokhicus rohusfipcs ( Pheasant)
Vegetation
Japan
Pheavint
Hacmaphysali^ Horse
flava
Dog
Nycterciitcs procyonottjes
liierrinus
(Raccoon Dog)
Vulpes v'ulpcs japomca (Red I-o\»
Sus scrofa leucomystax (Wild pig)
Ccn'us nippon (Sika Deer)
Domestic cattle
Vegetation
Niigata Shukunegi. Sado
(Is.)
Unie/u. Sado (Is.)
Kagoshinia Akune
Niigata
Aomori Towada
Tochigi
Mie
Iwate
Iharaki
Kuzuu
Towada
Kanagawa Yamakita-cho
Fukushima I ukushima-shi
Ohinado
Kanagawa Miyagasc, Kiyo-
Kawamura
Yamanashi Nohara, Doshi-
tnura
Miyazaki Tomisaki
Kyoto Hanase-mura
|
Aichi |
Tonc-mura |
|
Tochigi |
Kinugawa |
|
Shiga |
- |
|
Ihuraki |
- |
|
Mic |
- |
|
.Shi/uoka |
Misakubo-cho |
|
Iwala-gun |
|
|
Kagawa |
Zcntsliji |
|
Tokyo |
Hachijo (Is.) |
Saito, 1959b:20I.Coll.SaiIo. Sop. '56. (I 1-. 18 M, 32N. 74L).
Saito. 1959h:201. Coll. Saito, Sep '57 - Nov '58, (.1 I-, 5N, 60 L).
Saito, I 959b: 1 95. Coll. Saito. May-Oci '57,(1 M).
Kccgan & loshioka, 1957: 18. Coll. Toshioka, 10 Jan 5f), (5 M) (as //. sp. incertac ccdis. 1 1 ).
Kccgan & Toshioka. 19.S7 IS. 11 I).
#55J-055^ Coll. Akivama. I.I OcfSS.iJF. 1 M. .i^, I /,).
#56-J-059, Coll. Toshioka & Aki- yama, 23 l-eb'56, (/ /■").
#56-J-062, Coll. Toshioka, 19 Mar '5b. a F, I M).
#55-J-016. Coll. Akiyama. 1 i Oct •55.(,yFl.
#55-J-01S.Coll, Akiyama. 4 Nov ■55.1/ /•; /A/1.
#67-J-0145. Coll. Kanno. 5 Dec '66, ( / /•'!.
#67-J-0 144. Coll. Ushishigc, 8 Nov '66. (/ /■•).
#67-J-0089, Coll, lloliibcc & MlMi- sawa. 18 Apr '67.(7 I . 18 M. / F).
#67-J-029(l. Coll. Sasagawa & Lslin, 9 Nov '67, (15 I-, 2M.9U.JF. 3M).
#55-J-lll5. Coll. Nagayoshi, Jan '55, (/J A').
#67-J-0087, Coll. Yamaguti, 8 Apr '67.(8 [', 26 M, / F).
#67-J-nl77,Coll. Yamaguli, 8 Apr '67, (2 M».
#67-J-0 181. Coll. Yamaguti. Mar '67, (/An.
#67-J-0294, Coll. Suzuki. 15 Dec '67, (2 r, 19M. 3N, IF).
#56-J-060, Coll. Toshioka & Aki- yama. 25 lcb'56,(.''A/).
#67-J-0179, Coll. Yamaguti, Mar '67. (4 Ml.
#67-J-(l 182. Coll Yamaguli, Mar •67, (5 F, 7 Aft.
#67-J-0 184, Coll YaiiKigiili. Mar '67,(7 M).
*67-J-0292,Coll. Suniki, .1 Dec '67. (8 V. 70 M, h F. h Aft.
-56-J-063, Coll. I suchiyama, Sep '56, (/ F).
Jf68-J()309, Coll. Utsumi. 6 Apr '68,(4 1-, 6 M, 14 N, / F. I M. 2N).
196
BntCHAM Young University Science Bulletin
Haemaphysalh
jlava
I.epus hrachvunts
angxiHtidi'fis
(Hare)
Horse
Dog. liorsc. kow. hare, vegelalion
Horse, vegetation, Lcpus sp, (Hare)
Cattle, horse, dog
Japan
Dog
Cattle or horse Wild pig
Dog. horse, cattle, wild boar, serow
Kagoshiina
Iwate
Tokyo?
Saga-'
Miyagi?
Hiroshima
Shiniane
Kyoto.'
Mic
Hiroshima Chiba
Ogawa. Kokutni-'
Okunakayaina
I aniba
isG
Kiyosumiyama, Awa-gun
^68-J-l)339, Coll. Yaniamoto. 16 Jan-.l Mar '68, (9 I. 42 M. 18 N. 6L).
(No. 25.'t5), Coll. Watanabe. 16 VIII 1939.(1 I", labelled as //. waianahei n. sp.).
Neumann. 1897:333-336.(11 I",
7 M. I 3 N. L - type scries-), presented by Hrot. Suto at Tokyo, Yariiagiiehi at Saga, Miskina at Sendai. Neumann states nymphs were from hare & cow, larvae from eow.
Neumann. 1905:238, 141 1:1 12. (as flaia var. annata oi jlava armata) cited by N. & W., 1915.
Donit/, 1905:105-134. Donitz expressed doubt con<.crning Neumann's idcntitiLation tor the specimens m Herhn Museum.
VVarburtun, 191)8:510-512. Same specimens as N. & W.. 1915. Piesented by Donit/..
NuttailA Warburton. 1915:408- 410.(1 I . 1 M).N. 893. Also cited Neumann's record (1911).
Kishida. l922a:852-853. Coli. Aoki, 30 Oct '09.(1 M)(as//. /.).
Kishida. 1927:985.
Dog, cattle & horse
Cattle and horse
Cattle & dog Horse and cattle
Horse
Horse and dog
31 species hsled
Domestic dog. Sch'nantos thihclanus japonicus (Asiatic Hi.ick
|
Korea |
S. Zenra |
Kan-toh and Sy |
|
|
(-(^holla- |
toh |
||
|
namdo) |
|||
|
Japan |
|||
|
(Honshu |
_ |
||
|
Kyushu, |
& |
||
|
Shikoku |
|||
|
Japan |
Hiroshima. Mic |
- |
|
|
Kumamoto |
|||
|
Tokushima |
|||
|
" |
lokyo |
Miyake (is.) |
|
|
,, |
Aomon |
. |
|
|
hvatc |
|||
|
,, |
Iwate |
Okunakayama |
|
Aomon |
Sambomgi ta-gun |
Kainiki |
|
U\.itc |
1 aki/awa. gun |
\\atc- |
|
Miyagi |
Motoyoshi \(tsht-giin |
Moto- |
|
kunuiinoto |
- |
|
|
llokkaulo |
||
|
and all |
||
|
over |
||
|
Honshu |
||
|
Chi ha |
. |
|
|
Akila |
||
|
1 ukushima |
Kishjdj. 1936: 142. According to Kishida, recorded by Mr. Doi of Keijo in 1935. (Kagakukan Hoh. •35, p. 9).
Sugimolo 1936c:580.
Sugniiolo, 1935:23-24, I937a:308. Cited records of N.&W., 1915.
Nakamura & Yajima, 1937: 157. Other known hosts were also listed.
Nakatsun. 1942:294.
Yajinia. 1942:5117-5119.
Vajinia, 1942:5(10-5(12. (as wdtanabci) (2 1). Coll. Watanalie, Ih Oct '39.
Itagaki, Nod.i cV Yain.ij:uchi. 1944. 1959.
Shigetnon. ,\so & Yajinia. 1953: 290-293.
.^sa^llnla,cl ai . 1 955: 1 27-1 2«,
.Asanuina. 1956:91-92.
Biological Series, Vol. 15, No. 1
Ticks
197
Hacmaphysalis jlava
Hear). I.ipiis brachyurns (Marc). Sciurus In (Squirrel). Phasianus colchictis tohkaidi (Green Pheasant). Phasianus soemnwrnngii scintillans (Hondo Copper Pheasant)
Dog
Japan
Miyagi
Yaniagata
Nagano
Kyoto
Saitamu
Shi/uoka
Kcegun & loshioka. 1957:14. Ineluding Holotype o\' H. orien- talis Kishida, l*)34. (rom Tokyo Prcleclurc.
Haemaphysalis formosensis
Man. domestic dog, Nyctcrcutcs procyonoidcs (Raccoon Dog). Mustt'la sibirica itatsi (Weasel). Selcnarctos tfnbclanus japonicus (Asiatic Black Bear), icpus brachyurus angitstidens (Hare), l.cpus brachyurus lyoni (Hare), Apodcmus spcciosus sado- cnsis (1 leld mouse), Phasianus cokhicus rohustipcs (Green Pheasant), (iarrulus glandarius toku- gawac (Japanese Jay), t'rn- beriza spodocephala per- sonata (Japanese Bunting)
Cattle, l.cpus brachurus angustidcns (Hare). Meles m. anakuma (Badger). Nye- tereutcs procyonoidcs vivcrrinus (Raccoon Dog), Strcptopclia o. oricn- talis { I astern Turtle- dove), Phasianus cokhi- cus robust ipes (Green Pheasant) and vegetation
Sus scrofa riukiuanus (Wild pig)
Pcnlalagus furncssi (Ryukyu Rabbit)
Vegetation
Haemaphysalis calves
fujisana
Haemaphysalis Domestic pig
hvsrhcis
Sus scrofa riukiuanus rWild pig)
Japan (Honshu. Shikoku, Kyushu)
Japan
Niigata
Ryukyu Islands
Kagoshinia
Japan Shi/uoka
Kagoshima
Mainland and Sado (Is.)
Because o! ubiquitous distribu- tion, name of prclcctures and collecting data arc not given here.
Saito. 1959h: 193-209. Coll. Saito, "56-'59. Immature forms on small mammals and birds. (A. N.L).
Kakudayama. Saito et al., 1965:145-146. Coll.
Sasagamine Pasture, Saito. II Sep '62-17 May "65.
Ohsadorynkan Pas- (I-. M. N. I), turc
Ukenson. Amami Oshima
Yuwan. Amami Oshima
Ukenson. Amami (Jshima
Yuwan. Amami Oshima
I oot of Mt. Fuji (700 m.) near Tujinom,iya
Yuwan, Amami Oshima
Iriomote (Is.)
Iriomotc (Is.)
Ukenson, Amami Oshima
#67-J-U216,Coll. Iipton &. Mizu- sawa. 18 Mar "67. (/ F).
#bS-J-()32U. Coll. Mi/usawa. 5 Apr "68.(7 Af).
#68-J-0324. Coll. Mi/usawa. 5 Apr "68, (I N).
#68-J-0325. Coll, Mi/usawa. 5 Apr '68.(1 N).
#67-J-0217.Coll. Tipton & Mizu- sawa. 23 Mar '67.(7 AD.
#67-J-0266. Coll, Mi/usawa. 6 Oct "67,(7 m.
Kitaoka. 1970:73. (oil. H. Toyota. (A. N, L).
#65-J-0003. Coll. Hajime. Nov '65A5F.JM).
#55-R-021. Coll. Kcegan & Toshi- oka. 10 Nov '55. (iF. 5 A/).
#55-R-022, Coll. Kcegan & Toshi- oka.5Dcc'55.(/ F. I M. I IV).
#67-J-(J066. Coll. Tipton & Mi/u- sawa. I 3-19 Mar "67. (9 i". 3 M. 9F, WM. fiX).
#67-J-0075. Coll, lipton & Mi/u- sawa. I 3- 1 9 Mar "67, (.;/■'. / M.l /V).
198
Bhicham Younc Univehsity Science Bulletin
Hacmaphysalis hyslricis
Siis scroja riukiuamis (Wild pig)
Jjpan Kaj:oNhiinu
llkcnson, Aniatni Oshima
#67-J-(UI7«, Coll. Iipcon & Mizu- sawa. 13-14 Mar -ft?, (6 1-. 2 M. h F. :: M)
R>uk> ti Islands
Vainatosoii. Amaini Oshiina
#67-J-()07l.{ oil. Mi/usawa, 21 Mar '67.(4 1.4 M. 2(1 N. / F. 4 M. J .V).
Asiiikcn. Aniaini Osliinia
Yiiwaii. Aiiiami Osliinij
V67-J-024.S, Coll. Mi/.dsawa. 2(1 Sep 7,7.(6 \ .jl F. JOM)
a(,7-J-ll255.('oll. Mi/usawa. 1 Oct '67. (/ /. / 1/1.
*67-J-02.S7.(oll. Mizusawa. I Oct '67,(1 N. // h. 4M).
Ashikcii. Ainanii Oshinia
-67-J-027().<oll. Mi/usawa. 14 Oct '67.(-Mfl.
Dog
Yliwan. AmaTiii Oshinia
I'kcnson, Anianii OsIiinia
Yuwan. .'\ni.iiiii Oshirna
a6.S-J-l)ll(l4. (oil llaMnic. Dec '65.(.W-'1-
ffh7-J-(l21S,( oil llajimc. 1,1 Aug'67.(/ AD.
#67-J-(l214. Coll. Ilajinic. 1,1 Aug '67. (2 Fl.
»67-J-(l274, Coll Mi/usawa. Apr
#67-J-li24S. Coll. Mi/usa\w. Ill Sep '67. ( / Fl.
#67-J-ll2h.!,Coll KiinuraAi Mi/u- sawa. 4 Oil '6 7. ( / /I.
Pcnlala^ts furncs^i (Ryiikyu Rabbill
Vegelution
I'kenson, Am.inii Oshiina
#67-J-ll(16X. Coll, Tipton & Mi/u- sawa. 24 Mar '67.(1 N. / ,V).
#67-J-(l(l7(). Coll. Tipton & Mizu- sawa. 2.1 Mar '67.(1 I . 5 /V).
"Iiiwan. .^inann #67-J-0244. Coll, Mizusawa. 26
Oshiina Sep '67. (J f).
#67-J-0264, Coll. Kiniura & Mizu- sawa. 4 Oct '67. (2 An.
#67-J-027N. Coll. Kiniura & Mizu- sawa. K Oct '67, (/ /■■).
■ff67-J-027,S. Coll, Mizusawa. Apr '57. (/.A/I.
#67-J-0.1IIX.( oil Mizusawa. 6 Oct '67.(74 I.. / /• reared).
.V»,^ scrttfa riiikiuaniis (Wild pigl
Inoda, Ishigaki Keegan & loshioka. I4,S7: 1 5, Coll,
( Is. I Keegan & loshioka, 4 Nov ',S.S,
(I I , .5 M, 1 N,4 1 I,
Cow
Mountain near Keegan i loshioka. 1 457: 1 5, Coll,
Inoila, Ishigaki Keegan & loshioka. 8 Nov '.S.S,
(Is.) (7 I . 17 M, 26 N. IS L),
Yonakunills,) Siiginioto. 1437a 31. V 14371,604,
Coll. (Ilka. 6 Oct '36.1 I I ),
.Vi/,v scrofa nuktuanus (Wild pigl
Sus scrofa ssp. (Wild pig)
Shrubs
Otoini, Irioinote lis I
Mountains near Inoda, Ishigaki (Is.)
C liiiukj. Okinawa
kju.ishiina, 1463 1113 Coll Kawasliiina. IS Aug '62. |2 I . 1 M, 2N, I 1.1.
Iloogstraalel al.. 146.S:477, (2 1,4 M, 6 N. I L). (-RMI .14123): Coll, Keegan & Toshioku. 8Nov'.S.'i.(2 M. 1 I . 2 N. 1 1) (-IIII 53261
Moogslraal et, al,. 1465:477, (oil, llardcaslle. 31 Aug '46. 1 1 M). JfRMl. 27531.
Hioi.oGicAi. Sf.ries, Vol. 15, No. 1
Hacmapbysalis (Iround
hvstricis
Ryukyu KbtuK
Ticks
Tanyu-yaina. Okinawa
199
Hoopstraal ct al., 1965:477. Coll. Werner. 13 Jnl'51.(2 l).(MCZ).
Japan
Haemaphysalis laponica
Harniaphysalis kitaokai
Caphcornis crispus (Scrow)
Ci'nus nippon (Sika Deer)
OonK'stK t..ittlc
Dog
DonK'stK cattle
I.i'pus hrachvurns brachvurns (Harel
Horse
Cattle & horse
Dog. horse, cattle, wild pig, Capricornis crispus (Serow)
Capricornis crispiis (Scrow)
Ccr\-us nippini (Sika Deer)
Domestic cattle
Korea
Jjpjn
Nagano Kanagawa Tochigi Hokkaido
Nagano
Hanigyung- pukto
Mic
Shiga
Shunane
ShiiTiajuna
Tanzawa Mis.
Kiniigawa
Ktittan. lokachi
Ko/awa, Shiribeshi
Higashimokoto. Abashiri
llnggi & Kyonghung
Neumann. I 9U 1:2(> l-2()2. (as hispinosa. 2 I).
#53-J-005. Coll. Sakaguchi. 24 Nov'53, (/ A/).
#6K-J-l).V)4.Coli. Isucliiyj. 17 Mar "68, l2 m.
*56-J-(l34, Coll Toshioka & Aki- yama. 2,S I cb'.Sf.j/ A/).
#67J-()I^X. Col! jjuigichi. 13 May '67. ( 14 N,/") A, (> M).
#67-J-()322. Coll. Tosliioka & Sasa- gawa. IS Jul '67. (/ /■').
#55-J-()64, Coll loslnok.i & Aki- yania, 13 Aug '55. (/ M).
Asanunia. 1 95K: 274. Coll. Asanuiiia. '5S, ( 1 N).
Keegani Toshioka. I 957: 1 7. Coll Yajinia. May '4(1.(1 I . 1 M) (labelled as cuncinna).
Kecgan & Toshioka. 1457:17. Coll. Asanuma. Jur4i . ( I M). (labelled asjczoemis).
llagaki. Noda& Yamaguchi, 1444. 1959.(as/czmviA/.v in 1944, japonica in 1959).
Nikuppu, Hidaka Memambetsu. Abashiri Abira. Yututsu. Ihuri
Sapporo &. Kushiro
Hondo
Ouchiyjina
Sanbe. Oda-viii
OguraAc lakada. 1927:206. (as jczoensis).
Warburton. ! 908:5 I 2-5 I 3, (as japonmca). Numerous males. type specimens, found \\\\h flava ut the Hritish Museum (see discussion).
Neumann. 1911:112. {.\^H. Jlava
Jlava)
Nutlall& Warburton. 1915:402-403. (as//, japonica) Same specimens as Warburton's types. Coll, The Duke otliedtord.
#54-J-006. Coll, Asanuma. 17 Jan '54, (.?/•*. JM).
#67-J-Ol7X.Coll. Yamaguti.K Mar '67.(7 Af).
#67-J-OIX5, Coll. Yamaguti. 22 Mjr'67.(6 1 .8M. /M/-'. 9 A/).
#67-J-(n80. Coll, Yjmaguti. S Mar 'blA'iF. JAf).
#66-J-OOI9,Coli. Sugiyam.1 & Molu- bee. 6 Oct '66. (Aft h').
#670-0286, Coll. Yamaguli. 4 Nov '67.(32 I-. II M,5N,.Vf. 2 M).
#67-J-0289, Coll. Yamaguti. 4 Nov '67. (//■•).
200
Bhioiiam Young University Science Bulletin
Haemaphysatis kilaokai
Capriconiis crispus (Scrow)
Caltlf
Dcct
Jjpan
Vegetation
Cattle
Capncurnis cnspus (Serowj
Horsi
Kanaga\^a Nagasaki
Mie
Siiiniane
1 . I an/d\va Mts
Shiinanc
ruki>ka (su) (= I ukuoka)
I iikiiuka
?f6H-J-l)3(l5,Ci)i:, Isuihiya, 5 Mar '68.(1 I ).
Nakano-mura. Kita- Keegan & i'osliioka. 1957: 16. (as
[iiatsuura-gun
Sanhe
//. iiwrmis) Coll. Yajmia. 28 Jun '52.(1 I).
Keegan & Toshioka. 1957: 16. (as //. incrmis) Coll. Asanunia. I 7 Jan '54, (.idl . 27 Ml.
Iwasaki el al.. 1964. las//. iticrnm). Coll 1 suelile. .Sep '62- Aug '6.V
Suglmolo. 19.17a:315. (as//. ttwrmts) Probably cited froni Neumann { 19(16).
KiIaoka& Morii, 1967a: 145-I52. (I , M), Nov '62, (as ambi^w).
Neumann. 19(16:217, 1911:109. (1 I) (as incrmis and ambigita. may be the same specimen).
Nutlall & Warburlon. 19 I 5: 367-36H. (as itwrmis). Cited Neumann's rect)rd.
Nuttall& Warburlon. 1 9 1 5: .t67-.!6X. (as incrmis var. aponomvidcs) (1 I ). Berl. Mus. 17.1, also examined b> Hoogstraal. 1969.
Keegan & loshioka. 1957: 16. (as incrmis var aponomindc\). (lied Irom Nutlall & Warburlon. 1915.
Cow
Ccrxns nippon (Sika Deer)
Horse and Caprivonns crispus (Serowl
tlacmaphysalis Horse
longicornis
Shimane
Mie
Nagasaki
Hokkaido
Sanbe
Kanunokuni lliyjiua
lomioka
Kaininokuni. Hi\ .iin.i
lua/ato. Isushima
Korea
Misaki. Abashiri
I udii ri. Cbeju Do
lloogslraal. I 96 1 :3| 7-31 8 (as incrmt\ aponttmoidcs)', 1962:195- 1 99 (as jpunoinoidcs). Cited from Nutlall cV Warburlon. 1915.
lloogslraal, 1969:21 1-221 (as kitookai), Holotype, allotype, and paratypes.
Hoogstraal. 1969:21 1-221. Coll. Yamagutl, 22 Mar "67, (9 M, 19 11. Paratypes.
Hoogstraal. 1969:21 1-221. (2 II. Identitied by Neumann ds amht^ia. Hrit. Mus. #1367. #139. (1 I ). Identitied by N. & W\ as incrmis vdT. apononioidcs. Berl Mus. 173.
=-55-J-()()S. Coll. losliioka* Aki- yama. 17 Aug '55. iJ I').
#55-J-()32, Coll. Toshioka & Aki- yama, 17 Aug '55, ( 1 1.4 !■')
#55-J-(Hii. (nil Toshioka \ Aki- yama. 2 Aug '55. IV /•').
= 55-.l-ll49.( oil Tosliioka *: Aki- yama. 17 Aug '55. 1/ /•')
:;67-J-(ll 211, Coll. liptonii Mi/u- sawa.6 Mav '67,(52 I , 19 M, 9N,.M/. -/.V).
#67-J-()123.Coll. hploiiA Mi/u- sawa, b May '67.(9 I . 3 M, X N.
/ F. I ^n.
#67-J-l)189. Coll. loshioka & Sasa- gawa. 13 Jul '67, (15 I , 29 N, .V /■").
=67-K-(IIKil.( oil. Munsu.5 May '67, (.ff. 2N).
Biological Series, Vol. 15, No. 1 Horse
flacfnaphysalis longicumis
Cattle
Korcu Japan
Ticks
Hirosliima
Hokkaido
Nagasaki
201
Kj^oshinij
Shinianc
A?um;iy;iniij
Migasliiniokoto, Abashiri
Sliinlokli. iukaLhi
Oshinia-gun. Iki (K.)
Hatsuyama. Ikl (Iv)
Takcnotsuji. Iki (Is.)
Gonoura. Iki (Is.)
Iwazato, Tsushima
kliinan
Sakura Jinia
Sanbc
*ft7-K-()0()y. CoU. Munsu. 5 May •67, (.5 K h m.
#55-J-()09. Coll. Toshioka & Aki- yuma. 19 .Sup "55. (/ Af).
#55-J-OI I. Coll. loshioka & Aki- yama, 19 Sep '55. (ft /•■, .i M).
#.S5-J-(12.';.Coll. losliioka* Aki- yania, 1.1 Aug ■.s.s,( 1 3 I , 9 N. 2J F).
#67-J-0193, Coll. losliioka&Sasa- pawa. 15 Jul '67. (4 1.2 N..<F).
»67..|.|)II).S. ( ui: I iptun & Mi/u- sawa. 4 May '67. (6 1 , _^ I').
# 67-J-(ll06.Coll. ripton& Mw.u- sawa. 4 May '67.(22 1 , 5 M. 42 N, / /■'. ft A/. ft.V).
#67-J-010X, Coll. Tiplon ii Mi?u- sawa. 4 May '67,(6 !■, 5S N.4 F. UN).
#67-J-() ill. Coll. Tipton & Mizu- sawa,4May '67,(1 1,7 N, J F, / M. UN').
#67-J-01l2.Coll, Tipton & Mizu- sawa, 4 May '67, I 3 1 . 27 N, 2 F. 2 M. 21 IV).
#67-J-()l 1.1. Coll. Tipton* Mizu- sawa, 4 May '67,(4 1 ,4 M. U N, 23F.4UM. 7N).
#67-J-01 15, Coll. Tipton &. Mizu- sawa. 4 May '67, (2 !■, 4 M, 7 N, 2F.3M.8N).
^7-J-Ol 16, Coll. Tipton & Mizu- sawa, 4 May '67,(1 N,i/-', / M. I IV).
#67-J-014.1. Coll. Tipton & Mizu- sawa. 2 May '67, (2 I . / Ft.
#67-J-01 19. Coll. Tipton & Mizu- sawa. 6 May '67.(2 1 . 1 1 N, / F. 211 m.
#67-J-lll27,Coll. lipton & Mi/u- sawa. 1(1 May '67. (/ Fl.
#67-J-ll 1.1(1. Coll. Iipton & Mi/u- sawa. Id May '67. ([ \ . 2 F).
#67-J-0 1.13, Coll. Tipton & Mizu- sawa. 10 May '67.(3 V.Jh').
#67.J.(1 136. Coll. liplon & Mi/u- sawa. 1(1 May '67. (J Fl,
»67-J-(ll39. Coll. Mizusawa, 14 May '67.1/ Fl.
#67-J-()l52,Coll, Holuhcc & Sugi- yania. 15 May •67.(80 1 . 3 M, 28 N, / /■!.
a66.J-(l()23.C oil iloluhii & loshioka, 13 Jul '66.1/ Fl.
#6(.-J-(l(l25.Coll. Ilolulx-i & loshioka. 13 Jul '66. (/ F"l
-66-J-0020. Coll. llolubccS loshioka. 6 ()Lt '66, 12 M. 2 AT.
#68-J-ll327. Coll. Yaiiiaguti & Hk-tha. 4 Jun '68.(2 1.4 M.65 N).
202
Uai'ttuiphysaUs Cattle
longicornis
Japan
ShiiiianL'
Iwatc
lokyo
Korea
HHt(:nAM Doling Uninersity Science Bulletin
Sanlje
lU'ppu. ok I (Is.)
San be
Ki'nuHlii Pasture.
Takayaina-niura.
Agatsuma-gun
Sotoyania
Ua> asaka H.iLhijn Koshnna Aogoashinia Ih.i-Ri.Cheiii Do Ha-Gl, ( heju Do
#6«-J-0.t29, Coll. Yaniagutl & Kleeha, 4 Jun '(<S.(7 1,4 M. 4N).
#66-J-l)026,Coli. Yaniagiiti. 12 Sep '66. (X I . .1 M. J2 N. 1110 L,
J /•■. .' ^f).
^6(i-J-()02l,(oll. Vainaglitl. 12 Sep 'M.il /••).
=hS-J-ll.V1(). Coll ^'ani.iguli& HIeeha. 4 Jun 'hS. ( 15 I . 2 M.
1 Nl.
=h6-J-(lll2'),( oil, Yaniagutl. .1 Sep '6ft. (/ F).
=?fth-J-l)ll22. ( oil Yaniagutl, 3 Sep ■ftft. ( / /■•).
#67-J-OI4h, Coll Salto. 2h Apr •62.(7()N. / /-'I.
= h7-J-l)2l I. Coll Yalllagull. Ill Aug '(i7, (17 1).
ff67-J-(l212. Coll "laniagutl. 10 Aug '67. (>) I . I Nl
#67-J-y2l.?.Coll Yaniagutl. 10 Aug"67.l« I . 2 Nl
=f.7-J-(l214. (oil 'laniagutl. 1(1
Aug 'h7. (2 1 . 1 N. 1 Ll
5h7-J-(l2ll7. (oil taniaguti. ]^ Jul'h7.(2 I . / /-'I
Sh7-J-ll2(l>(. (oil lakallashl. 2ft Jul '67. ( Id I . / /■)
Sft7-K-l)()l)2.Coll Yi Muiisu. .S Mav ■ft7, |I4 N. / M. I .VI.
?)!b7-K-ll003,Coll. Driggers. 5 May •hl.iJh: J At. /.V).
Naedo-Ki. ( lie|N Do ^7-K 0004. Coll. Dnggers. .S May 'h7. It, M).
#ft7-K-()00.'i. Coll Driggers. 5 May ■•67,(2 1 . I N.-4F\.
Ilalliin. Cheju Do
Jf ft7.K-0()06, Coll. Driggers. 5 May ■ft7.i.11-, 2 M.4K I M).
#ft7-K-0007, Coll. Parsons. .S May ■ft7.(2 i: I M. /.V).
Bay-uni. Clieiu Do
«h7-K-00()S. Coll Parsons. .S May ■67. (X I . .S N. / /■. 2.VI
Song-C;a, Cheju Do •!;ft7-K-00l 0. ( oil Driggers, 5 May
•ft7,(.'; 1-. .1 M, 7 N, 7 h\ I M. :.V].
^67.K-I)0I l.( oil YiMunsu.5 May '67.(2 I . 4 N. / K J m.
Aewol, Cheju Do
-67-K-1I012. (oil. Parsons. .S May '67.(1 I-. 1 N.-//-. / .1/1
-67-K-(ini3, Coll. Yi Munsu, 5 Mav '67,(4 P. 4N, //■', I M. .<.V).
#67-K-OOI4,Coll. Driggers. .s Mav '67,(2 I-, 1 M, JF).
*67K-OOI5. Coll, Parsons, 5 May
'67,(1 1-, I M, 9 N, :f. J .W,
BioLOGicAi, Series, Vol. 15, No. 1
HacmapllvsaUs C 'at t Ic
Ticks
203
Ccrvits nippon (Sika Deer)
Dog
J J pa II
Donic'stiL- sheep
Vegetation
Cattle, horse, slieep goat, dog
Korea
Japan
Korea
Japan
Dog, eow
CatllL-
Horse
Cattle. hor>e. dog. badger, deer
I othigi
Hokkaido
lokyo
Hokkaido
Shizuoka
Miva/aki
Tottori
Hokkaido
At)mori
Iwate
Gumma
Nagano
Nabum-Ni. Cheju Do
Kumik. Cheju Do
Kumsong-Ni. Chejii Do
Sogwan. Cheju Dt) Kwason. C'heju Do
Kiiuigawa Misaki, Abasliin Kozawa. Shiribeshi
Haehlju (is.)
Ishidor ) arm. Cheju Do
Misaki. Abashin
Ch'onsong-Ni
Cheju Do and Koje Do
K>ago i. Haragi
Izu
Kvushu
Takanabe
Daisen
#67-K-lMll6.Coll ViMunsu.S May '67,(1 1-. I N. .'/•■, J M. I ,V).
#67-K-0()2(), Coll, Parsons, (. Mav '67,(4 I. 1 N,.;/-'l
=67-K-()021, Coll, Parsons, (, May '67,(1 \. I M)
ir67-K-()(i:6, Coll Parsons, f, May '67. Cl 1 . : N. 2 h. .',1/1,
#67-K-()()2X. Coll, Driggers. 6 May '67.(1 M. 2 N. 2 1,.,V/--, / A7, 2 .V),
rf56-J-()6I.Coll, losliiokaS Akl- yama. 25 1 eb '56, (/ \U
#67-J-(ll')II.Coll losliiokaii ,Sasa- gawa, \2, Jul '67. ( 14 I . 5 N.JF),
#67-J-(12()l.Coll. Toshioka& Sasa- gawa. IK Jul '67, (35 1-. 33 N, Ih F).
#67-J-0203, Coll, Toshioka & Sasa- gawa. l(i Jur67,(/F),
#66-J-(>24(). Coll, Yamaguti. 3 Sep '66, (/ F. -■',V|
#67-J-()208,Coll. LItsumi. 13 Jul '67,(2 1 , / /■•).
#67-K-0023, Coll, Driggers. 6 May '67,(1 N.5M).
#67-K-()()24. Coll, Driggers & Yi Munsu, 6 May '67, ( / K / A/1,
#67-J-() 191, Coll, Toshioka & Sasa- gawa, 13 Jul '67, (3 1 . 4 N. / F).
Hoogstraal et al,. 1 MhKl 2114- 1 2(16, Coll, Traub. 16 Jun '52. (1 11. RML #33(1'):,
Moogs'raal et al,. 196S: 12(14-1206. Coll, Parsons and/or Driggers and \i Munsu. Other data, see Hoogstraal el al,. l')6H
Hoogstraal et al.. 14(>S: 12(14-12116, Lcetotype temale, paraleetotype male & synlypes, (6 1 , 2 M) N, 1425 in Neumann eolleetion,
Hoogstraal et al,, 1968: 12(14- 1206 (3 1, 2 M) marked "TYPi;", ( I 11 e\ iou. Kyago. (lb) e\ dog. Haragi. in Merlin Miiseuiu
Hoogstraal el al,. 196K 12114-1206. II 1 I lint, Mus, #1 I. 12, 13. 14.
Hoogstraal et al,. I 96S: 1 204-1 2(16, Coll, Sugimoto. 14 Jul '33. (4 1 . 1 Ml RML #151X5,
Hoogs(raal et al.. 196S 1 204- 1 206. (9 I . 1 Ml N, 2409. Nuttall ( ollestion in Unt, Mus,. same speeimens m N, & VV,. 1915,
Hoogstraal el al,. 196X: 1204-I2()6. (X M N, 290X. Nuttall Colleetion in Brit, Mus,. same speeimens in N.& W,. 1915,
Hoogstraal el al,. 196X,
204
HaemaphysaUs longicornis
Cattle, horse, dog. badger, deer
Japan
Tokyo
Naru
Hiroshima
Shimane
Nagasaki
Kagoshima
Bhigham Young University Science Bulletin
Hoogstraalct al.. 1968.
Cattle, horse, dog
various provinces
Dog
Man
Korea
Japan
I.cpus hrathyunis brachvunis (Hare)
Cattle
Cattle and horse
Horse
Vegetation
Cattle, horse, sheep, dog, goat
Kyoto
Nagano
Kagoshima
Okayama
Hokkaido
Tokyo
|
Japan |
Hokkaido |
|
& |
Aoniori |
|
Korea |
hvate |
|
Miyagi |
|
|
Chiba |
|
|
Hyogo |
|
|
Okayama |
Kcijo ( = S(.'OuI(
Gmkakuji-cho. Kyoto-shi
Osunii
Sjnbc
A/unu\ jtnu
Higashiniokdto- niura, Aba'.hiri
Ishikari
Oshinia
Donitz. 1905:127-129. (UN//. ncumanni) Type specimens.
Neumann. I 9 II : 1 09, (as // m-umamu).
1 rapido, 19h-'i: IhO; Hoiigslraal & Irapidi). 1966: 1 192. (as//. ncumanni} (fi 1 , 2 Ml. Nn. 1425. Trapido seleeted 1 I lectotype from (he lot ( = syntypc ol Donitz's ncumanni). Hoogsiraal & Trapido redescnbed the leelotype and paraleetotype.
Kishida. 19.16:139-142. (as//. ncumanni)-
Nagahana & Matsuo, 1962:1 19-121. (as// hispintisa) ( 1 I I. 21 Jul '61.
Asanuina. 195«:279. (as//. bispinosa) Coll. Asanuma. 'SS. (I-, N).
Kcegan & loshioka. 1457: 11-12. (as //. hisptnosa) ("oil, .Asannnia. 9 .Sep ■4X. (IX I ).
Keegan & Toshioka. 1957:11-12. (as//, hispinosa) (.'oW. .Asanuma. 19 Jur55,(4 II,
Keegan& Toshioka. 1957 11-12. (as// htsptnctsa) CoW. loshioka. 14 Jul '55. (.1 I 1.4(16 MGL^l & (1 1 1.406 MCI =4.
Keegan & Toshioka, 1957:11-12. (as // hispinosa) Coll. Ttishioka & Akiyama. IN Sep '55. ( 15 l\ 6 N. 102 1.1.
Keegan& loshioka, 1957: 11-12. (as//, hispinosa} Col\. Yajima. l.^i Jun '54, l2 1 . 2 Ml
Keepan & loshioka. 1957:1 1-12. (as// htspinosa] Coll. loshioka & Akiyama. 19 Sep "55. 406 MGL #.1.1.(7 1 . 5 M,4 N. 14.1 LI.
Keegan & Toshioka. 1957:1 1-12, (as// /)/iyjiV;r;,vfll Coll. loshioka Si. Akivama. 1 .1 Aug '55. 406 MGL #11.(.16 1 .9 N.4 II.
Keegan & loshioka, 1957:11-12. (as// bi^ptnosa) Coll, Toshioka & Akiyama. 17 Aug '55. 406 MGL *7,(1.17 1 . 11 N. .1 LI.
Keegan & Toshioka. 1957: 1 1-12. (as //. bispinina) ( 1 II. 5 Jun '55, 4116 MGl -II),
Nakatsujl. 1942.294. (as//. htspinosa).
Itagaki, Noda & Yamaguehi, 1944. 1955. (as// hispinosa).
Biological Series, Vol. 15, No. 1
Ticks
205
Haetnaphy sails t aitk\ horse, dog.
longicornis sheep, goat
Japan Korea
Shiinane
Oila
Miya/aki
Kagoshima
Hamgyong-
pukto
Itagaki. Noda & YainagULhi, 1955. (as//, hispinosa).
1944.
Korea
Hamgyong-
naindo,
Kyonggi-do.
Kyongsang-
puklo,
Kyongsang-
nariido.
Cholla-
nanido
Cattle, ht»rse. human, wild rabhil
C attic & horsi
Japan
& Korea
Japan
lukuoka, Kumamoto Miyazaki Kagoshima
Honshu. K\ iisfui Hi Hokkaido
Sugimoio. 1936e:580. (as//. hispinosa).
Sugimoto. I937b:605. (as//. hispinosa) (F. M, N. L). Jul-Oct '33 and Jul-Nov '36.
Lepus hrachyums angiisttdens (Hare), Meles meles anakuma ( Badger). \'yctereu!es procvonoides viverrimts (Raeeoon Dog), cattle & vegetation
Cattle, horse. Ai. hare
Cattle, horse, goat
Korea
Japan
Cattle, horse
Cattle, horse, hare. Sox. &. vegetation
Horse
Niigata
all over the peninsula
Kakudayama. Sasagamme Pasture Si. Ohsadorynkan Pasture
|
Japan |
Hokkaido |
Hakodate, Sapporo. |
|
& |
Muroran. Asahikawa, |
|
|
Korea |
Aomori Iwate Kumamoto |
Abashiri 1 usan (=Pusan) |
|
Japan |
Hokkaido |
|
|
& |
Iwate |
|
|
Ryukyu |
lukushmia |
|
|
Islands |
Ibaraki Cluniina Hyogo Shnnane Okayama Hiroshima Tokushima Oita Nagasaki Miyazaki Kumamoto Okinawa |
|
|
Korea |
Chhungt^hhong- pukto. Kyongsang-do. Ph\ongan-do. Hamgyong- pukto. Kangwan-do. Cholla-nanido |
|
|
Japan |
Iwate. Ai>rnori |
- |
|
Kumamoto |
_ |
Kishida. 1936:139.142. {ds H. hispinosa) ALLOrding to Kishidu. Akusawa recorded hispinosa in 192K.
Saito.et ai.. 1965:145-146. (as H. hispinosa) Coll. Sailo, 8 Aug '61-23 Apr '65, (1, N, L).
Ogura & Takadu, 1927:2(l4-2()5. (as //. hispinosa)-
Nakamura & Yajima. 1937: 15 1. ( as //. hispinosa).
Yajinia. 1942:507-509. (as//. hispinosa).
Shlgemori. Aso & Yajima. 1953:290- 293. (as//, hispinosa).
Nutlall & Warburton. 1915:433. (as//, hispinosa] ( 1 I . 1 M) N. 129, presented to N. & W. by Uonitz as bis ncumanni.
Nuttall & Warburton. 1915:433. (as//, hispinosa) l\) N, 2907a. 15 Jul '12. lound witb Dcrmaccntor sp.
206
Rrigiiam Young University Science Bulletin
Hacmaphysalis longicornis
Horse
Japjn
Cuttle
Hacmaphysalis Sus scrofa leuiomyslax
mcgaspinosa (Wild pig)
Cen\(s nippon (Sikj Dc'cr)
Miya/aki
Tokyo
Mic
Shiga Miya/aki
K a ruga w a
Takanabf
Kitagawa-rmiru & Aya-machi, Higashi- rnorokata-gun
Miyagasc
NulIalUSc Warlnirlun. 1415:433. (as//, htspinosa) I A. I.) N. 19()«. 2 Aug ' 1 2. tnund with IxoJcs ricmus (- ' /. nipp(mcrisis). also examined hy Hoog\traal.
Nuttall& Warburton. 1915:433. (a-.//, hispmosa) (I ) N. 1909, 5 Jun ■ 12 Oct '1 2. aKo cNannned by Hoogstraal.
Nuttall& Wurburlon. 1915:433. (av//. hispmosa) (1 1 ) N. 29 11 a. 1 6 Oct '11, tbund with Hoophilus \p,. submitted by Dr. Miyajima.
Coll, Yamagiiti. N Mar 'hi. (ISM).
Coll. Yamaguli. Mar 7i7. (4 1 . 1 9 M ).
Coll 'laiiiaguti. Mar 'Ul . ( 1 } M).
Coll. Vajinia. 15 Mar '61.(2 i).
Coll. Kilaoka. 27 Nov '68. (31 I . 15 M.X4 N.8 L).
Coll. Kitaoka. 5 Jan-23 I eb '69. (Numerous adults)
Haemaphy salts wellingloni
Capricornis crispus (Serow)
Vegetation
Hacmaphysalis Pcnlalam s furncs-ii
pentalagi (Ryukyu Rablntl
Nanasawa. Alsugi- machi, Naka-gun
Mmeyama. Kuribara. Isehara-machi, Naka- giin
|
Ryukyu |
Kagoshmia |
Ydinaloson, Amami |
|
Islands |
Oshima Yuwan-dake, Amam Oshmia |
Turdiis carUn cordis (Japanese Grey Itirush)
Japan
Yuwan-dake. Amanii tjshmia
Shi/uoka Hokugo-rnura. Sunto-
gun
Ixodes aaililarsus C'atlle
Naganti
Iwate
Maila-niura
K.i\\ .il-imira
=6S-J-0307, Coll. Tsuehiya. 5 Mar •f)8, (I Ml.
#68-J-().Wl.Coll. Tsuehiya. 17 Mar'68. {24 1-.28M).
Saito, l%y:87. Coll. Sallo.
5 Oct '68. ( I Ml Hololype. 1 1 I )
Allotype. 1.1 I . .1 Ml.") Apr '69.
Sailo, 1969:87. Coll. Sallo. Ill Apr '69.(1 M).
#67-J-IJI)h.l, Coll. Mi/usawa, 17 Mar '67.1/ .Wl.
JF68-J-II-1 1 1 . Coll. Mizusawa. ."! Apr 6S.li: I , 1 M. i N. 1 L, / r. I M).
*68-J-0.112.Coll. Ml/u^awa, 4 Apr '68 I 2 1.6 M, ."i N, / /•■. -^ M. J. VI.
if68-J-0.1l .1. Coll. Mizusawa. 5 Apr •68,(1 M, .K)N. //■■. 2,M. .'i-V).
S6X-J-0.'tl5. Coll. Mizusawa. 6 Apr
'68.(5 1 ,.15 M.. IN, /:/■: I M.
Pospelova-Shtrom, 19.15: 21.1. Coll. Sehmldl, '27,(1 Ml.
Keegan & Tosliloka, 1957:4-28. Ulseussed the loeality of Pospelova-Shtrom's record.
Hoogstraal & YamagutI, 197(1. Coll, Mizusawa, Apr '68, ( 1 2 1 , 411 M, 2.1 N, .15 LI.
Asanuma& Kosaka, 1954:1114-1(17. Coll Toshioka, 29 Apr '54, (1 Ml
Asanunia& Kosaka, 1954:1114-1117. Collected 19 Apr '51, (1 Ml.
Coll. Yajima, 8 Aug '49, ( / /-'I.
I Biological Series, Vol. 15, No. 1
Ixodes acu litarsu i M a n J a [la n
Capricorn is crispus (Serow)
Serow. cattle & dog
Ixodes an^istus Clcthrionomys rufocanus bcdfordiae (Rcd-backctl mouse!
Vegetation
Clvthnonornys rut this (Red-backed mouse)
Apodemus speciosus aimi (I leld mouse)
Clethnonomys rufocanus bedfordiae ( Red-backed mouse) & Apodemus speciosus ainu (Meld mouse)
,, Clethnonomys mfceanus
bedfordiae (Red-backed mouse)
Clcthrionomys rufocanus hedjordiae (Red-backed mouse), Apodemus speciosus ainu (Field mow^c). Apo- demus argenteus hokkaidi (Field mouse)
Ixodes granulafus Rati us raitus (Rat)
Suncus murinus nukiuanus (Shrew)
Rattus norvcgieus (Norway Rat)
Rattus sp. (Rat)
Under log
|
Ticks |
||
|
Ovik.i |
- |
Ohara& laiuura, l'^5S. CollcLted Aug '58. |
|
Wjkayanu |
Ml. Gornaiian |
KeL'ganAi losh.oka. 1457:18. Colk-ttcd 4 Jul '49. 1 1 1 ). |
|
Sailanui |
Shikanoyu. |
Kccgan & Toshioka. 1457: 18. |
|
Cliichibii-shi |
Coll. Asanuma. ( 1 1). |
|
|
caiu. |
Kishida. 19.?(la:l-5. |
|
|
Njgano & |
||
|
'lo> aula |
207
Hukkaido
Ryukyu Islands
Onncto. Ncniuro
Rcbunto, Soya Sarobetsii. Soya Otoineppu
Toyotomi
Ishikan. Neiiiuro, Abashin, Soya
Hiroo and Temc
Asanuma. 1947b:972.
Karsdi. 1X8(1: 142. Coll. Hllgcndorl. Type. (1 1 ),
Neumann. 1911:16.
Coll. Ono. 9 Sep '56,(1 L).
Coll. Ono, 22 Otl '56. ( 1 Nl.
Coll. Ono, 1 Jul "6 1,(1 1 ),
Asanuma, 1957:234 and Asanuma & Sckikawa, 1952: 1 1 1 . Coll. Asanuma. Jul '5 1 as Ixodes sp. .^ (larva) and /.vot/cs sp. 61 (nymph). (L. Nl.
Asanuma. 195 7:234 and Asanuma and Sckikawa. 1952:111,
Ono. 1966:62-68. Coll. Ono. Jun '55 -Oct '62, (A, N, L).
Asanuma. 1955:1240-1242. IN). as Ixodes sp. 61.
Hiroo, Sounkyo. Asanuma. 1955: 1 240-1242. (Ll. as
Otoineppu. Toyotomi. Ixodes sp. 3 Ishikari, Onaobetsu
Okinawa
Naha-shi. Okmawa ishigaki (Is.)
Nishihara. Ishi- gaki (Is I
Ohaia, Irioniote (Is.)
Okinawa
Nago. Okinawa
Takabanare Shima. Okinawa
#55-R-028. Coll. Keegan & Toshi- oka. 2nOet'55,(A/-', /;V. .?/,).
#R-00II7. (/ /■•).
Kitaoka'sColleetion. 15 May '69. l.n . 1 Nl.
Keegan & Toshioka, 1957: 19. Coll. Keegan & Toshioka. 10 Nov '55. RML =96.(1 N. 23 LI.
Keegan H. Toshioka. 1957: 19. Coll. Keegan & Toshioka, 29 Oct '55, ( 2 1-4116 MGL #75, 4 N-406 MGL #167).
Keegan & Toshioka, 1957: 19. Coll. Keegan. 3 Nov '55. 406 MGL #90, (1 N).
Keegan & Toshioka, 1957: 19. Coll. Hardeastle. May '45. RML »27522 (3 1).
Keegan & Toshioka. 1957: 19. Coll. Posekany. 25 Apr '45, RML #2751 1 (2 1).
Keegan & Toshioka, 1957: 19. Coll. Posekany, 15 Jun '45, ( I I-, 2 M),
208
Bhigham Young Univehsitv Sc:ience Bulletin
Ixodes gramilalus Railiis rallus i Rat I
Ryukyu Islumls
Okinawa
Kcegan & Tinhioku. 1957; 19. Coll. Posekany, 29 Apr '45. RMI, #27512
nil.
Kcugan& Toshioka. 1 957: 19. Coll. Povekany. 5 May '45, (2 1 -RML ^27514, ,1 H-RML #275 13).
Kccgan& I oshioka. 1957: 19. Coll. Posekany, 16 May MS. RML #275 I S (2 1,2 M, I Nl.
>'jinamiwj. Okinawa
Nago, Okinawa
Kccgan &. 1 oshioka. 1957: 19. Coll. Hardcasllc, 10 May 45. RML s:275l()(4 1 ).
KcL-gan& Toshioka. 1957: 19. Coll. Hardiastlc. 2.1 May '45. RML #27519(3 V).
Kecgan & Toshioka. 1957:19. Coll. Hardcasllc. May '45. RML #27521 (I I ).
Rattus noryegicus (Norway Rat)
Japan Tokyo
Naha. Okinawa llachijo (Is.)
Miyaki (Is.)
llachijo Koshinia
lachljo (Is.)
Kawashiina. I9(,.l: 1(1.1. Coll. Kawashinia. 25 Jun '62. ( I I ).
.^sanunia& Sckikawa, 1952:107-1 16, 1953:99-1 12. Coll, Asanurna, Dcv "51 & Mav '52, (A. N. 1.1.
Asanunia& Sekikawa, 195 2:107-1 16, 1953:99-112, Coll. Asanuma, Mar '52. (N.LI.
Asanuma & .Sekikawa. 1952: 107-1 16. 1953:99-1 12. Coll. Asanuma. Sep '50 & Aug '51, (N, L).
Asanuma* Sckikawa, 1952:107-116. 1953:99-1 12. Coll. Asanuma. May '50 & May '52. (A).
Asanuma & Sckikawa. 1952: 107-1 16. 195 3:99-112. Coll. Asanuma, Mar '50. Aug'51.& May '5 2.
(A. N. L).
Miyaki (Is.)
Asanuma & Sckikawa. 1952:107-1 16. 1953:99-1 12. Coll. Asanuma. Mar "52. (N. L).
ApoJcniu^ \pi'ti(tsus 1 1 icid Mouse)
Kochi OkmoShima Asanuma & Sckikawa. 1 952: 107- 1 1 6.
1953:99-112. Coll. Asanuma. Aug '52. (Al.
Tokyo Mi>akc(ls.l Asanuma & Sckikawa. 1952: 107-1 16.
1953:99-112. Coll. Asanuma. Mar '52. (N. L).
Ixodes lividus
Dog& car
Siincus muriiuts (Shrew)
I'urdus cchcru>pus
felat'nopiis
(Seven Islands Ihrushl
Apodcmus ayronus ssp. (lield mouse)
Di'lichofl urhica dasypus (Japanese House Martini
Kanagawa
Tokyo
R>uk,\u Islands
Japan lokyo
Chihara
Kurihama
1/u-shichito Nago. Okinawa Miyakc (Is.)
Paedun-ni. Panstmg. Chindong-ni
J:ipan Shimanc
lliiioinisaki
Asanuma & Sckikawa. 1952:107-1 16. 1953:99-1 12. Coll. Asanuma. Jan '53. (N).
Asanuma* Sckikawa. 1952:107-116. 1953:99-1 12, (oil, Asanuma. Jan'53.(Ll.
Asanumai. Sckikawa. 1953:99-1 12. Coll. Asanuma. (2 A).
Kecgan it loshioka, 1957: 19, Collected 26 May '45.(1 II.
Asanuma & Kosaka, 1955: 195. Collected (II.
Arthur. 1957:6N3-6S6. Coll. 37th Med. Co.. Jun & Jul '54. (lOM. 30 1 ).
-66-J-(l027. Coll. Toshioka. Hctch- Icy & Holubec. 9-15 Jul '66. (.5 /•").
Biological Series, Vol. 15, No. 1
TlCK.S
209
Ixodes lividus
Dclkhoti iirhua Jasypus (Japjncsc Mouse Mjrtin)
Riparia riparia ijimac (Fastern Sand Martin)
Japan
Delichon urbica Jasypus (Japanese House Martin)
Ixodes Vegetation
monospinosus
Man
Ixodes nipponensis Cattle
Cenus nippon (Sika Deer)
Dog
Lcpus hrachyurm aripislidens i Hare \
Vegetation
Dog
I.epus hrachyums angitsiidens i Hare)
( altle. horse, dog, badger, hare, weasel, man. small rodents & birds.
Guriima
Hokkaido
Mic
Kochi
Niigatu
Minakanii
Hamamokoto. Abashiri-shi
|
Nagano |
Hakuba-iniira |
|
Kitaa^umi-giin |
|
|
(« |
Otari-mura, |
|
Kitaazumi-gun |
|
|
Mie |
Odaigahara, |
|
Owasc-shi |
|
|
Niigata |
hdc Mts.. |
|
Kitakanbara-gun |
|
|
Shiga |
Otsu |
Sanbc
#66-J-0n2S. Coll. Su/uki, 19 Jul '66. (.?/■"!.
#6S-J-()334, Coll. Su/uki & .Sasa- gawa, 5 Jul '68. (50 I-, 20 M. 4N.5/-'. 5M.F\ I.).
#67-J-l)192, Coll. Toshioka & Sasa- gawa. 14 Jure?. (2 I-. 4 M, 3 N, / F.JM).
Ono. 1967:217. Coll. Ono. 3(1 Mar '63, (20I-, 25 M. 1 N).
Ono, 1967:217.Coll.Ono, 4 Aug '66.(2 I).
llchrkawa& Sato. 1969:96. Coll. Uthikawa. 3(1 Dec '67, ( 1 N).
Uchikawa & Sato, 1969:96. Coll. Uchikawa. 3 1 Dec '67 & 7 Apr '68, (2 K, 13 N,5 Ll.
Kitaoka's collection, Coll. Yokoyaina, 27 Apr '69. ( 1 I).
Saito. 1967. (1 1 ). 2X Jun '67.
#55-J-()33.Coll. Akivama, 10 Dec '55, (/M).
#66-J-01S. Coll. Ilolubec & Sugi- yama. 6 Oct '66, (1(1 1 . 10 M. 4:h. J/A7».
#67-J-0164, Coll. Holubec& Sugi- yama. 16 May '67, (/ Fl.
#67-J-0326. Coll. Holubec & Sugi- yania. 16 May '67. (/ F).
t*7-J-0281.Coll. Yamaguti, 4 Nov 'bl.iJ F. I M).
#67-J-0282, Coll. Yaniaguli. 4 Nov '67. (//O.
#67-J-0287, Coll. Yamaguti. 4 Nov '67.(5 F. 10 M).
#6S-J-033 I . Coll, Yamaguti & Blecha. 4 Jun '68. (/ /■).
#67-J-0 186. Coll. Yamaguti, 22 Mar '67. (.?/■").
|
Hiroshima |
Kamed |
ani |
*55-J-029, Coll. Akivama. 12 Nov '55,(JF). |
|
Shiga |
1 maz u |
#55-J-031.Coll. Akiyama. 12 Dec •5i.ll F. /.W). |
|
|
Tokyo |
Omc |
#55-J-040. Coll. loshioka. 14 Jan '56. (/ F). |
|
|
Kagoshima |
Ogawa |
Kokiihu- |
#6S-J-0338. Coll, Yamamolo. 16 |
|
shi |
Jan-3 Mar '68. (2 1 . 9 M. 1 Nl. |
Sanbe
lligashiliar;inil
Kakudayaina. Minamikatnbara-gun
Honshu, Kyushu. Shikoku & Sado Is.
^68-J-0328. Coll. Yamaguti & Blecha. 4 Jun '68 (/ F).
#67-J-034l.( oil, llolubcL & Sugi- yama. 19 May '67. ill).
Kitaoka& Saito, 1967:74. Coll. Saito. 1 1 Jan '65. (1 I I. parent ot holotype.
Kilaoka& Saito. 1967:82. Immature stages on .small rodents & birds.
210
Rmu;ham ^'oung University Science Bulletin
Ixodes nipponensis Clethriononiys mfocanus hvdfordiac ) Rcd-buLkcd nicnisc). C. ntiilus mikado (Rcd-bucked mouse), Apodcmus spcciosus amu {McUi mouse). ,-1, ar^cn lens (licid mouse).
J J pan
Hokkaido
[shikan ^: )lulaka
Ono. 1966:62-68. Coll. Ono. Oct '55 - Aug '62. (N. L). as Ixodes sp. NA & I-A.
Ixodes ovatus
Dog
Rodenis &. shrews
Cut lie
|
Miyjgi |
- |
|
|
HiikkjicUi •t |
Muruyama Shi Shiranuka. Kushjro Ko/awa. Shinbeshi |
Sapporo |
laro-cho
Ono. 1962^:25. Coll. Olurj, (Al.
Ono, 1962b: 155. Coll. Ono. as Ixodes sp. LA & NA.
#55-J-026.Coll. loshioku& Akiyanu, 14 Aug '55. I-/ /•■).
*67-J-().1()3. Coll. Toshioka & Sasagawa. 18 Jul '67. Li h'. 2 M).
#67-J-()2.1 1 . Coll. Toshioka & Sasagawa. 17 Sep *67. (/ /•').
#67-J-(K)yh, Coll. I oshioka & Sasag.iw,i. M Mj\ '(,7. II 1.7/-". / .Wl
.Shirn.ini'
Gcnbi. iLhinoseki
•=■(17^-0099. Coll. 1 oshioka & Sasagawa. 15 May '67. (.?/■").
#67-J-0158. Coll. Holubct & Sugi- yama. 15 May '67. 1/ /•').
Horse
Dog
Hokkaido
Miyagi
Aoniori
Tochigi
Shi/unai. Hldaka
Towada-inura
I'Lithi/awa
1/uru
ff67-J-(ll67, Coll. HoIuIk-l & Sugi- yarna. 16 May '67. (/ h. I Sf\.
*)7-J-()173. Coll. HolulK-i & Sugi- yama. 16 May '67. (/ /■').
#67-J-()302. Coll. Toshioka & .Sasagawa. 16 Jul '67. (7F, 23 M).
#55-J-(MI7.Coll AsjTuiTiij, 19 Jul '55,(J/-', JAfl.
#55-J-027.Coll. Akiyama, \} Aug '55. l//-").
#56-J-039. Coll. Toshioka & Aki- yama. 24 l-cb"56. (/ /•").
lanL-yatiid
Asama
S67-J-ll(195. Coll. Toshioka & Sasa- gawa. 14 May '67, (6 1. 2 h'\.
#67-J-llll9X. Coll. losliioka & Sasagawa, 15 May '67. ( 16 I . / /■').
#67-J-n239. Coll, Yamagull. '66.
1/ .in.
Kanagawa
Kl\ okawa
K.ilsuura. I'usano
I oyounu'-nnira. lawara/u
llig.isliih.iraiiii
#67-J-ll241.( oil Y.iinaguli. I I Apr '67.1/ h. : \1)
#67-J-l)243.Coll. 'lain.iguti. 25 Apr '67. i: I : \h
#67-J-(IIWl.( oil llolulH-i & Sugi- yama. IM Apr '67, | 1 M. / /•. / M).
#56-J-043, Coll losliioka & Aki- yama. 28 Icb '56, 1 / /•'!
#56-J-()44,Coll. Kecgaii* loslii- oka. 15 Jan '56.1/ /-I.
a67-J-lllX7,ColI. HolulH-i ii Sugi- yallia. 19 May '67, (J/-'l.
I.i-piis bravhyunta angttaridi-iis i Hare)
Kiimamolo
Kagoshima
Min.iiiial.i. I iiino\ u
Ogawa, Koktihu-shi
^56-J-ll4l,Coll. loshioka. 9 Jan '56, ( / /•■).
*ft8-J-()34l). Coll. Yamamoto, 16 Jan- 3 Mai ■6X. 1/ /•'. / ,1/).
Biological Series, V'ol. 15, No. 1
Ixodes ovatus Vcticlalioii
Apoiletnus spcci(}sus aimt (licld inuLisc)
Clethriononiys mfociuius ht'dfordiav (RL-d-backfd nunisc)
J^ipan
Ticks
Hokkaido
Tsukisuppu. Ishikari
Teinc. Islnk.m
211
#55-J-()47. (oil. Toshioka & Aki- yuma, 5 Jun '55, (/ F),
(\)ll Orui. IN Oct "55. (/ A). Coll. Olio. l^J Apr '5(1. (/ ,V).
Caltli.', hor^c & human
Horse, cattle, dog, hare. & human
Cattle & luirse
Man. cattle, dog. cat, Mustela sibirica itatsi (weasel). Xvclcrculcs procyonoidcs viicrrinu s (Raccoon Dogj. Moycra wogitra \vogi4ra (Mole), I.i-pus hrachyunm lyoni (Hare), Raltus noncgi- cwA(Nor\va> Rul), f.i'pus hrachyunis anfiustidens (Hare), Mtcrotus monte- belli (Korean Meadow Vole), M. monicht'lli brcvicorpus ( Meadow Vole). Apodcmus spcci- osus sadoensis (i ield mouse). Crocidura dsi- nezumi (Shrew), & vegetation
Rodents & shrews
Clclhnonomys mfocanus bedfordiae (Red-backed mouse), C. mtilus mikado (Red-backed mouse), Apodcmus argcnteus (1 ield mouse). A. speciosus amu (Field mouse).
Clethrionomys rufovanu s bedfordiae (Red-backed mouse).
A kit a.
Shi/uoka,
Kyoto,
Saga,
Tokyo.
Kanagawu,
Hokkaido
Niigata,
Akitu.
Shimane,
Hyogo.
(lumma,
rukushima.
Hokkaido
Aomori, Iwate
Aomori.
Iwate,
rukushima,
Miyagi.
Nagano.
Hokkaido
Niigata
Kanagawa
Hokkaido
Hakodate, Sapporo, Ogura & lakada, 1927;201-2()3 Muroran, Atsunai (as /. jrequem) {V, M).
Kishida. I93()a; 1-5. as /. frcquens. ovatus. japoriensis. & carinatus. Record oi Jrcqucns is cited from Ogura & lakada (1927).
Nakaniura& Vajimu, iy37:145-MK. as japonerisis & frequens.
Niigata Pret. in Honshu
&
Sado (Is.)
Mt. Ketsu, Iseharacho
Supporo-sln
Oshima, Ishikari. Hidaka, Abashiri, Tokachi. Rumoi
leme
Yajinij. 1442:507-504. ds frequens.
Itagaki, Noda & Yamaguchi, IM44, 1959, ds frequens.
Sailo, 1959b: 199-203. 1964:60; Saitoet al., 1965:145-146. (as /, japonensis) Coll. Saito. Jul '55- 17 May "65, (A. N, L).
Immature forms on small rodents.
Ono. 1962b: 155. (as/, sp. 6) Coll. Ono. (L).
Ono. 1966:62-68. (as/, sp. 6 & 52) Coll. Ono. Jun '55 - Oct *62.
(N.L).
Asanuma, 1955: 1 240-1242. (as /. sp. 52) (N.).
Apodcmus spceiosus (I-ield mouse), A. argenleus ( lield mouse). Hothenomys smilhi (Vole). Mus museulus (House mouse), Mierotus montehelli (Korean Meadow Vole)
Aomori,
Yamanashi,
Chiba.
Kanugawa.
Toyama.
i'ukui,
Shizuoka.
Kyoto,
Gita.
Asanuma & Sekikawa. 1952:107-1 16.
1953:99. (as/, sp. 6 & 52)
(N.L).
212
Ixodes ovatus Dop & hare
Japan
Chiba, Akila. lukushima, Miyagi,
Nilgata
Bhicham Young Univehsity Science Bulletin
Asanuma. 1956:92-93. (as/. iap'iuviiMs) {A I,
Kcctian & Toshioka. I957;2(l. (as /- japoyjcnsis) Coll. Saito, 27 Jul ■55.(2 II.
C'altk
Okayama
Dog
Ratlits nurvc^lcus (Norway Rat)
/ cjuts hrui hytinis brat hvttruK ( Marc)
Man
Niigata
Nagano
Kochi
Tokyo
Gini'undaira
K.iniiko^hi
Kecgan & loshioka. 1957:2(1. (as /. japunftisn) Coll. Vajnna. 1 ^ Jul •55. ( 1 1 ).
Ktcgan & Toshioka, 1957:2(1. (as /. japom'iisis) Coll. Yajinia, 10 Jul '36. (111. (labflli-d d-. frcQiu'ils).
Kccgan & Toshioka. 1957:20. (as /, japonensis) Coll. Sailo, 31 Jul
'56.(1 Ml.
KcLgan & losliioka. 1957:20. (as /. japoni-nsis) Coll. .Saito, 26 Jul '55,(1 1 ).
kccgan & loshioka. 1957:20. (as /. /aponcrisis ) Coti. llironiatsu. 4 IX-c '52,(1 1 , ION),
Kccgan & Toshioka. 1957:20. (as /. lapont-nsis) Coll. Hironiatsu. 21 1 cb '56.(5 I-, 1 Ml.
Kccgan & Toshioka. 1957:26. (as /. japoiu'iisis) Coll. Kishida, (III. (Hoiotypc ol cannatus Kisliida. 19301.
Horse, hare, dog
Horse, dog
Horse, dog
Ixodes persulcattis Cattle
Hor\c
Akita, Saga
Saga
Tokyo ?
Saga
Hokkaido
Hokkaido
near Tokvo
Shikotsu lake Iburi
Shintoku, Tokachi
Kuttan, lokadii
Ko/awa. Shiribcshi
Shintoku. lokathi
Shi/unai. Hidaka
Nciniiann. 1S99:1 16-1 IX. Coll, lagULlli (Akital & Yaniaguehl (Sagal. (4 1 I. Type scries contained 2 I of r/c7>(MS (=? pcrsitlcatus or mppotwnsis).
Neumann, 1904:452-453. 191 1 U& Diinitz. 1905:132, (2 1 ol 4 1 above I.
Nuttall& Warburton. 191i:15S.
(2 II. As ricinus var. ovatus.
same specimens as Neumann I 19041
.Neumann. 1904:45X. 191 1:22. Nuttall& Warburton, 191 1 :20>l-209. (as/, japortf lists) Coll, Harmand, 1901,( 1 1 I, type specimen, (Pans Museiiml.
Morel. 1963:925-928. (2 I I. same speciniens as Neumann ( 19041. Morel synonymi/ed japonensis under ovattts
Schul/e, 1930:294-303. (as/ frequens) Coll, S. Schocde. 16 Jul '06,(1 Ml.
#67-J-0197.Coll. laniguchi, 17 May '(.l.ifi F. IJM. I A).
■Sr67-J-0199. Coll. laniguchi. I ,i May '67,(i/-'l.
#67-J-0202, Coll, Toshioka & Sasapawa. IS Jul '67, (/ /•'!.
*67-J-(l242.Coll, Yaniaguli.6 Jur67.(/ F).
=67-J-OI95, Coll, laniguchi. 10 May '67.1/ /■•).
#67-J-0200. Coll loshioka & Sasagawa. 16 Jul '67. iS /-"I.
Biological Series, Vol. 15, No. 1
Ixodes fXTsulcatus Horse
Vrsin arctos yt'sot'nsis (Bear)
Japan
Dog
Man
Ticks
Hokkaido
213
Tokushimj
Sliiraniika. Ku'ihiro
Shintoku, Tokachi
Kozawa, Shiribcshi
Mt. 1 surugi
#55-J-05 I . Co!!. Toshioka & Akiyamu. 14 Aug '55. ( J /•").
#670-0196. Coll. Taniguclu. 19 Jun "67. (i/-; 2 m.
ff67-J-02f)4. Coll. Toshioka & Sasagawa. IS Jul '67, (/ f-\ I M).
#67-J-0:44, Coll. Yaniaguli, 25 Apr "67, (/ h').
Keegan& loshioka. 1957:21-22. Coll. Tanaka, 16 Jur56.(2 \\
1 N).
Horse
Dog
I.epus hrachvurns hrachvums (Hard
Weasel
Dog, Lt'pus brachyums angiislidens (Hare), /.. braihyunts lyoni (Hare), eattlc. N yctcrcutes procyonoides (Raeeoon Do'^), Micro tus man [c belli brcvicorpus (Meadow Vole). Apodemus spccioaus sajocnsis (lield mouse). Rattus norrt'^f^s (Norway Rat), Croadura dainczumi (Shrew). Phasianus colchicus rubu stipes (Pheasant), i vegetation
Rodents &. shrews
Clcthrionomys ntfoeanus bedjordiae (Red-backed mouse), C nifihis mikado (Red-batked mouse Apodemus speciosus amu ( lield mouse). A. ar^enteus (I ield mouse).
Clethnonomys ntjoeamis bcdfordiae { Red-batked mouse). Apodemus specio- sus aimt (Field mouse), A. argenteus hokkaidi il ield mouse). Mus musculus yesonis ( House mouse).
Apodemus speciosus airw (I ield mouse). A. argentues (Field mouse), ,4. speciosus (Field mouse), holhenomys smilhi (Vole). Cicihnonomys rufoeamis (Red-backed mouse), Mus musculus yesonis (House mouse)
I wale
Niigata
KoLhi
Niigata
Kanagawa
Hokkaido
Hokkaido
Aomori
Akitj
Saitamu
Yamanashi
(lunniij
Hokkaido.
Niigata,
Akita,
Yamagatu.
Nagano.
Crumma.
Gilu.
Ginzandaira
Aki-shi
Niigata Pret. in Honshu & Sado (Is.)
Mt. Ketsu. Isehara- cho
Keegan & Toshioka. 1957:21-22. Coll. Yajima. 27 Jun\l«.(l !■, 1 M).
Keegan & loshioka. 1957:21-22. Coll.Saito. 25 Jur55.(2 1).
Keegan & Toshioka. 1957:21-22. Coll. Hiromatsu, \^ Nov '55-10 Mar "56. (I\ M).
Keegan & Toshioka. 1957:21-22. Coll, Hiromatsu. (1 I ).
Saito. 1969b. 199-20.?. Coll. Saito. Jul'55 - I eb '59. (A.N.L),
Ono. 1962b: 155. Coll. Ono, (L) (as Ixodes sp. 2).
Ono. 1966:62-6S. Coll, Ono. Jun '55 -Oet '62. (N. L).
Sapporo-shi
Oshima, Ishikari. Hidaka. Soraehi, Kaniikawa, Tokaehi, Kushiro. Nemuro, Abashiri.
Shintoku, Hiioo. Asanuma. 1955:1240-1242. (N. L).
Nohoro, Sounkyo, as Ixodes sp. 2 & 54.
Otoineppu, Toyotomi, Shiranuka. Peine. Nonaka.
Sugayu Tazawa Chiehibu
Asanuma & Sekikawa. 1952:111, 1953:99 & 105. Coll. Asanuma. Ma> "53 -Jun "53. (N. L), as Ixodes sp. 2 & 54.
Yamanashi & Yamanaka O/e
Kitaoka& Saito. 1967:82.
214
Ixodes pcrsuhalus
Bricham Young University Science Bulletin
Ixodes philipi
Htirsc & hare
Varunis aminals
Horse
Oiltlc. horse &. nun
Human. i.attle. horse, hare, deer. dog. fox. roe-deer & leopard (troni /uo '). marten, hedgehog, weasel. sheep, goal.
Horse ?
Horse, cattle, dog
(altle. horse
Cattle, horse, dog
hijjinus ieucomclas or Occanadroma caslro
Ixodes sigTiatus l.ants erassirostris
(HIack-tailed C^ull)
Phalacrocorax pela^- cus (Pelagic Shag)
Japan
Korea
Japan
Korea
Jap.in
Korea
Japan
Hokkaido
Akita. Saga
Aornon. Hokkaido.
Hokkaido.
Tokyo,
Nagano,
Kyoto.
Hyogo,
L-'hime,
Saga.
N. Korea
Miya/.dki
Aornon
Shikolsti lake. Iburi
Dai sen
Schul/e. !y,H»: 294-303. Coll. Schoede. 16 Jul '06. (2 I ).
Neumann. 1899:116. 1904:452; NuttallA Warhurton. 1911:156. (as/, nanus) Nuttall & Warburton cited Neumann's record.
Neumann. I'M 1 1 M 3. (as/. ncinus).
Nuttall & Warhurton. 1415:433. (as /. ricinus) lound with //. bispinosa {=11. longtcornis) . 1 Aug '12.
Ogura& Takada. 1927:201. (as /. rieinus) (I . M).
KIshida, 193()a:2.(as/ ricinus)
Kishida, 1916: 142. (as/. ruinusl
Sugimoto, i937h:603-6()4. (7 I ). ds ricinus var. mivazakicnsis.
|
Kumamolo. |
. |
Nakamura& Yajima. 1937:142-14 |
|
Miya/aki. |
(as/, ricinus). |
|
|
Okayama. |
||
|
Akita. |
||
|
(iumnia. |
||
|
Niigata. |
||
|
Fukushima, |
, |
|
|
Iwate. |
||
|
Shimanc. |
||
|
Hokkaido. |
||
|
Hamgyong- |
||
|
pukto |
||
|
Aomon, |
- |
Yajniia. 1 942;5()7-50S. (as/. |
|
hvalc |
ricinus). |
|
|
Hokkaido |
Kiishiro. Kamikawa. |
llagaki. Noda& Yaniaguchi. 1944 |
|
Abashin |
1959. (as/, ricinus). |
|
|
lualc |
i aki/awa |
|
|
Aomon |
Sanbongi |
|
|
Miyagi |
Naruko |
|
|
I iikushnna |
liandai-nnira |
|
|
Mic |
Watarai |
|
|
Miya/aki |
Koyu |
|
|
llamgyong- |
Unggi. Kyonghung |
Itagaki.NodacV Vainaguchi, 1944 |
|
pukto |
1959. (as/ nctnus). |
SjnganiinKi Iskinii
Ku•ra^^ .inil Kohls. 1970:725. (A).
(= llokkaidu)
K.ihuiiriu. II.Khiniihc- ff.'iS-J-DSO, Coll, Toshioka & Aki- shi yamu. IK Aug '.'i.S. (,SW /■', -K A/l.
40h MGI -4ll,(oll. Aki.v.Mii,i. 14 Otl '55. (4h I . 7: M. 1 1 Nl.
*66-J-p.!2.l. Coll. losliiok.i & .Sasagavva. 15 Aug 'fth. (41.2 M, (i N, 2 1.. :i M).
J?ft7-J-IKW2. C oil. loshiuka i Sasagawa, 12 May 'b7. (2 1 , 2 M. I7SN, 12(12 L. ioF. I m.
Neumann. 19(11:2X4, 19(14:451. 1911: 21. (» I), (lypc. Hanib. Mus. i. \s parai'irostriK in 1901, vuilypliJis var. sinnariis in 1 904. Ji cudypndi\ si^iani\ in 1911.
Biological Series, Vol. 15, No. 1
Ticks
213
Ixodes si^ialus Phalacrocorax pcla^-
cus (Pelagic Shag)
Japan
Yc/o
(= Hokkaido)
Nultall& Warburton. 191 l:261-2ft4. As sifjnarns. L*\aininL'ii Neumann's specimens as well as sif^ialus sent them by Hirula. and tound them identical.
/.arus crassirosrris (Black-tailed Gul!)
Crevices in rocks near nests ot Lams crassi rostns (Black-tailed Gull)
PhalacroLorax pclagi- ctts (Pelagic Shag)
Pfialacrocorax capillalus (Temminck's Cormorant)
Japanese Cormorant
Aomori
Hokkaido
Hokkaido. Kanagawa
Hokkaido
Zumpt. 1952: IH. Merely lisis the previous record.
Kabujima. Hachinohc- Asanuma. Okubo & lukiida, 1955; shi K5-K6. Cull, lukudu & Asanuma.
May '51 - Ma> "56. (A.N. L).
Kabujima. Hachinohc- Keegan & Toshioka. 1957:22. Coll. shi Toshioka & Akiyama. IS Aug '55.
(70 I, 53 M, 3 N).4()6 MGL ^H.
Coll. Akiyama, 14 Oct '55. (96 I .
72 M. II N). 406 MGL #40.
Asanuiiij Si I ukud.i. 1957 147-159.
Asanuma, 1965a: 1 16.
Kishida. I930a:2. Kishida did not reter to the scientific name of the host. May be citation from Neumann or Nuttall & Warburton.
Ixodes simplex
PipistrcUus ahramus (Bat)
Okayama
Coll. Yamaguti. 9 Mar '57,(1 F).
Ixodes tanuki
Ixodes turdus
Myotis mavrodaetylus (Little Brown Bat)
Bat
Myotis macradac(ylu\ (Little Brown Bat)
Minioptcnts sehrei- hersi (Long-winged Bat)
Nyctercuies proeyonoi- des viverrimts (Raccoon Dog)
Turdus celaenopus eclaenopus (Seven Islands Thrush)
Motacilla cmerea easpi- ca (Eastern Grey Wagtail)
Turdus celaenopus celaenopus (Seven Islands Thrush)
Iwate
Nagano Shiga
Kanagawa
Niigata
Tokyo
Yamaguchi
Tokyo
near Kameaka
Kamakura
Asahi-nmra. Ivva-
tune-gun
Kurokuwa-mura, Kitakambara-gun
Sasagami-mura. Kitakambara-gun
I/u-shichito
Tokuyama-^hi Hachijo (Is.)
Coil. Lndo. 26 Jul '67. (/ ;V).
Kishida, 19.^()a:.l.
Keegan & Toshioka. 1957:23-24. Coll. Asanuma, ( I I).
Arthur. 195^: 1S9. 30 Sep "52, (I I , 3 L), RML #30943.
Arthur. 1956: 1H9. 24 Dec '45. (I N). RML #22366.
Saito, 1964:59-66. Coll. Saito, 23 Nov '59. (I F).
Saito. 1964:59-66. Coll. Saito.
22 0ct '59, (2 1-. I M).(l M: Holotype, I liAllotypc).
Saito, 1964:59-66. Coll. Saito,
23 Nov '63.(1 V).
#55-J-024. Coll. Asanuma, 13 Aug '55. (4 N, IF).
Kitaoka's C'ollection, 26 Nov '66. (3 l).
Nakatsiiji. 1942:291-295. Coll. Tono. 25 Dec '40. (7 I ).
Keegan & loshioka. 1957:24. Coll. Asanuma. 12 Dec '55.(1 I. 6N). 406 MGL # i4K.
Aogashima
Keegan & Toshioka. 1957:24. Coll. Asanuma. Nov '54.(1 I, 1 N).
Turdus c. celaenopus (Seven Islands Ihrush), Turdus ehrysolaus ehry- \olans (Japanese Brown Thrush), k'rnheriza ele^ans elegans ( Yellow-throated Bunting). Emhertza eioides ciopsis (Japanese Meadow
|
„ |
Aogashima, Miyake |
|
(Is.). Oshima. Hachi |
|
|
jo (Is.). Hachijo |
|
|
Koshima. Niijima |
|
|
Chiha |
Niihama |
|
I ochigi |
Nikko |
AsaniiriKicS Kosakj, 1955:194.
&
Asanuma & Nakagawa, 1955:549. ,11 May 'SI, Jan '52. I i-b •5.-!, May 'S.!. Jan '54, 1 cb - Mar •54. Nov'54.(l-. N. L).
216
Ixodes turJus
IxoUcs uhac
Ixodes vcspcrtilionis
Hunting). Efnbcriza spodo- cvphala pcrsonata (Jupancsc Hunting)
Ccttia diphonv cantons (JapancsL- Hush Warbler). Ccttia diphouc ijimac {Jjpjncsc Hush Warbler). Bambusicola thoracica thoracica
(Chinese Hjmboo Phea- sant),
Coccothratistcs cocco- [firaustcs japonicus (Japanese Hawfineh). Hypsipctcs atnaurotts amaurotis
(Hrown-eared Hulbul). Pyrrhula pyrrhiila jiriscivcNlris (Japanese HuIIIinLh). Pfiylloscopus ijimac (Ijima's willow warbler) Apodcmus spcciosus (I ield mouse)
,\'yctalus sp. (Hat)
l.unda cirrhata ( Tufted puttin)
Rhinolophus jcrrum cquinum nippon (Morseslioe bat)
Japan
Niigjij
Nagano
Hokkaido
iothigi
Rkicham Young University Science Bulletin
Asanunia & Seikjkawa, 1 45 2: 1 I I . as Ixodes sp I . , Asaniirna & Kosaka. 1955:194: Asanunia & Nakagawj. 1955:549. .11 May '51. Jan "52. Ieb& May '5.1. Jan - Mar & Nov '54. (I . N. L).
|
Shi/uoka |
Hokugo-nuira. Sunto- |
|
gun |
|
|
Tokyo |
Aogashnna. Miyake |
|
(Is.). Oshinu. Mathh |
|
|
jo (Is.). Hachijo |
|
|
Koshinia. Nuinna |
|
|
( hiba |
Niihania |
|
1 OL-higi |
Nikkti |
|
Shi/uoka |
Hokugo-nuira. Sunio |
|
gun |
Kisliitla. IM.^{tj:2. js/j»rw\. Douhtliil rcutrd.
Asarluinu, l')(.l: IXI. IMhSj: | Ih.
-5(vJ-l)37, (nil loslimk.i i \ki- y.iMia. :4 I ch'5(,.(_lN. 5 L. / /-I
Rhinolophus cornutus cormttus (florscshoc bjt)
#5h-J-(i«.C.)ll losli)ok.i& Aki- yaiiia. 24 I oh '5*,. | / ,W).
( hilDsL-, Ibun #h.l-J-(l(ll4, Coll. Hdtlon. 9 Oct
■63. (/ ,V).
#63-J-()()l.'l.Coil ILitluri. 2.1 Ocl '6.1. (/ F).
Iw.m.ii, .Slntihi'shi #65-J-Olll .S. Cull. Hjlturi. 7 May
'6.'i,(//V).
#65-J-(l(llh, Cull Hanon, 7 May ■65. (J/ I.
#64-J-IMII7.(oli. Ilatlon. 12 Jun '64, (/ /'I.
Mhinnlophti\ Jvnutu cquinum (llorscslHic lull
Rhinolophus jcrntm- cqumuni nippon lllorsishiK- hall
kuto
Japan
Iwalc
Kangwan- do
KiH hj
Ryuscndo. Iwai/umi- ^67-J-()iW4. Coll. i'oshioka & cho Sasagawa. 1 ,1 May '67, {2 h').
rabhan tavi-, Wonjii 2,S Jun '6.1, ill).
Shjmi/u, .'Xki-shi
Kwgan & Toshioka, 1957:25. Coll. Hironialsu, 1(1 May '5.1. ( 1 N).
Wakiyaina tavc
KccfianA loshioka. 1 457: 25. Coll. llirunialMl. '51. (2 Nl.
"lo^hnio
KiL-gan Sl loshioka. 1157:25. Coll. loOiioka, :i Mar '56,(2 L), 4(16 MCI *2(I4.
Rhinolophus cornutus cornutus (Horseshoe bat)
I ukuslnina
Kecjian & loshioka. 1 457: 25. Coll. Akiyarna. 2(1 IH-i '55, I I 1 I 4116 Mc;i «111.
Yoshino
Kccgan & Toshioka, 1957:25. (oil loshioka. 21 Mar '56. (2 L). 4(ih MC;i #2(11.
Pipistrcllus ahramus (Hall
ihinif
Kuroiwa i.uvc. Kamiukcnu-gun
Kei-gan &. loshioka. 1957:25. Coll. Monkawa, 21 Jan '56. (I II.
liioLOGicAL Series, Vol. 15, No. 1
IxoiJcs icspcrlilionis
Rhipiccphalus san^iinvus
Rhinolophus fcmun- cquinum (Ho^sc^hut• hat)
l.t'pus h. hraihvums (Hare)
Rhinolophus fvrnim- cquinum (Horsosiioi.' bat)
Dog
Man
Japan
Ryukyu Islands
Japan
Ticks
lochigi
Nayano
Myiagi (sic) (= Miyagi)
217
Kiinagawa
Okinawa
(amp McrLV. Okinawa
Yokohaina-shi
Ogaishi-inura, Aso- gun
Kunianioto-shi
Osaka-shi
Kccgan*. Tiisliioka. 1457:25. ColL loshioka & Akiyanu, 25 Icb '56 ( I N. 4 L). 4(16 M(iL #191.
Kccgan & liKhiiika. 1457:25. Coll. HironialMi. 2 Mar '28, (IN,.? L). This rciord seems to be doubtful.
Arthur, 1956: 1X4; lloogstraal, 1956: 569; Kcegan& loshioka, 1957:25. Coll, Nieholson. I.? May '52, (I I), RML#.12112.
#66-R-()(l05. Coll, Doucette & .Sugiyania. 19 Oct '66, {4 F. / ^T).
#6K-K-II(IIK, ( oil Hubert, .10 Sep '68,(2 I ).
«64-J-llll79, (oil. I lint, 7 Get '64, (2S I , 75 Ml.
Sugimoto, 19.17b:6l()-6 12. Sep '.1.1, (.11).
Sugiino((i. I9.17h 61(1-612, l8Jul •.^5,(.l 1,5 Ml,
Keegan & loshioka, 1957:26-27, Coll, Holland, Sep '56,(1 1).
Numbers in p.irentheses represent specimens senl (o Mi (;ien \1 Kolils lor contirniation ol delermin.ilions
218
Bhicham Young University Science Bulletin
APPENDIX 3
List of Maps and Text Figures
|
Map |
|
|
1. |
Map of |
|
2. |
Known |
|
3. |
Known |
|
4. |
Known |
|
5. |
Known |
|
6. |
Known |
|
7. |
Known |
|
8. |
Known |
|
9. |
Known |
|
10. |
Know n |
|
11. |
Known |
|
12. |
Know n |
|
13. |
Known |
|
14. |
Known |
|
15. |
Known |
|
Ih. |
Known |
|
17. |
Known |
|
18. |
Known |
|
19. |
Known |
|
20. |
Known |
|
21. |
Known |
|
22. |
Known |
|
23. |
Known |
|
24. |
Known |
|
25. |
Known |
|
26. |
Known |
|
27. |
Known |
|
28. |
Known |
|
29. |
Known |
|
30. |
Known |
|
31. |
Known |
|
32. |
Known |
|
33. |
Known |
|
34. |
Known |
|
35. |
Known |
|
36. |
Known |
|
37. |
Known |
PagL- 3
Japan, Korea and the R\ iikyu Islands
distribution of Argas japonicus 8
distribution oi Argas vespenilionis 14
distribution of Ornithodoms capensis 15
distribution of Ainblyununa guoemydae 23
distribution of Amhiyomma nitidum 25
distribution of Amhlyomma testudinarium 30
distribution of Boophihis microphis 36
distribution of llaenwphysalis campainilala 48
distribution of llacmaphysalis concinna 54
distribution of Hacmaphysalis sp. (H. cornigera group) 59
distribution of Hacmaphysalis doeniizi 61
distribution of llacmaphysalis flava 62
distribution of HaemaphysaUs fnrmosensis 68
distribution of Hacmaphysalis fujisana 77
distribution of llacmaphysalis hystricis 82
distribution of Hacmaphysalis japonica 87
distribution of Hacmaphysalis kilaokai 93
distribution of llacmaphysalis longiconiis 100
distribution of Hacmaphysalis megaspinosa 105
distribution of Hacmaphysalis pentalagi 110
distribution of Hacmaphysalis wellingtoni 113
distribution of Ixodes acutitarsus 115
distribution of Ixodes angustus 1 19
distribution of Ixodes granulatus 1 19
distribution of Ixodes lividiis 124
distribution of Ixodes monospinosus 129
distribution of Ixodes nipponensis 135
distribution of Ixodes ovalus 141
distribution of Ixodes persulcatiis & I. neimis 147
distribution of Ixodes phitipi 1 48
distribution of Ixodes signatus 155
distribution of Ixodes simplex simplex 158
distribution of Ixodes taruiki 158
distribution of Ixodes turdus 161
distribution of Ixodes vespcrtilionis 170
distribution of Rhipicephahts sanguineus 175
Fig.
1.
2.
3.
4.
5.
6.
7.
8.
9. 10. 11. 12. 13. 14. 15. 16. 17. 18. 19. 20. 21. 22. 23. 24. 25. 26.
Key characteristics of ticks - I (genus Ixodes) 4
Key characteristics of ticks -2 5
Key characteristics of ticks -3 6
llustratcd key to the genera of ticks of Japan, Korea and the Ryukyu Islands 7
llustration of Argas japonicus. female & male 9
llustration of Argas japonicus. larva & nymph 10
llustration of Argas vespenilionis. female & male 12
llustration of Argas vespenilionis. larva 13
llustration of Ornithodoros capensis. female & male 16
llustration of Ornilhorodos capensis, larva & nymph 17
llustration of .Amhlyomma geoemydae . female 19
llustration of Amhlyomma geoemydae. male 20
llustration of .Amhlyomma geoemydae. nymph 21
llustration of Amhlyomma geoemydae. larva 22
llustration of .Amhlyomma nitidum. female 25
llustration of Amhlyomma tesludinarium. female 26
llustration of Amhlyomma tesludinarium. male 27
llustration of .Amhlyomma testudinarium. nymph 28
llustration of A mhlyomma tesludinarium. larva 29
llustration of Boophilus mieroplus. female 32
llustration of Boophilus microphts. male 33
llustration of Boophilus mieroplus. nymph 34
llustration of Boophilus mieroplus. larva 35
llustration of Dermacentor sp., female 37
llustration of Dermacentor sp., male 38
llustration of Dermacentor sp., female 39
|
HioLocicAL Skuien, |
|
|
27. |
Ilustration of |
|
28. |
llustration of |
|
29. |
Ilustration of |
|
30. |
llustration of |
|
31. |
Ilustration of |
|
32. |
llustration of |
|
33. |
llustration of |
|
34. |
llustration of |
|
35. |
llustration of |
|
36. |
llustration of |
|
37. |
llustration of |
|
38. |
llustration of |
|
39. |
llustration of |
|
40. |
llustration of |
|
41. |
llustration of |
|
42. |
llustration of |
|
43. |
llustration of |
|
44. |
llustration of |
|
45, |
llustration of |
|
46. |
llustration of |
|
47. |
llustration of |
|
48. |
llustration of |
|
49. |
llustration of |
|
50. |
llustration of |
|
51. |
llustration of |
|
52. |
llustration of |
|
53. |
llustration of |
|
54. |
Ilustration of |
|
55. |
llustration of |
|
56. |
llustration of |
|
57. |
llustration of |
|
58. |
llustration of |
|
59. |
llustration of |
|
60. |
llustration of |
|
61. |
llustration of |
|
62. |
llustration of |
|
63. |
llustration of |
|
64. |
llustration of |
|
65. |
llustration of |
|
66. |
llustration of |
|
67. |
llustration of |
|
68. |
llustration of |
|
69. |
llustration of |
|
70. |
llustration of |
|
71. |
llustration of |
|
72. |
llustration of |
|
73. |
llustration of |
|
74. |
llustration of |
|
75. |
llustration of |
|
76. |
llustration of |
|
77. |
llustration of |
|
78. |
llustration of |
|
79. |
llustration of |
|
80. |
llustration of |
|
81. |
llustration of |
|
82. |
llustration of |
|
83. |
llustration of |
|
84. |
llustration of |
|
85. |
llustration of |
|
86. |
llustration of |
|
87. |
llustration of |
|
88. |
llustration of |
|
89. |
llustration of |
|
90. |
llustration of |
|
91. |
llustration of |
|
92. |
llustration of |
|
93. |
llustration of |
|
94. |
Illustration of |
|
95. |
llustration of |
|
96. |
llustration of |
|
97. |
llustration of |
Vol. 1.5, N(
Ticks
219
Dennacentor sp.. male 40
Dermacentor sp., male 41
Haemaphysalis campamilata. female 44
Hacmaphysalis campamilata. male 45
Haemaphysalis campamilata. nymph 46
Haemaphysalis campamilata. larva 47
Haemaphysalis concinna. female 50
Haemaphysalis concinna. male 51
Haemaphysalis concinna. nymph 52
Haemaphysalis concinna. larva 53
Haemaphysalis sp. {H. cornigera group), female 55
Haemaphysalis sp. (H. cornigera group), male 56
Haemaphysalis sp. (//. cornigera group), nymph 57
Haemaphysalis sp. (H. cornigera group), larva 58
Haemaphysalis doenitzi. female 60
Haemaphysalis flava. female 63
Haemaphysalis flava. male 64
Haemaphysalis flava, nymph 65
Haemaphysalis flava. larva 66
Haemaphysalis formosensis, female 69
Haemaphysalis formosensis. male 70
Haemaphysalis formosensis. nymph 71
Haemaphysalis formosensis. larva 72
Haemaphysalis fujisana. male 73
Haemaphysalis fujisana. female 74
Haemaphysalis fujisana. nymph 75
Haemaphysalis fujisana. larva 76
Haemaphysalis hystricis. female 78
Haemaphysalis hystricis. male 79
Haemaphysalis hystricis, nymph 80
Haemaphysalis hystricis. larva 81
Haemaphysalis japonica. female 84
Haemaphysalis japonica. male 85
Haemaphysalis japonica. nymph 86
Haemaphysalis kitaokai. female 88
Hacmaphysalis kitaokai. male 89
Haemaphysalis kitaokai. nymph 90
Haemaphysalis kitaokai, larva 91
Hacmaphysalis longicornis. female 95
Hacmaphysalis longicornis, male 96
Haemaphysalis longicornis, nymph 97
Haemaphysalis longicornis, larva 98
Haemaphysalis rnegaspinosa, female 101
Haemaphysalis megaspinosa, male 102
Haemaphysalis megaspinosa. nymph 103
Haemaphysalis megaspinosa. larva 104
Haemaphysalis pentalagi. female 106
Haemaphysalis pentalagi. male 107
Haemaphysalis pentalagi. nymph 108
Haemaphysalis pentalagi. larva 109
Haemaphysalis wellingtoni. male Ill
Ixodes acutitarsiis, female 114
Ixodes angustus. female 1 16
Ixodes angustus. nymph 117
Ixodes angustus. larva 118
Ixodes granulatus. female 1 20
Ixodes granulatus. male 121
Ixodes granulatus. nymph 122
Ixodes granulatus. larva 123
Ixodes lividus. female 1 25
Ixodes lividus. male 1 26
Ixodes lividus, nymph 127
Ixodes lividus. larva 128
Ixodes monospinosus, female 1 30
Ixodes nipponensis. female 131
Ixodes nipponensis. male 132
Ixodes nipponensis. nymph 133
Ixodes nipponensis. larva 1 34
Ixodes ovatiis. female 1 37
Ixodes ovatus. male 1 38
Ixodes ovatus. nymph 139
220 Brigham Voung University Science Bulletin
98. Illustration of Ixodes ovatus. larva 140
99. Illustration of Ixodes pcrsulcalus. female 143
100. Illustration of Ixodes persulcatus, male 144
101. Illustration of Ixodes persulcatus, nymph 145
102. Illustration of Ixodes persulcatus, larva 146
103. Illustration of Ixodes philipi. female 149
104. Illustration of Ixodes si^natus, female 151
105. Illustration of Ixodes signatus. male 152
106. Illustration of Ixodes signatus. nymph 153
107. Illustration of Ixodes signatus, larva 154
108. Illustration of Ixodes simplex simplex, female 156
109. Illustration of Ixodes simplex simplex, nymph 157
1 10. Illustration of Ixodes lanuki, female 159
111. Illustration of Ixodes tanuki, male 160
1 12. Illustration of Ixodes turdus, female 162
113. Illustration of Ixodes turdus. nymph 163
1 14. Illustration of Ixodes turdus, larva 164
115. Illustration of Ixodes vespertilionis, female 166
116. Illustration of Ixodes vespertilionis, male 167
117. Illustration of Ixodes vespertilionis. nymph 168
118. Illustration of Ixodes vespertilionis. larva 169
119. Illustration of Rhipicephalus sanguineus, female 171
120. Illustration of Rhipicephalus sanguineus, male 172
121. Illustration of Rhipicephalus sanguineus, nymph 173
122. Illustration of Rhipicephalus sanguineus, larva 174
Biological Sekies, Vol. 15, No. 1
Ticks APPENDIX 4
221
Figure
Species
Information on Specimens Illustrated
Remarks
5 Argas japonicus
Argas japonicus
10
Argas vespertilionis
Argas vespertilionis Ornithodoros capensis
Ornithodoros capensis
1 1 .4 mblyomma geoemydae
1 2 A mblyomma geoemydae
1 3 A m hlyom ma geoemydae
14 A mblyomma geoemydae
15 Amhlyomma nitidum
16 A mblyomma testudinanum
17 Amblyomma tcstudinarium
18 Amblyomma testudinarium
19 Amblyomma testudinarium
20 Boophilus microplus
2 1 Boophilus microplus
22 Boophilus microplus
Adult from Coll. #66-J-0037 - From nest of Hirundo daurica japonica (Japanese Striated Swallow), Niimi, Okayama Pref., Japan, 21 September 1966, Hatoyama & Mizusawa. Holotype female & allotype male, a-d, female; e-g, male, a, whole body (dorsal & ventral view); b, e, capitulum (ventral view); c, f, genital area; d, g, tarsi MV (lateral view). Adapted from Yamaguti et al., (1968).
Larva & nymph from Coll. #66-J-0037 - Same as above, a, b, larva; c, d, nymph, a, whole body (dorsal & ventral view); b, Haller's organ; c, capitulum; d, tarsi I-IV (lateral view).
Female from Coll. #67-J-0215 - From Vespertilio superans (frosted bat), Shojo, Fukushima Pref., Japan, 11 August 1967, Yamaguti. a-g, female: h. genital area of male, a, whole body (dorsal & ventral view); b, periphery (lateral view); c, posterior quadrant; d, tarsi I-IV (lateral view); e, spiracular plate; f, capitulum (ventral view); g, anal valve.
Larva - From Pipistrellus abramus (Bat), Fukuoka-shi, Japan, 25 June 1967, Wada's collection.
Adult from Coll. #66-1-0054 - Under rocks near nest of Larus crassirostris (Black- Tailed Gull), Kyojima, Shimane Pref., Japan, 9 July 1966, Toshioka & Betchley. a-d, female: e-g, male, a, whole body (dorsal & ventral view); b. e, capitulum (ventral view); c, f, genital area; d, g. tarsi 1-IV (lateral view).
Larva & nymph from Coll. #66-J-0054 - From Calonectris leucomelas (Streaked Shearwater), Ohakajima, Shimane Pref., Japan, 13 September 1966, Yamaguti's collection (larva); #66-1-0054 (nymph), a, b, larva: c, d, nymph, a, whole body (dorsal & ventral view); b, Haller's organ; c, capitulum (ventral view); d, tarsi I-IV (lateral view).
Female from 406 MGL CoU. #57 - From Cyclemys flavomarginata flavomarginata (Turtle), Futanaka, Ishigaki (Is.), Ryukyu Islands, 6 November 1955, Keegan & Toshioka. Reproduced from Keegan & Toshioka (1957), pi. 4.
Male from 406 MGL CoU. #57 - Same as above. Reproduced from Keegan & Toshioka (1957), pi. 5.
Nymph from Coll.#68-R-0015 - From Geoemyda spengleri japonica (Turtle), Yona, Okinawa, Ryukyu Islands, .April 1968, Mizusawa.
Larva from Coll.#68-R-O015 - Same as above.
Female from RML Coll. #35991 - From Sea Snake. Naha, Okinawa, Ryukyu Islands.
Female from Coll. #55-R-017 - From Wild pig, mountains near Inoda, Ishigaki (Is.), Ryukyu Islands, 4 November 1955, Keegan & Toshioka. Reproduced from Keegan & Toshioka (1957), pi. 6.
Male from 406 MGL CoU. #82 - Same as above. 2 November 1955. Reproduced from Keegan & Toshioka (1957). pL 7.
Nymph from Coll. »68-J-0318 - From Sus scrofa leucomystax (Wild pig), Yuwan, Amami Oshima, Kagoshima Pref., Japan, 11 April 1968, Mizusawa.
Larva from CoU.#68-J-0317 - From vegetation, Yuwan, Amami Oshimu, Kagoshima Pref., Japan. 10 April 1968. Mizusawa.
Female from 406 MGL Coll. #73 - From cow, Hateruma (Is.), Ryukyu Islands. Reproduced from Keegan & Toshioka (1957), pi. 8.
Male - Same as above. Reproduced from Keegan & Toshioka ( 1957). pi. 9.
Nymph from CoU. #67-J-0261 - From cow, Yuwan, Amami Oshima. Kagoshima Pref., Japan. 3 October 1967, Kimura & Mizusawa.
Bkic.ham Vounc University Sc ience Bulletin
23 lioophiliis microplus
24 Dermaccntor sp.
25 Dermaccnlor sp.
26 Dermaccnlor sp.
27 Dermaccntor sp.
28 Dermaccntor sp. 65
29 Haemaphysalis campanulata
30 Haemaphysalis campanulata
31 Haemaphysalis campanulata
32 Haemaphysalis campanulata
33 Haemaphysalis coneinna
34 Haemaphysalis coneinna
35 Haemaphysalis coneinna
36 Haemaphysalis coneinna
37 Haemaphysalis sp. (W. cornigera group)
38 Haemaphysalis sp. (//. cornigera group)
39 Haemaphysalis sp. (//. cornigera group)
40 Haemaphysalis sp. (//. cornigera group)
41 Haemaphysalis doenitzi
42 Haemaphysalis flava
4 3 Haemaphysalis /la va
44 Haemaphysalis flava
45 Haemaphysalis flava
46 Haemaphysalis formosensis
47 Haemaphysalis formosensis
48 Haemaphysalis formosensis
49 Haemaphysalis formosensis
50 Haemaphysalis fujisana
5 1 Haemaphysalis fujisana
Larva from Coll. #67-R-0011 - iTom cow, Hirai, Ishigaki (Is.), Ryukyu Islands, 8 February 1967, Tipton.
Female - From horse, Unggi. North Korea, June 1942, Asanuma's collection. Reproduced from Keegan & Toshioka (1957), pi. 10.
Male - Same as above. Reproduced from Keegan & Toshioka ( 1957), pi. 11.
Female - From horse, Okunakayama, Iwate Pref., Japan, 16 October 1938, Yajima's collection. Reproduced from Keegan & Toshioka (1957), pi. 12.
Male - Same as above. Reproduced from Keegan & Toshioka ( 1957), pi. 13.
Male from 406 MGL Coll. #101 - From Wild pig, Shizuhara, Kyoto Pref., Japan, 12 December 1955, Akiyama. Reproduced from Keegan & Toshioka ( 1957), pi. 14.
Female from Coll. #52-J-004 - From dog, Miura Peninsula, Kanagawa Pref., Japan, Asanuma. Reproduced from Keegan & Toshioka ( 1957), pi. 17.
Male from Coll.#41-J-002 - From Man, Takarazuka, Hyogo Pref., Japan, 15 January 1941, Hiromatsu. Reproduced from Keegan & Toshioka (1957), pi. 18.
Nymph from Coll. #67-J-0 188 - From dog, Chiba Pref, Japan, 12 June 1967.
Larva from Coll.#67-J-01SS-F| - Reared from material m Coll. #67-J-0 188.
[■emale - From cow, .'\tsukeshi, Hokkaido, Japan, July 1966, Kitaoka's collection.
Male - Nozukesaki. Nemuro-shi, Hokkaido, Japan, May 1964, Kitaoka's collection.
Nymph - Reared specimen loaned by Kitaoka.
Larva - Same as above.
Female - From cow, Hachijo (Is.), Tokyct, Japan, 29 May 1949, Asanuma's collec- tion. Reproduced from Keegan & Toshioka (1957), pi. 19.
Male - Same as above. Reproduced from Keegan & Toshioka ( 195 7), pi. 20.
Nymph - Reared specimen loaned by Kitaoka.
Larva from CoU. #67-J-0?06-F| - From cow, Hachijo Koshima, Tokyo. Japan. 13 July 1967, Yamaguti.
Female from Coll. #56-J-057 - From pheasant, Akune, Kagoshima Pref., Japan, 10 January 1956, Toshioka. Reproduced from Keegan & Toshioka ( 1957), pi. 28.
Female from Coll. #55-J-018 - I'rom dog, Yamakita-cho. Kanagawa Pref., Japan, 4 November 1955. Akiyama. Reproduced from Keegan & Toshioka ( 1957), pi. 21.
Male from Coll. ff55-J-018 - Same as above. Reproduced from Keegan & Toshioka (1957). pi. 22.
Nymph from Coll.#67-J-0294-F| - Reared from material in Coll.#67-J-0294.
Larva from Coll.#67-J-0294-F'| - Same as above.
Female from Coll. #67-J-0216 - From Sus scrofa riukiuanus (Wild pig), LIkenson, Amaini Oshima, Kagoshima Pref., Japan, 18 March 1967, Tipton & Mizusawa.
Male from Coll. #67-J-021 7 - On vegetation. Ukenson. Amami Oshima, Kagoshima Pref., Japan, 23 March 1967. Tipton & Mizusawa.
Nymph from Coll.#67-J-O2l6-1', - Reared from material in Coll.#67-J-()216.
Larva from Coll. #67-J-0216-l'| - Reared tfom material in Coll. #67-J-0216.
Female from cow, foot of Mt. I uji, near I ujinonin.i. Slii/uoka. Illustrations from Kitoaka (1970).
Male from cow. foot of Mt. I u|i. near 1 ujinonuya, Shi/uoka. Illusiralions from Kitoaka (1970).
Bioi.ocicAi, Skkies, Vol, 15, No.
52 Hacinaphysalis fujisana
5 3 ilacinaphysalis fujisana
54 Hacmaphysalis hystricis
55 Hacmaphysalis hystricis
5 b Hacmaphysalis hystricis
57 Hacmaphysalis hystricis
5 8 Hacmaphysalis japonica
5 9 Hacmaphysalis japonica
60 Hacmaphysalis japonica
6 1 Hacmaphysalis kitaokai
62 Hacmaphysalis kitaokai
63 Hacmaphysalis kitaokai
64 Hacmaphysalis kitaokai
65 Hacmaphysalis longicornis
Ticks
223
66
Hacmaphysalis longicornis
Nymph Iroin cow, foot of Mt. luji, near lujlnoiniya, Sliizuoka. Illustrations from Kitoaka(1970).
Larva from cow, foot of Mt. Fuji, near Fnijinomiya, Shi/uoka. Illustrations from Kitoaka(1970).
Female from 406 MGL Coll. #92 - From Wild pig, mountain near Inoda, Ishigaki (Is.), Ryukyu Islands, 8 November 1955, Keegan & Toshioka. Reproduced from Keegan & Toshioka (1957). pi. 23.
Male from 406 MGL Coll. #92 - Same as above. Reproduced from Keegan & Toshioka (1957), pi. 24.
Nymph frorti Coll. #67-1-007 I - From Sus scrofa riukiuanus (Wild pig), Yamatoson, Amami Oshima, Kagoshima Pref., 21 March 1967, Mizusawa.
Larva from Coll. #67-1-007 1-F, - Reared from material in Coll. #67-1-0071.
Female from Coll. #67-1-0198 - From cow, Kuttari, Tokachi, Hokkaido, lapan, 13 May 1966, Taniguchi.
Male - From Capricornis crispus (Serow), Shimajima, Nagano Pref., Japan, 29 November 1953. Sakaguchi. Reproduced from Keegan & Toshioka (1957), pi. 27.
67 Hacmaphysalis longicornis
68 Hacmaphysalis longicornis
69 Hacmaphysalis mcgaspinosa
70 Hacmaphysalis mcgaspinosa 1 1 Hacmaphysalis mcgaspinosa 11 Hacmaphysalis mcgaspinosa
73 HacmaphysaUs pcnlalagi
74 HacmaphysaUs pentalagi
75 Haemaphysalis pentalagi
76 Haemaphysalis pentalagi 11 HacmaphysaUs wcllingtoni 1 8 Ixodes acu titarsu s
Nymph - Same as the lot of CoU. #67-1-0198.
Female from Coll. #54-1-006 - From Cervus nippon nippon (Sika Deer), Ouchiyama, Mie Pref.. lapan, 17 January 1954, Asanuma. Reproduced from Keegan & Toshioka (1957). pi. 25.
Male from Coll. #54-1-006 - Same as above. Reproduced from Keegan & Toshioka (1957), pL 26.
Nymph from Coll.#67-1-0286-F, - Reared from material in Coll. #67-1-0286.
Larva from Coll.#67-1-0286-F, - Same as above.
Female from Coll. #55-1-049 - From horse, pasture near Hakodate, Hokkaido, Japan, 17 August 1955, Toshioka & Akiyama. Reproduced from Keegan & Toshioka (1957), pi. 15.
Male from CoU. #55-1-009 - From cow, Azumayama pasture, Hiroshima Pref., Japan, 19 September 1955, Toshioka & Akiyama. Reproduced from Keegan & Toshioka (1957), pi. 16.
Nymph from Coll. #67-1-0191 - On vegetation. Misaki, Abashiri, Hokkaido, Japan, 13 July 1967, Toshioka & Sasagawa.
Larva from Coll.#66-J-0022-Fi - Reared from material in Coll.#66-J-0022.
Female from Coll,#68-J-0301-F, - Reared from material in Coll.#68-J-0301.
Male from Coll. #68-1-030 l-F, - Reared from material in Coll. #68-1-0301.
Nymph from Coll.#68-J-0301-l-, - Same as above.
Larva - Reared specimen loaned by Kitaoka.
Female from Coll. #68-J-0315 - From Pcntalagiis furnessi (Ryukyu Rabbit), Yuwandake, Amami Oshima, Kagoshima Pref.. lapan, 6 April 1968, Mizusawa.
Male from Coll. #68-1-03 15 - Same as above.
Nymph from Coll. #68-1-031 1-F, - Reared from material in CoU. #68-1-03 1 1.
Larva from Coll. #68-1-031 1-F, - Same as above.
Male - Adopted from Asanuma & Kosaka (1954). pp. 105-106. figs. I & 2.
Female from RML Coll. #27491 - From Man, Mt. Gomadan, Wakayama Pref., lapan, 4 luly 1949. Reproduced from Keegan & Toshioka (1957), pi. 29.
224
79 Ixodes angiistus
80 Ixodes angiistus
8 1 Ixodes angustus
8 2 Ixodes gran u la tu s
83 Ixodes gramdatus
84 Ixodes granulatus
85 Ixodes granulatus
86 Ixodes lividus
87 Ixodes lividus
88 Ixodes lividus
|
89 |
Ixodes lividus |
|
90 |
Ixodes monospinosi |
|
91 |
Ixodes nipponensis |
|
92 |
Ixodes nipponensis |
|
93 |
Ixodes nipponensis |
|
94 |
Ixodes nipponensis |
|
95 |
Ixodes ovatus |
|
96 |
Ixodes ovatus |
|
97 |
Ixodes ovatus |
|
98 |
Ixodes ovatus |
99
Ixodes persulcatus
|
100 |
Ixodes persulcatus |
|
101 |
Ixodes persulcatus |
|
102 |
Ixodes persulcatus |
|
103 |
Ixodes philipi |
|
104 |
Ixodes signatus |
HiuGiiAM VouNG University Science Bulletin
Female - Crawling on a man, Sarobetsu, Soya. Hokkaido, Japan, 1 July 1961, One's collection.
Nymph - From Clethrionomys rufocamis hedfordiae (Red-Backed Mouse), Robunto, Soya, Hokkaido, Japan, 22 October 1956, One's collection.
Larva - From Clethrionomys rufocanus bedfordiae (Red-Backed Mouse), Onncto, Nemuro, Hokkaido, Japan, 9 September 1956, Ono's collection.
Female from 406 MGL Coll. #75 - From Rattus rattus (Rat), Nishihara, Ishigaki (Is.), Ryukyu Islands, 29 October 1955, Keegan &. Toshioka. Reproduced from Keegan & Toshioka (1957), pi. 30.
Male Lee.
From Rattus coxinga (Rat). Grass Mt.. lormosa. 20 May 1958. Yung I'oh
Nymph - Adopted from Asanuma & Sekikawa (1952). fig. 4 & ( 1953), figs. 9 & 10.
Larva - Collected on Grass Mt.. Formosa. 20 May 1958. Yung loh Lee.
Female from Coll. #67-J-0192 - From Riparia riparia ijimae (Eastern Sand Martin), Hamamokoto, Abashiri, Hokkaido, Japan, 14 July 1967, Toshioka & Sasagawa.
Male from Con.#67-J-0192 - Same as above.
Nymph from Coll. #68-J-0334 - From the nest of Delichon urhica dasypus (Japanese House Martin). Minakami, Gunma Pref.. Japan, 5 July 1968, Suzuki & Sasagawa.
Larva from Coll.#68-J-0334 - Same as above.
Female - On vegetation. Odaigahara. Mie Pref.. Japan. 27 April 1969, Yokoyama (Kitaoka's collection).
Female from Coll. #66-J-0018 - From cow, Sanbe pasture, Shimane Pref., Japan, 6 October 1966, Holubec & Sugiyama.
Male from Coll. #55-J-033 - From cow. Otsu, Shiga Pref.. Japan. 10 December 1955, Akiyama. Reproduced from Keegan & Toshioka ( 1957), pi. 34.
Nymph from Coll. #68-1-0338 - From Lepus hrachyurus angustidens (Hare). Ogawa, Kokubu-shi, Kagoshima Pref., Japan, 16 January - 3 March 1968.
Larva from Coll.#67-J-034l-F, - Reared from material in Coll.#67-J-()341.
Female from Coll. #55-1-007 - From dog, Miyagi Pref., Japan, 19 July 1955, Asanuma. Reproduced from Keegan & Toshioka ( 1957). pi. 31.
Male from Coll. #55-J-0O7 - Same as above. Reproduced from Keegan & Toshioka (1957). pi. 32.
Nymph - I-rom Clethrionomys rufocamis hedfordiae (Red-Backed Mouse), Teine, Ishikari, Hokkaido, Japan, 19 April 1956. Ono's collection.
Larva - From Apodemus speciosus ainu (Field Mouse), Teine, Ishikan, Hokkaido, Japan, 18 October 1955, Ono's collection.
Female from Coll. #67-J-0 197 - From cow. Shintoku, Tokachi, Hokkaido, Japan, 17 May 1967, Taniguchi.
Male from Coll. #67-J-0 197 - Same as above.
Nymph from Coll. #67-J-0 197 - Same as above.
Larva from Coll.#67-J-02()0-F, - Reared from material in Coll.#67-J-0200.
Female from Puffinus leucomelas (streaked sheerwater) or Oceanodroma castro (madeiran storm petrel). Sanganjinia. hvatc Prefecture.
Female from Coll. #55-J-050 - Collected in crevices in rocks near nest of Larus crassirostris (Black- Failed Gull), Kabujima, Hachinohe-shi, Aomori Pref., Japan, 18
Biological Series, Vol. 15. No. 1
Ticks
225
1 05 Ixodes signatus
1(16 Ixodes signatus
1 117 Ixodes signatus
1 08 Ixodes simplex simplex
1 09 Ixodes simplex simplex
110 Ixodes tanuki 1 1 I Ixodes tanuki 1 1 2 Ixodes turdus
1 1 3 Ixodes turdus
1 14 Ixodes turdus
1 1 5 Ixodes vespertilionis
1 I 6 Ixodes vespertilionis
1 1 7 Ixodes vespertilionis
1 1 8 Ixodes vespertilionis
1 19 Rhipicephalus sanguineus
120 Rhipieephalus sanguineus
1 2 1 Rhipicephalus sanguineus 1 22 Rhipicephalus sanguineus
August 1955. Toshioka & Akivama. Reproduced from Keegan & Toshioka (1957), pi. 35.
Male from 406 MGL Coll. #40 - Same as above, 14 October 1955. Akivama. Reproduced from Keegan & Toshioka (1957). pi. 36.
Nymph from Coll. #67-1-0092 - Same as above. 12 May 1967, Toshioka & Sasagawa.
Larva from Coll. #66-1-0323 - Same as above, 15 August 1966, Toshioka & Sasagawa.
Female from RML Coll. #30943 - From Myotis macrodactylus (Little Brown Bat), near Kameaka, Honshu, Japan, 30 September 1952. Reproduced from Keegan & Toshioka (1957), pi. 37.
Nymph - From Myotis macrodactylus (Little Brown Bat), Iwatc Pref.. Japan, 26 July 1967, Yaniaguti (Endo's collection).
Female - Adopted from Saito (1964), p. 62, tigs. 1-5.
Male - Adopted from Saito (1964). p. 64. figs. 6-9.
Female from Coll. #55-J-024 - From Turdus celaenopus celaenopus (Seven Islands Thrush), Hachijo (Is.), Tokyo, Japan, 12 December 1955, Asanuma. Reproduced from Keegan & Toshioka (1957), pi. 38.
Nymph from Coll. #55-1-024 - Same as above.
Larva - Adopted from Asanuma & Sekikawa ( 1952). p. 112, fig. 5.
Female from Coll. #56-1-037 - From Rhinolophus ferrum-equinum nippon (Horse- shoe Bat), Izuru, Tochigi Pref., Japan, 24 February 1956, Toshioka & Akiyama. Reproduced from Keegan & Toshioka (1957), pi. 39.
Male from 406 MGL Coll. #191 - Same as above. Moulted from a nymph of the lot above. Reproduced from Keegan &. Toshioka (1957). pi. 40.
Nymph from Coll. #66-1-0015 - From Rhinolophus cornutus cornutus (Horseshoe Bat), Iwanai, Shiribeshi, Hokkaido, Japan, 7 May 1965, Hattori.
Larva from Col). #66-J-0016 - Same as above.
Female from Coll. #66-R-0005 - From dog, Okinawa. Ryukyu Islands. 19 October 1966. Doucette & Sugiyama.
Male from Coll. #64-1-0279 - From dog. Yokohama-shi. Kanagawa Pref.. Japan, 7 October 1964, Flint.
Nymph - Reared specimen sent from Texas, USA.
Larva - Same as above.
226
BUIGHAM ^OUNG UNIVERSITY SciENCE BULLETIN
APPENDIX 5
Corrected Scientific Names of Hosts Recorded in the Literature
Some of the scientific names of hosts cited in the Htera- ture require revision in accord with current knowledge of vertebrate taxonomy. Therefore, corrected names were used in Appendix 2. Revisions were based on the following pub- lications; Yoshinori Imai/umi (1960). Colored illustrations of the
mammalsof Japan. Hoikusha Publ. Co. Ltd., Osaka. 196 p. Kenji Nakamura & Shun-ichi Ueno (1963). Japanese reptiles
and amphibians in color, Hoikusha Publ. Co. Ltd.. Osaka,
214 p.
Keisuke Kobayashi (1965). Birds of Japan in natural colors, Hoikusha Publ. Co. Ltd., Osaka, 231 p. Rev. and Knl. Ed. The Ornithological Society of Japan (1958). A handlist of the Japanese birds, 4th Rev. Ed.
The English common names were taken trom the follow- ing publication: Ernest P. Walker ( 1964). Mammals of the world. Vols. 1 & 2.
Name cited
Corrected Name
Apodemus geisha
Apodemtis sylvaticus ainu
Apodemus sylvaticus
speciosus
Cervus nippon nippon
Crocidura russula
Cvclemys Jlavnmar^inala
Hori'iiiS canlans cantans
Horeires canlans ijimae
l.xos amauroiis amauroiis
l.epus linudus an^usfidens
I.cpus timidus brachyurua Lepus tinudus lyoni Meles anakuma Microhyla Jissipcs Nemorrhacdus cnspus
Phasianus versicolor
Phasianus versicolor tohkatdt
Phylloscopiis occipi- talis ijtmae Sciurus vulgaris lis
Sus leucomystax or
Susl. leucomystax
Syromaticus socmmcrringn
scintillans
i'rsus thihctamis /aponicus
PuJJinus leucomelas
Apodemus argenteus Apodemus speciosus ainu Apodemus speciosus
Cervus nippon Crocidura dsinezumi
C 'yclemys fla vo mar^ina la
Jlavomarginafa
Cctlta diphonc canlans
Cciiia diphonc ijimac
Hvpsipclcs amauroiis
amauroiis
I.cpus brachyurus angus-
tidens
I.cpus brachyurus
l.epus brachyurus lyoni
Meles melcs anakuma
Microhyla ornata
Capricorms crispus
cnspwi
Phasianus colchicus
rabustipcs
Phasianus colchicus
tohkaidi
Phyltoscopus ijimac
Sciurus hs
Sus scrofa leucomystax
Pha stanu s vo cm m crringii
\cinitllans
Sclenarctos thibclanus
japomcus
Caloncctris
Icucomclas
("ommon Name lield mouse I ield mouse licid mouse
Sika deer Shrew
Turtle
Japanese Bush
Warbler
Japanese Bush
Warbler
Brown-cared
Biilbul
Hare
Hare Hare
Badger I rog Japanese Serow
Pheasant
(jreen Pheasant
WilknvWarbler
Squirrel
Wild Pig
Hiindo Copper
Pheasant
Asialie Black
Bear
Streaked
Shearwater
Literature ^___
Asanumu & Sekikawa ( 1952). (1953) Asanuma & Sekikawa ( 1952) Asanuma & Sekikawa ( 1952)
Hoogstraa! (1969)
Sailo (1959b). (1964). Saito et al.
(1965)
Keegan & Toshioka (1957)
Asanuma & Kusaka ( 1955)
Avinunia & Kos.ika ( 1955)
Asanunu & Kosaka i 195 5). Asanuma &
Nakagawa (1955)
Saitol 1959b). Saito et al. (1965),
kilaoka & Saitu ( 1967). Asanuma,
Sakurai et al. (1955)
Asanuma ( 1956)
Saito ( 1959b). Saito et al. ( 1965)
Saito et al. (1965)
Sugimoto (1937a)
Neumann (1906). (1911). Nutta!l&
\V,irlHirlon ( 1915)
Saitu ( 1959b). Saito ct al ( 1965)
Asanuma ( 1956). Asanuma, Sakurai et al.
(1955)
Asanuma and Ktisaka (l'>55)
■Asanuma ( 1956). .Asanuma, Sakurai ct al
(1955)
Asanuma, Sakurai et al. ( 1955).'
Kishida (1922a)
Asanuma ( 1956). Asanuma. Sakurai et al.
(1955)
Keegan & Toshioka ( 1957), Asanuma ( 19561.
Saito (1959b)
Kenans and Kohls i 1970)
Page in Appendix 2
210,212.214
207.208.210. 211,214. 197.209,216
|
189. 195, 206. 2 in. 212. 214 |
199, |
200, |
203, |
|
188 216 |
|||
|
216 |
|||
|
216 |
|||
|
196. 20.S, 214. |
210. |
211, |
212 |
|
197 197,212, 197,205, 189 199. 200. |
214. 206, |
207 |
|
|
195,214 |
|||
|
197 |
|||
|
216 |
|||
|
197 |
|||
|
1X8, 195, 197 |
206, |
||
|
192, 197 |
|||
|
187,215, |
MUS. COMP. ZCOL. LIDRARY
FEB ISiqyp Brigham Young University h ^Ar o
Science Bulletin
A LIST OF ARTHROPODS OF MEDICAL
IMPORTANCE WHICH OCCUR IN UTAH
WITH A REVIEW OF ARTHROPOD-BORNE
DISEASES ENDEMIC IN THE STATE
by
Vernon J. Tipton
and Robert C. Saunders
BIOLOGICAL SERIES — VOLUME XV, NUMBER 2 AUGUST 1971
BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN BIOLOGICAL SERIES
Editor: Stanley L. Welsh, Department of Botany,
Brigham Young University, Provo, Utah
Members of the Editorial Board:
Vernon J. Tipton, Zoology Ferron L. Anderson, Zoology Joseph R. Murdock, Botany Wilmer W. Tanner, Zoology
Ex officio Members:
A. Lester Allen, Dean, College of Biological and Agricultural Sciences Ernest L. Olson, Chairman, University Publications
The Brigham Young University Science Bulletin, Biological Series, publishes acceptable papers, particularly large manuscripts, on all phases of biology.
Separate numbers and back volumes can be purchased from Publication Sales, Brigham Young University, Provo, Utah. All remittances should be made payable to Brigham Young University.
Orders and materials for library exchange should be directed to the Division of Gifts and Exchange, Brigliam Young University Library, Provo, Utah 84601 .
Brigham Young University Science Bulletin
A LIST OF ARTHROPODS OF MEDICAL
IMPORTANCE WHICH OCCUR IN UTAH
WITH A REVIEW OF ARTHROPOD-BORNE
DISEASES ENDEMIC IN THE STATE
by
Vernon J. Tipton
and
Robert C. Saunders
BIOLOGICAL SERIES — VOLUME XV, NUMBER 2 AUGUST 1971
TABLE OF CONTENTS
INTRODUCTION I
Colorado Tick Fever 1
The Encephalitides 2
Rocky Mountain Spotted Fever 2
Plague 3
Tularemia 3
Malaria 4
Miscellaneous 5
ACKNOWLEDGMENTS 6
LIST OF ARTHROPODS 7
Arachnida 7
Insecta 8
BIBLIOGRAPHY 19
Acariiia 19
Anoplura 20
Araneida 21
Coleoptera 21
Diptera 21
Hemiptera 24
Hvmenoptera 25
Lepidoptera 25
MaUophaga 25
Orthoptera 25
Scorpionida 25
Siphonaptera 26
Colorado Tick Fever 26
Encephalitis 26
Malaria 27
Plague 27
Rocky Mountain Spotted Fever 28
Tularemia 28
Miscellaneous 29
A LIST OF ARTHROPODS OF MEDICAL IMPORTANCE
WHICH OCCUR IN UTAH WITH A REVIEW OF
ARTHROPOD-BORNE DISEASES ENDEMIC IN THE STATE
by
Vernon J. Tipton and Robert C. Saunders"
INTRODUCTION
The information contained in this paper has been assembled for the express purpose of demonstrating lacunae in current knowledge of health problems associated with arthropods in Utah. Obviously it is not definitive, nor is it intended to be, but rather it should serve as a base for future research investigations. It will be modified as additional infonnation becomes available.
Not all of the arthropods in this list are of proven medical importance— in fact many of them are of doubtful importance— but they are included in the list because they are close rela- tives of species which are of medical signifi- cance in other areas and their potential for af- fecting the health of man has not yet been fully investigated.
Some difficulties are inherent in a review of the history of arthropod-bome diseases, particu- larlv in rural areas. Records are incomplete and their accuracy is suspect. Some arthropod-bome diseases are not included on the standard state report form and thus in manv instances are not reported. In rural areas, where many arthropod- bome diseases occur, people may not seek medi- cal care because of tradition or because no care is available. Diagnostic techniques are slow to reach rural areas and receive broad acceptance. Unless the index of suspicion is high, physicians are prone to ignore diagnostic tests necessary for specific discrimination. However, the history of tularemia in Utah provides ample evidence that rural medicine does not necessarily mean archaic methods and techniques. Dr. Richard A. Pearse, a Brigham City physician, published a clinical description of tularemia in humans which is considered to be the first account in the English language (Jellison, 1971). Many of the early epidemiological investigations of tularemia were conducted in the rural community of Delta and
'Department of /oology. BriRhani Yoiinp Ifniversity, Provo. Utah.
were prompted by the astute observations of local physicians.
Colorado Tick Fever:
The work of Becker (1926, 1930) demon- strated that Colorado tick fever (CTF) is a disease entity distinct from Rocky Mountain spotted fever (RMSF) but it was not until 1940 that Topping, Cullyford, and Davis (1940) pro- vided the first detailed clinical description of CTF. Consequently, accurate data on the inci- dence of CTF prior to 1940 are not available. Records of cases of CTF in Utah from 1940 to 1959 maintained by the Rocky Mountain Labo- ratory in Hamilton, Montana, are probably the most reliable but may be incomplete because information is based on CTF virus isolated from the blood of Utah residents at Hamilton and undoubtedly there were patients who were hos- pitalized elsewhere or not at all. Prior to 1960 the accuracy of records of CTF is in question because diagnostic techniques for the arbovi- ruses were in their infancy. Comparatively sim- ple but reliable tests are available but physicians may not utilize them either because of incon- venience or they may not be aware that the CTF virus persists for approximately 90 days after onset. Because of their epidemiological and clinical similarities CTF has been confused with RMSF. In children CTF may cause encephalitis- like symptoms and even death (Eklund, Kenne- dy, and Casey, 1961 ) but probably there are few inapparent infections in a population. It is pos- sible that many mild cases have escaped detec- tion and have not been reported. According to Pratt and Rice (1969) there were only 96 cases of CTF reported in Utah during the period from 19.56 to 1969 compared with 1,717 cases in Colorado during the same period. However, the low incidence of CTF in Utah mav not be a
Brigham Young University Science Bulletin
true measure of its significance. There is no immunization available and treatment consists of supportive care.
Dennacentor andersoni Stiles is considered to be the most important vector in the epidemio- logical pattern involving man. However, CTF virus has been isolated from ticks of several other species indicating that they may play an important role in the cycle in nature. Ecological studies aimed at detennining epidemiological patterns in nature and particularlv the identity of reservoir animals are needed.
The Enci:i'iialitu)es:
Cases of western encephalitis (WE) and perhaps other encephalitides have probably oc- curred in humans and horses in Utah prior to the outbreak of 1933. However, the etiology and epidemiology of the encephalitides had not yet been elucidated, and one can only speculate about the prevalence of arthropod-bonie viral agents in horses and humans prior to that date. In the Register of Deaths at the Salt Lake City Health Department, Bureau of Vital Statistics, brain fever was listed as the cause of death in several instances during the 1847-1865 period. It is possible that at least some of these deaths were due to infections with arboviruses.
The 1933 episode described by Madsen ( 1934 ) occurred in two waves, both of which began in the West Point area of Weber County. The first wave began about Julv, reached a peak about 10 August, and had almost completely subsided by 1 September. Madsen estimated that 1,139 sick horses were involved in the first wave, of which 43.9 percent died. The second wave, which began the middle of September and subsided about 1 November, involved 2,819 horses, of which about 53.2 percent died. The main focus of infection was in Salt Lake, Davis, Box Elder, Cache, and Weber counties in Utah and Franklin County in Idaho, but there were isolated cases in Rich, Summit, Morgan, Utah, and Tooele counties. An outbreak among hu- mans occurred twenty-five years later (in 1958) in the same general locality in northern Utah and represents the largest number of cases re- ported in Utah for a single year (Jenkins and Donath, 1959). Serological studies revealed a high number of inapparent infections. Thomas and Smith (1959) conducted a survey on infec- tion rates in mosejuitocs, birds, and mammals and found that the highest infection rates in chickens and inos(]uitoes occurred in the geo- graphical center of the human outbreak. In connection with the 1958 outbreak, Rees ct ai. (1959) concluded:
( 1 ) There was in Utah in 1958 a recognized outbreak of Western Etjuine Encephalitis in man; (2) It was accompanied hy a tremen- dous increase in the numbers of C. ttirsalis mosciiiitoes; (3) Culi-x tarsalis moscpiitoes were avidly feeding on man in consideralile numbers during this period; and (4) .Some C tarsalii mosijuitoes were harboring tlic Western equine strain of encephalitis \irus.
Local physicians reported an extensive outbreak of human encephalitis of unconfinned etiology in Box Elder County in 1936. In Weber County in 19.56 there was a serologically confinned case of St. Louis encephalitis and in 1957 a fatal case of SLE plus two cases of WE, one of which was fatal. There were 525 cases of equine en- cephalitis from 1955 to 1969, including 244 cases in the outbreak of 1958. There is no evi- dence of arbovirus activity in Weber County from 1933 to 1955. During recent years along the Wasatch front there have been occasional cases of a disease, difficult to diagnose but typi- cal of western encephalitis. Physicians do not routinely submit acute and convalescent sera necessary for definitive diagnostic tests. Some additional information is needed on feeding preferences of mosquitoes and the role of pas- serine birds in the epidemiology of WE in areas where there have been active foci in the past. The fate of arboviruses during periods between epidemics is a perennial problem recjuiring in- vestigation.
Other arbovinises which have been isolated in Utah include Hart Park-like isolates from Ctilex tarsalis, California encephalitis group iso- lates from Anopheles freehorni, Citliseta inor- nata, Aedes dorsals, A. ninromaculis, Culex enj- tlirotliorax, Psorophora si^niipcnnis, and Culex tarsalis and Cache Valley isolates from Culiseta inonmta and Anopheles freehorni (Holden and Hess, 1959; Crane et al., 1970; and Elbel et al, 1971).
Rocky Moi'xtain Spotted Fe\er:
Beck (1955) reviewed the history- ot Rocky Mountain spotted fever (RMSF) in Utah. It is not certain when RMSF first occurred in the state, but probably the disease was prevalent among early settlers. Beck ( loc cit. ) reproduced ;i newspaper article, published in 1941, in which Dr. William M. McKay, acting commissioner of health for the state of Utah during that period, speculates that Brigham Young was afflicted with RSMF at the time he entered Salt Lake Valley. Some support for this view is supplied by Bvington (in Beck, 1955) who believed that "mountain fever" was the same as RMSF. In the Register of Deaths at the Salt Lake City- Health Department, Bureau of \'ital Statistics,
Biological Series, Vol. 15, No. 2 Utah Arthropods
several deaths are recorded for wliich the cause is listed as "mountain fever." Aldiough there is insufficient evidence to establish a definitive re- lationship between the two, the season during which the deaths occurred is consistent with the epidemiology of RMSF. We do not overlook the possibility that "mountain fever" could be Colo- rado tick fever which also occurs in Utah under similar circumstances. The deaths for which "mountain fever" was listed as the cause oc- curred as follows: two in July and one in August of 1849; one in June and five in July of 1850; two in early November of 1854; and one in late May and two in September of 1855. In at least one instance the cause of death was given as "intermittent mountain fever." In the majority of cases the victims were adult males.
Dermacentor andersoni Stiles is considered to be the most important vector involved in human cases of RMSF in Utah although there may be other species important in the perpetuation of the disease in nature. Beck ( 1955 ) and Coffee (1953) gave data on seasonal and altitudinal distribution as well as life cycles of tick vectors of RMSF.
Jellison ( 1945 ) considers the cottontail rab- bit. Stjlvilaous nuttaUi Bachman to be an im- portant component of the RMSF biocenose. It is a vagile animal with a fairly high degree of ecological tolerance. Studies of the population dynamics of the ectoparasites of the cottontail rabbit may suggest ecological patterns which help to perpetuate the disease in nature.
From 1915 through 1969 the Utah State De- partment of Health, Bureau of Vital Statistics, recorded 496 cases of RMSF in Utah among both residents and nonresidents. Cases occurred most frequently during the months of June, July, and August, although others were reported as early as April and as late as November. Among arthropod-borne diseases with endemic foci in Utah, RMSF is second only to tularemia in total number of cases reported in the state.
Plague:
The recorded history of plague in Utah is not dramatic, but nonetheless, plague represents a disease of great potential significance because of the widespread distribution of capable vectors and reservoirs in the state and the increasing number of fishermen and campers who invade the plague biocenose each year. There have been only two confinned human cases (in 1936 and 1966) and one doubtful case (in 1939) re- ported in the state.
According to Allred (1952) and Beck (1955), capable vectors of plague are indige-
nous to every county in the state, and there are 41 proven reservoirs of plague in Utah, of which 36 are species of rodents. Stark lists 43 species of fleas which are classified as "capable (natu- ral)" or "potential (experimental)" vectors. XenopsijUa cheopis (Rothschild), the most im- portant vector of plague on a worldwide basis, has not been collected in large numbers in Utah. DUimanus mantanitx (Baker) has been con- sidered the most significant vector of plague in Utah, but Parker (1971) oljtained Pasteurella pestis isolates repeatedly from Malaraeus sino- inus (Jordan), Opisodasijs kceni (Baker), Mo- no]).niUiis etimolpi (RothschOd), and Epitedia stanfordi Traub. The number of isolates was highest from specimens of M. sinomus and O. keeni associated with species of Peromijsciis. Parker believes that plague is not limited to squirrel-flea complexes or to particular vegeta- tive associations and topographic patterns. It is evident that vector efficiency varies consider- ably and is influenced by several environmental factors. Thus, there may be several species of rodent fleas with the potential to function effec- tively as vectors of plague as environmental con- ditions change and meet the requirements for transmission by a particular vector species. A study extending over several years relating population fluctuations of fleas on ground squir- rels, on wood rats, and in their nests to environ- mental changes may be helpful in understanding vector and reservoir capabilities.
Tularemia:
Tularemia is a zoonotic disease which has probably been present in the wild fauna of Utah for hundreds of years. Thus it is tempting to speculate about its importance among the pio- neers during the last half of the nineteenth cen- tury. A large segment of the male population in pioneer Utah was engaged in agricultural pursuits, particularly clearing of land, which suggests an invasion of an ecosystem in which transmission of tularemia was likely of common occurrence. Before the advent of white settlers, Indians may have been victims of tularemia inasmuch as rabbits, proven reservoirs of tula- remia, probably constituted a significant part of their diet.
However, recorded history of tularemia in Utah begins about 1908 as indicated by Francis (1925):
There has e.visted in Utah, at least since 1908, a human disease known locally as deer-fly fever. What I believe to be the first clinical reference to luinian cases of tularemia is con- tained in a paper read before the Utah State Medical Association, Salt Lake City, October
Bricham Young University Science Bulletin
3, 1910, 1)\ R. A. Pearse, Brigliam City, Utah. Dr. Pearse refers to si.\ cases, which occurred in the month of August, caused by the bite of a fly, on the exposed parts of the body (neck, ear, clieek, wrist, ankle, and hand ) . After an incubation period of from two to five days ... In 1919 and 1920, 1 studied .seven cases of deer-fly fever near Fillmore, Millard Coun- ty, Utah, and found them positive for tula- remia, clinically, culturally, and serologically. The cases occurred in June, July and August during the seasonal prevalence of the fly Chrijsops discalis. The sites of the fly bites were the neck, temple, ear, and posterior sur- face of the lower third of the thigh. In all cases, suppuration occurred in the glands drain- ing the bitten area. All patients had fever; one died on the twenty-si.\th day of illness. I heard ol [X'rhaps two dozen other cases in the general community in which I worked. From seven- teen jackrabbits, sick or dead, in the commu- nity I isolated Bacterium tularense, thus estab- lishing the great reservoir of infection.
In an earlier publication, Francis (1922) gives a more specific location of tularemia foci in Utah.
So far as known there have been but two foci of infection in Utah. The focus here reported is in Millard County, 5 miles (8 kilometers) west of Holden, .5 miles northwest of Fillmore, 25 miles ( 40 kilometers ) southeast of Delta, and 120 miles (193 kilometers) south of Salt Lake City. The other focus has received clini- cal confirmation and is located near Brigham, a town 20 miles ( 32 kilometers ) north of Ogden in Box Elder County. Both foci have probably existed for at least fifteen years.
Although tularemia does not usually occur in epidemic form, Hillman and Morgan (1937) reported an outbreak of 26 cases among a group of 170 enrollees of a Civilian Conservation Corps camp "located on the treeless plains north of Great Salt Lake." They suggested that the epidemiological evidence available pointed to deer flies as the vectors and jackrabbits as the reservoirs. The cases were diagnosed between 11 and 30 July 1935. There was a noticeable increase in the population of deer flies the week before the onset of the first case. Several men in the camp experienced multiple bites, and lesions on tularemia victims were on uncovered portions of the body. Jackrabbits were numer- ous; many were dead and several were lethargic.
Locomotive Springs, the site of the Civilian Conservation Corps camp, is in the general area of Tremonton where Pearse had seen cases in 1908 and 1910.
Russian workers have proposed subspecific designations for the causative agents of tula- remia which have been accepted by most North
American workers. Francisella tularensis tula- rensis of North America is usually associated with rabbits and arthropods while the more cos- mopolitan tonii, Franci.seUa tidarcnsi.s- palearctica appears to be transmitted independent of arth- ropods and has been isolated from acjuatic or semiaquatic rodents. An organism isolated from a water sample collected in Utah was given the name Francisella novicida ( Larson, VVicht, and Jellison, 1955). All three fomis have been found in Utah. Francisella tularensis tularensis is the principal cause of human tularemia but Fran- cisella tularensi.i palearctica, isolated from musk- rats, should be mentioned because of its im- portance in Utah.
According to the records of the Utah State Department of Public Health, Bureau of Vital Statistics, there have been 986 cases of tularemia in the state during the 45-year period from 1925 through 1969. Approximately three-fourths of these cases occurred during the twenty-year period from 1935 to 19.54.
Chnjsops discalis has been shovra to be an efficient experimental vector of tularemia; it has been known to bite man (Jellison, 1950). For these reasons it has been suspected of being the most important deer fly vector of tularemia in Utah. However, Cox (1965) found C. discalis to be less abundant than C. fulvaster and C. aestuans in study areas near Utah Lake. More- over, he isolated F. tularensis from three of 73 pools of deer flies. Two isolates were obtained from two pools of C. julvaster and one isolate from one pool of C. aestuaivs.
There is a particular need for investigation of seasonal and geographic distribution of spe- cies of Chnjsops and the animals on which they feed, the duration of infection in reservoirs and vector species, and serological survevs of human populations in areas where there are high den- sitv populations of deer flies.
M.\L.\niA;
Most cases of malaria which have occurred in the state were contracted elsewhere, but Marshall and Rees (1948), in their excellent review of malaria in Utah, have provided sub- stantial evidence that local transmission has taken place, particularly in southern Utah. They point out that most of the early Utah settlers came from the Mississippi Valley where ma- laria was prevalent. Contact with the outside world was maintained through continuing im- migration, returning missionaries, and settlers passing through on their way to California or Oregon. Perhaps the only case of malaria in Utah sufficiently well documented to be consid-
Biological Series, Vol. 15, No. 2 Utah Arthropods
ered autochthonous is cited by them as follows: "In April, 1947, a vivax infection was reported in a two-year-old child of that area who had never been out of the state." The "area" referred to is southern Utah. Anopheles freeborni Aitken is widespread throughout the state and Anopheles franciscanus McKracken is widespread through- out the southern half of the state ( Nielsen, 1968). Both are considered to be efficient vec- tors of malaria, especially the fonner.
In the Register of Deaths at the Salt Lake City Health Department, Bureau of Vital Statis- tics, there are several entries in which "malig- nant fever" and "bilious fever" are listed as the cause of death. Early physicians made the dis- tinction among "fever," "mountain fever," "ma- lignant fever," "bilious fever," and "typhoid fever," and although it would be inaccurate to associate malignant fever or bilious fever with malaria, there is a possibility that a persistent fever occurring during the srunmer months could be malaria.
In a five-year period from 1943 through 1947, 723 cases of malaria were reported in Utah and reflect the impact of returning servicemen on the incidence of disease within the state. There was another less dramatic rise in the incidence of malaria in Utah, associated with the Korean War, during the period from 1951 to 1955 when 75 cases were reported. Nevertheless, with an adequate reservoir of infection, capable vectors, and a susceptible resident population, malaria has not become established in Utah. Rapid diagnosis and treatment of servicemen, improved mosquito control, and an infonned public are the principal factors which mitigate the impor- tance of malaria in Utah. The feeding habits of the mosquito vectors in a rural setting may be another factor of some importance. Cattle and horses are the preferred sources of blood meals for some Anopheles species.
Miscellaneous:
Relapsing fever is virtually unknown in Utah, although it has been reported on several occa- sions from surrounding states. Davis (1939) reported a single case which occurred near Salt Lake City in 1928. Both Ornithodoros parkeri and O. turicata, proven vectors of relapsing fever, occur in the state although their distribu- tion is not completely known.
Coxiella burnetti, the causative agent of Q fever, has been isolated from rodents {Di- podomijs ordii, D. microps, and Pewmysrus maniculatus) and a tick {Demwcentor paru- mapterus) in the Great Salt Lake Desert in Utah. C. burnetti antibodies were demonstrated
serologically in Lepus californicus, Omjchomijs leucogaster, and Eutainias minimus ( Stoenner et al., 1959). There is no record of Q fever in
Ca.se.s of Selected Arthropod-Bome Diseases in Utah 1915-1969
|
Year |
CTF |
WE |
Malaria |
RMSF |
Tularemia |
|
1915 |
35 |
||||
|
1916 |
34 |
||||
|
1917 |
15 |
||||
|
1918 |
5 |
||||
|
1919 |
10 |
||||
|
1920 |
9 |
||||
|
1921 |
8 |
||||
|
1922 |
18 |
||||
|
1923 |
15 |
||||
|
1924 |
10 |
||||
|
1925 |
2 |
6 |
1 |
||
|
1926 |
• |
5 |
4 |
||
|
1927 |
7 |
1 |
|||
|
1928 |
1 |
10 |
1 |
||
|
1929 |
11 |
0 |
|||
|
1930 |
13 |
4 |
|||
|
1931 |
10 |
2 |
|||
|
1932 |
22 |
4 |
|||
|
1933 |
5 |
6 |
|||
|
1934 |
1 |
12 |
6 |
||
|
1935 |
3 |
14 |
37 |
||
|
1936 |
8 |
5 |
|||
|
1937 |
2 |
12. |
41 |
||
|
1938 |
10 |
19 |
73 |
||
|
1939 |
2 |
1 |
24 |
44 |
|
|
1940 |
2 |
2 |
5 |
15 |
55 |
|
1941 |
5 |
0 |
13 |
45 |
|
|
1942 |
0 |
5 |
8 |
48 |
|
|
1943 |
12 |
313 |
12 |
34 |
|
|
1944 |
5 |
157 |
10 |
23 |
|
|
1945 |
2 |
112 |
9 |
28 |
|
|
1946 |
4 |
93 |
5 |
29 |
|
|
1947 |
6 |
48 |
5 |
36 |
|
|
1948 |
2 |
2 |
4 |
45 |
|
|
1949 |
1° |
3 |
2 |
14 |
35 |
|
1950 |
5 |
0 |
7 |
40 |
|
|
1951 |
1° |
.7 |
23 |
18 |
30 |
|
1952 |
1" |
8 |
38 |
9 |
20 |
|
1953 |
1° |
6 |
7 |
9 |
40 |
|
1954 |
2" |
6 |
4 |
4 |
22 |
|
1955 |
3" |
2 |
3 |
7 |
29 |
|
1956 |
3 |
1°° |
0 |
3 |
21 |
|
1957 |
4 |
3" |
1 |
9 |
30 |
|
1958 |
0 |
47 |
0 |
0 |
17 |
|
1959 |
4 |
38 |
2 |
1 |
10 |
|
1960 |
4 |
23 |
0 |
3 |
19 |
|
1961 |
2 |
6 |
0 |
1 |
17 |
|
1962 |
8 |
8 |
0 |
0 |
16 |
|
1963 |
12 |
12 |
0 |
0 |
3 |
|
1964 |
12 |
1 |
0 |
1 |
24 |
|
1965 |
11 |
0 |
1 |
0 |
10 |
|
1966 |
4 |
1 |
3 |
0 |
3 |
|
1967 |
11 |
0 |
3 |
0 |
7 |
|
1968 |
5 |
0 |
1 |
0 |
5 |
|
1969 |
16 |
0 |
3 |
2 |
16 |
"Represents isolations from patients hospitalized at Rocky Mountain Laboratory, Hamilton, Montana.
•"One case of SLE.
Brigham Young Univebsitv Science Bulletin
humans in Utah, but (liis may be due to faulty diagnosis or reporting, inasmuch as human cases have occurred in surrounding states.
Mohr { 1951 ), in his paper on the distribution of murine typhus and plague in the United States, gives no records of murine typhus for the state of Utah. The flea index of XenupsijUu cheopis on Rattus sp. apparently has never been very high in Utah, and probably accounts for the absence of the disease in the state.
Annstrong (1922) reported an epidemic of typhus on the San Juan Indian Reservation dur- ing the last half of 1920 and the first half of 1921 in which there were 6.3 cases of tyjDhus with 27 deaths among appro.ximatelv 7,000 Indians. The San Juan Indian Reservation is 5,884 square miles in the four-comers area of New Mexico, Arizona, and Utah.
One case of dengue was reported in 1942, but it was probably contracted outside the state.
One doubtful case of rickettsialpox has been
reported from Utaii (Pratt and Rice, 1969).
In Utah the incidence of bites and stings of arthropods and the number of cases of derma- titis caused by urticating and vesicating insects is unknown. Scattered cases of archnidism have been reported, including at least one death from the sting of a hymenopterous insect. Latrodectus liesperus Chamberlin and Ivie and several spe- cies of Hymenoptera are the most important venomous arthropods in the state.
Tick paralysis, caused by the bite of female ticks, Dennacentor andersoni Stiles, occurs most freijuently in an area comprising the north- em part of Idaho and adjacent portiorLs of Wash- ington and Montana. Isolated cases have been reported in other sections of the Rocky Moun- tains where Dennacentor andersoni occurs (Philip, 1969). Insofar as we are aware there have been no cases of tick paralysis reported in Utah, but the possibility of its occurrence should not be overlooked.
ACKNOWLEDGMENTS
We are indebted to many people who have contributed time and effort to minimize the number of errors in the list of arthropods and who have made suggestions concerning the re- view of arthropod-bome diseases in Utah. In particular we are grateful to Dr. Karl V. Krom- bain, chainnan. Department of Entomology, Na- tional Museum of Natural History, Smithsonian Institution and his associates at the National Museum and the Systematic Entomology Labo- ratory, United States Department of Agricul- ture: Dr. Ralph E. Crabhill, Jr., Dr. K. C. Emerson, Dr. Douglas C. Ferguson, Dr. William D. Field, Dr. Richard C. Froeschner, Dr. Ash- ley B. Gumey, Dr. Jon L. Herring, Dr. Arnold S. Menke, Dr. C. F. W. Muesebeck, Dr. Curtis W. Sabrosky, Dr. Reese I. Sailer, Dr. D. R. Smith, Dr. Robert Traub, and Dr. Willis W. Wirth. Others who have reviewed the manu- script and have made helpful suggestions are Dr. Don M. Rees, Dr. Lewis T. Nielsen, Dr. Robert Winget, and Mr. Bmce Knudson, De- partment of Biology, University of Utah; Dr. Wilford J. Hanson and Dr. George F. Knowlton, Utah State Universitv; Dr. James M. Brennan, Dr. William L. Jellison, and Mr. Glen M. Kohls. Rocky Mountain Laboratory, Hamilton, Mon-
tana; Dr. Robert E. Elbel, Deseret Test Center, Dugway, Utah; Dr. Dale D. Parker, Research Entomologist, EcoDynamics, Salt Lake City, Utah; Dr." Willis J. Gertsch, Portal, Arizona; Dr. Harold E. Stark, Center for Disease Control, Fort Collins, Colorado; Dr. Phyllis T. Johnson, Costa Mesa, Califomia; Dr. Dorald M. Allred, Dr. Ferron L. Andersen, Dr. Gerald L. Hay- ward, Dr. Vasco M. Tanner, and Dr. Stephen L. Wood, Department of Zoology, Brigham Young University, Provo, Utah; Dr. Donald N. Wright, Department of Microbiology, Brigham Young University, Provo, Utah; Dr. Maurice T. James, Dr. William Tumer, and Mr. Craig R. Baird, Wa.shington State University, Pullman, Wash- ington; Dr. Deane P. Funnan, University of Califomia, Berkeley, California; Dr. Kenneth J. Capelle, Bear River Research Station, Brigham City, Utah; Dr. Harry Hoogstraal, U. S. Naval Medical Research Unit No. 3, Cairo, Egypt; Dr. Bobbie V. Peterson, Entomologv Research Insti- tute, Ottawa, Ontario, Canada. We are also in- debted to Dennis I. Anderson and Lynden P. Baum for their assistance with accumulation of the scientific names for the list and to Phyllis Madden for typing the manuscript.
Biological Series, Vol. 15, No. 2 Utah AiniiHOPons
LIST OF ARTHROPODS
ARACHNID.\
1. Acarina: I.\odicles
A. Argasidae
1. Argas
A. cooleij! Kohls and Hoogstraal, 1960 A. gigantcus Kohls and Clifford, 1968 A. sanchezi Duges, 1887
2. Ornithodoros
O. concanensis Cooley and Kohls, 1941
O. ereviicus Cooley and Kohls, 1941
O. kelleyi Cooley and Kohls, 1941
O. parked (Cooley, 1936)
O. spanius KoMs and Clifford, 196.3
O. talaje (Guerin-Meneville, 1849)
O. turicata (Duges, 1876)
3. Otobius
O. megnini (Duges, 1884)
0. lagophilus Cooley and Kohls, 1940
B. Ixodidae
1. Dermaccntor
D. albipictiis (Packard, 1869)
D. andcrsoni Stiles, 1908
D. hitnteri Bishopp, 1912
D. parumapertus Neumann, 1901
2. Haemaphijsalis
H. leporispahistris (Packard, 1869)
3. Ixodes
1. angustus Neumann, 1899
/. jellisoni Cooley and Kohls, 1938
/. kingi Bishopp, 1911
I. marmotae Cooley and Kohls. 1938
/. 7r!i/r!,s Bishopp and Smith, 1937
I. ochotonae Gregson, 1941
7. pacificus Cooley and Kohls, 1943
7. sculptus Neumann, 1904
7. soricis Gregson, 1942
7. spinipaljus Hadwen and Nuttall, 1916
7. texaniis Banks, 1908
7. iLoodi Bishopp. 1911
4. Rhipiceplialus
R. sanguineus ( Latreille, 1806)
r. Acarina: Me.sostigmata
A. Dermamjssidae
1. Dermamjssus
D. gallinae ( De Geer, 1778 )
2. Hirstionyssus
H. bisetosus AUred, 1957
77. cynomijs (Radford, 1941)
77. eutamiue Allred and Beck, 1966
H. fcmuralis Allred, 1957
H. hilli (Jameson, 1950)
77. incomptus (Eads and Hightower, 19.52)
77. imbellinus (Oudemans, 1913)
77. latisctttatus (de Meillon and Lavoipierre,
1944) 77. longichelac Allred and Beck, 1966 77. neotoniae (Eads and Hightower, 1951) 77. occidentalis ( Ewing, 1923) 77. paraffinis Herrin. 1970 77. perognathi Herrin, 1970 77. staff ordi Strandtmann and Hunt, 1951
77. talpae Zemskaya, 1955 77. thomomys Allred and Beck, 1966 77. torus Allred and Beck, 1966 77. triacanthus (Jameson, 1950) 77. utahensis Allred and Beck, 1966
3. Lipomjssokh's
L. becki (Allred, 19.57) L. sanguineus (Hirst, 1914)
4. Myonyssits
M. montanus Furman and Tipton, 1955
B. Haemogamasidae
1. Brevisterna
B. montanus (Ewing, 1922) B. utahensis (Ewing. 1922)
2. Eulaclaps
E. stabularis (Koch, 1836)
3. Haemogamasus
77. alaskensis Ewing, 1925 77. barberi (Ewing, 1925) 77. nidiformis Bregetova, 1955 77. occidentalis ( Keegan, 1951 ) H. pontigcr ( Berlese, 1903)
4. Ischyropoda
7. armatus Keegan, 1951 7. furmani Keegan, 1951
C. Halarachnidae 1. Zumptiella
Z. bakeri (Furman, 1954)
D. Laelapidae
1. Androlaelaps
A. circularis (Ewing, 1933) A. crowd Jameson, 1947 A. debilis Jameson, 1950 A. fenilis (Megnin, 1876; A. geomys Strandtmann. 1949 A. glasgowi (Ewing, 1925) A. hollisteri (Ewing, 1925) A. leviculus Eads, 1951
2. Hi/poaspis
H. gurabcnsis (Fox, 1946)
77. lubrica Oudemans and Voigts, 1904
3. Laelaps
L. incilis Allred and Beck, 1966 L. kochi Oudemans, 1936 L. multispinosus Banks, 1909 L. nuttalli Hirst, 1915
E. Macronyssidae
1. Chrioptonyssus
C. rohustipes (Ewing, 1925)
2. Ornithonyssus
O. aridus Furman and Radovsky, 1963
O. hacoti (Hirst, 1913)
O. silviarum ( Canestrini and Fanzago. 1877)
3. Steatonyssus
S. antrozoi Radovsky and Furman, 1963
F. Rhinonyssidae
1. Paraneonyssus
P. icteridius Strandtmann and Furman, 1956
Bhicham Young University Science Bulletin
G. Spintumiciclae
1. Paraspinturnix
P. glohosus Rudnick, 1960
2. Spinturnix
S. orri Rudnick, 1960
Acarina : Orihatei Garabodidac
Passalozelcs
P. linearis Higgins and Woolley, 1962
Acarina : Trombidiformes A. Trombiculidac
1. Acomatacarus
A. arizonensis Kwing, 1942
2. Chatia
C. ochotona (Radford, 1942) C. setosa Brennan, 1946
3. Cheladontd
C. crossi Lipovskv, Crosslev, and Loomis, 1955
4. Euschoengastia
E. cordiremus Brennan, 1948
E. crkcticohi Brennan, 1948
E. ct/noriu/icola Crossley and Lipovsky, 1954
E. (iccipicm Gould, 1956
E. fa-soUa Brennan and Beck, 1955
E. fiirmcini Gould, 1956
£. hoffimmac Gould, 1956
E. lanccolata Brennan and Beck, 1955
E. land Brennan and Beck, 1955
E. luteodcma Brennan. 1948
E. oltcm Brennan and Beck. 1955
E. oregonensis (Ewing, 1929)
E. pomcrantzi Brennan and Jones, 1954
£. radfordi Brennan and Jones, 1954
E. rotunda Brennan and Beck. 1955
E. sciuricola (Ewing, 1925)
E. soricinus Gould, 1956
5. Euschoengastoides
E. hcerfii ( Brennan, 1948 )
E. hoplai (Loomis, 1954)
E. utahensis (Brennan and Beck, 1955)
6. Eutrnmbicula
E. hclhini (Gould, 1950)
7. Hexidioni.i
H. allredi (Brennan and Beck, 1955) H. doremi (Brennan and Beck. 1955)
8. Hi/poncocula
H. iircnicola (Loomis, 19S4) //. ntontanensis (Brennan, 1946)
9. Gahrlicput
G. (imericcina (Ewing, 1942)
10. Leeuwenhoekia
L. americuna (Ewing, 1942)
11. Leptotroml)idiuni
L. mijotis (Ewing, 1929)
L. panamenxi.'! (Ewing, 1925)
L. potosina (Hoffman. 1950)
12. MiijatromhicuUi
M. esoenii- (Sxsa and Ogata, 1953) M. sargenti ( Brennan, 1952
13. Neoschocngastki
N. americuna (Hirst, 1921)
14. Neotrond>icuUi
N. calif omica ( Ewing, 1942 )
N. harperi (Ewing, 1928)
N. jcwetti (Brennan and Wharton, 1950)
N. microti (Ewing, 1928)
N. subsignata (Brennan and Wharton, 1950)
15. Odontacarus
O. hirsutiis (Ewing, 1931 )
O. lin.sdalei (Brennan and |ones, 1954)
O. micheneri Grcenberg, 1952
16. Tromhicida
T. hakcri Ewing, 1946 T. kardosi Loomis, 1954 T. univari Brennan. 1965
17. Whartonia
W. perplexa (Brennan, 1947)
B. Mvobiidae Radfordia
R. hachai Howell and Elzinga, 1962 R. krninu (Koch, 1841) R. suhuliger Ewing, 19.38
II. Araneida
A. Laxoscelidae Loxosceles
L. unicolor Keyserling, 1887
B. Theridiidae Latrodectus
L. hesperui Chamlierlin and hie. 1935
III. .Scorpionida
A. Buthidae Centntroides
C. sculpturatus Ewing. 1928
B. Vejovidae
1. Atiuroctonus
A. phaiodactylus (Wood, 1863)
2. Hadrunis
H. arizonensis Ewing, 1928 H. spadix, Stahnke, 1940
3. Vejovis
V. hecki Gertsch and Allred, 1965
V. horeus (Girard, 1845)
X . roncusus Stahnke, 1940
V. ntahcnsis Williams, 1968
V. wapatkiensis Stahnke, 1940
INSEGTA
I . .\noplura
A. Haematopinidae
Ilaematojrinus
H. asini (Linnaeus, 1758)
H. cunjstemus ( Nitzsch, 1818)
H. stiis (Linnaeus, 1758)
B. Hoplopleuridae 1. Enderleinellus
E. murmotae Ferris, 1919
E. oshorni Kellogg and F"erris. 191.5
E. paralongiccps- Kim, 1966
E. suturalis (Osboni. 1891)
E. tamiasciuTi Kim, 1966
Biologic AL Series, \'ol. 15, No. 2 Utah Akthhoi'oos
2. Fahrcuholzia
F. piuimlii Kellogg ;\nd Ferris, 1915 F. reductu Ferris, 1922
3. Haemodipsus
H. lijriocephalus ( Burmeister, 1839 )
H. setoni Ewing, 1924
H. ventricostis (Denny, 1842)
4. Hoploplcum
H. actintlwpu.'! (Burmeister, 1839)
H. (irhnricolu Kellogg and Ferris, 1915
H. ciiptiosci Johnson, 1960
H. difjicilis Kim, 1965
;/. erratku (Osbom, 1896)
H. ferrisi Cook and Beer, 1959
H. hesiicroini/dix ( Oslxim, 1891)
H. ontjchonn/di.i Cook and Beer, 1959
H. pacified Ewing, 1924
H. reithrodontdmi/dis Ferris, 1951
H. sciuricoUi Ferris, 1921
H. trisjtinosd Kellogg and Ferris, 1915
5. Neohaenuitopinus
N. citellinus Ferris, 1942
N. inonuitus (Kellogg and Ferris, 1915)
N. laeviunculus (Gnibe, 1851)
IV. marmotiic Ferris, 1923
N. neotomtie Ferris, 1942
N. pacificus Kellogg and Ferris, 1915
N. sciuri Jancke, 1931
N. sciumpteri (Osbom. 1891)
N. semifasciatus Ferris, 1916
iV. spilosonmc Pratt and Stojanovicli, 1961
6. Pnhjphix
P. ahiskcnvi.s Ewing, 1927
P. auricularifi Kellogg and Ferris, 1915
P. borealis Ferris, 1933
P. serratu ( Burmeister, 1839 )
P. spinulosa (Burmeister. 1839)
C. Linognathidae
1. Linogiuithu.s
L. africamis Kellogg and Paine, 1911
L. pedalis (Oshom, 1896)
L. setosus (\on Olfers, 1816)
L. stcnopsix (Burmeister, 1838)
L. vitidi (Linnaeus, 1758)
2. Solenoptcs
S. Innipilosus ( Fahrenholz, 1916) S. capiUdtus Enderlein. 1904 S. ferrisi (Fahrenholz, 1916)
D. Pediculidae
1. Microphthinis
M. itncinatus (Ferris, 1916)
2. Pedicidus
P. humanus Linnaeus, 17.58
3. Pthinis
P. pubis (Linnaeus, 1758)
II. Orthoptera A. Blattidae
1. Arcm'taga
A. errdtica Kehn, 1907
2. Blatta
B. orientalis (Linnaeus, 1758)
3. Bktella
B. germanica (Linnaeus. 1767)
4. Punchlord
P. nivea (Linnaeus, 1758)
5. Pcriphmctd
P. dtnericdiid (Linnaeus, 1758) P. dustrdlidsiuc ( Fabricius, 1775)
6. Suppella
S. longipalpd ( l'"abricius, 1798)
III. Coleopiera
A. Leptinidae
1. Lcptinillus
L. vdlidus (Horn, 1872)
2. PUitypsijllus
P. cdstoris Ritsema, 1869
B. Meloidae
1. Epicauta
E. fdhricii ( LeConte, 1853)
E. ferrugincd (Say, 1823)
E. normalis Werner, 1945
E. puncticoUis ( Mannerheim, 1843)
2. Lijtfci
L. ciiduipcunis (LeConte, 1851)
3. Nemognathd
N. luridd LeConte, 1853 N. hitea LeConte, 1853
IV". Diptera
A. Calliphoridae
1. Aldrichina
A. grahami ( Aldrich. 1930)
2. Bufolucilia
B. silvarurn (Meigen, 1826)
3. CdUiphord
C. coloraden-sis Hough, 1899 C. lividd Hall, 1948
C. ferraenovcie Macquart, 1851
C. viciiui Robineau-Desvoidy, 1830
C. vomitorid (Linnaeus, 1758)
4. Cochliomtjid
C. hoininicordx (Coqueral, 1858) C. mdccllarid (Fabricius, 1775)
5. Cynomyopsis
C. Cdddvcriiui (Robineau-Desvoidy, 1830)
6. Eucdlliphord
E. lilaed (Walker, 1849)
7. Lucilia
L. iUustris (Meigen, 1826)
8. Phdenicia
P. sericata (Meigen, 1826)
9. Phormia
P. regina (Meigen, 1826)
10. PoUcnid
P. rudis (Fabricius, 1794)
11. Protoc(dliphord
P. aened Shannon and Dobroscky, 1924
P. asiovord Shannon and Dobroscky, 1924
P. cuprind (Hall, 1948)
P. hespcrid Shannon and Dobroscky, 1924
P. hirudo Shannon and Dobroscky, 1924
P. hirundo Shannon and Dobroscky, 1924
P. metalUcd (Townsend, 1919)
P. sialia Shannon and Dobroscky, 1924
BiUGHAM YouNc; University Science Bulletin
12. Proloplionniu
P. terraenovae ( Robineau-Desvoidy, 1830)
B. Ceratopogonitlae
1. Culicokles
C. haucri Hoffman, 1925
C. cochiscnsls Wirth and Blanton. 1967
C. cockercllii (Co(iiiillett, 1901)
C. crcpitsculari.s Malloch, 1915
C. frcchonii W'irlli and Blanton, 1969
C. liucmatojHttiis Malloch, 1915
C. hwrofili/pliiciis Malloch, 1915
C. nwntiintis Wirth and Blanton, 1969
C. obsoletus (Meigen, 1818)
C. palnwriie James, 1945
C. stellifer ( CoquiUett, 1901)
C. usinr^eri Wirth, 1952
C. iituhcnsis Fox, 1946
C. variipennis variipennis (Coquillftt, 1901)
2. Leptoconops
L. kerteszi Hieffc-r. 1908
C. Chloropidac" Hippelates
H. microccntriis Coqiiillett, 1904
H. muntunn.s Sabrosky, 1941
H. poUipcs (Locw, 1865)
//. piirliceps (Becker, 1912)
H. pu.sio Loew, 1872
D. Cuhcidae
1. Aedes
A. atropalpus (CoquiUett, 1902)
A. cumpestris Dyar and Knab, 1907
A. cdtaphijUn Dyar, 1916
A. cincrciis Meigen, 1818
A. conununk ( De Geer, 1776)
A. clorsalis (Meigen, 1830)
A. excnicians (Walker, 1856)
A. fitchii (Felt and Young, 1904)
A. fhvcsccm ( Muller, 1764)
A. hexodnntus Dyar, 1916
A. impiger (Walker, 1848)
A. implicatus Vockeroth, 1954
A. increpitu.i Dyar, 1916
A. intrudens Dyar, 1919
A. melanimon Dvar. 1924
A. nickeni O'Meare and Craig. 1970
A. ;iigr(;mnri(/(.v ( Ludlow, 1907 )
A. nipluidopsix Uvar and Knab, 1918
A. pulUitm (Coqiiillett, 1904)
A. schizopinax Dyar, 1929
A. sierrenxis (Ludlow, 1905)
A. spencerii idcihocnsis (Theobald. 1901)
A. sticticus (Meigen, 1838)
A. trivittatus (Cociuillett, 1902)
A. varipalpm ( Co(|uillett, 1902)
A. ventrovitlis Dyar, 1916
A. vexans (Meigen, 1830)
2. Anopheles
A. earlei Vargas, 1943
A. fr(mcisciinu.i McCracken. 1904
A. freehorni Aitken, 1939
3. Coquillettidia ( —Mansonia)
C. pcrturlxnu (Walker, 1856)
4. Culex
C. upiccdis Adams, 1903 C. erylhrothorax Dyar, 1907
C. pipiens pipicn.s Liimaeus, 1758 C. pipienx (piincjucjasciatus .Say, 1823 C. restuans (Theobald, 1901) C. tarsalis Coiiuillett, 1896 C. territmi.s Walker, 1856 C. thruDuhiis Dyar, 1921
5. Culiseta
C. iTtipaticns (Walker, 1848)
C. incidcns Thompson, 1868
C. mormita Williston, 1893
C. morsituns di/ari Coc|uillett, 1902
C. sihcstri.s ininncsotac Barr, 19.57
6. Orlhopodiiini/ui
0. si^nifcni ( Cociuillett, 1896)
7. Psorophora
P. signipennis (CoquiUett, 1896)
E. Cuterebridae Cuterebra
C. cmgustifrniu Dalmat, 1942 C. aj)j>roxiniala Walker, 1866 C. grusca Cixinillett, 1904 C. jcllisoni Curran, 1942 C. k'pusculi Townsend, 1897 C. polifa Cocpiillett, 1898 C. princeps (Austen, 1895) C ruficnis (Austen, 1933) C. tcnchrosa Cocpiillett, 1898
F. Gasterophilidae Gastcrophilus
G. haemorrhoklalis (Linnaeus, 178.5) G. intcstinaUs ( De Geer, 1776) G. nasalis (Linnaeus, 17.58)
G. Hippoboscidae
1 . Icosta
1. americana (Leach, 1817) /. hirsuta (Ferris, 1927)
/. nigra (Perty, 1833)
2. Lipoptena
L. deprcssu ( Sav, 1823)
3. Mclophagus
M. ovinu.s- (Linnaeus, 1785)
4. MijophthirUi
M. fiml)riat<i ( Waterhouse, 1887)
5. Ncolipopteiui
N. fcrhsi (Be(|uaert, 1935)
6. Olfersw
O. .'iordidii Bigot, 1885
7. Ornithinni/u
O. (iiichincuriii .Speiser, 1905
8. Ornithoicii
O. vicirw (Walker, 1849)
II. .Muscidae
1. Fannia
F. caniculam ( Linnaeus, 1761 ) F. scalaris ( Fabricius, 1794)
2. llacmatohia
H. irritans Linnaeus, 1785
3. Miiscti
XL tiuliinuudis De Geer, 1776 A/, domestica (Linnaeus, 1758)
Biological Sehies, Vol. 15, No. 2 IIiaii Akihhoi-ods
11
4. Muscina
M. (issimilis (Fallen, 1823) M. stdluihim (FalU'-n, 1817)
5. Stomoxij.t
S. ciilcitnius (Linnaeus, 1758)
I. Nycteribiidae Basilia
B. aiUrozoi (Townsend, 1893) B. con/iwrltini (Ferris, 1916)
B. forcipata Ferris, 1924
J. Oestridae
1. Cephcncmijia
C. jclliioui Townsend, 1941 C. pratti Hunter. 1916
2. Hypoderma
H. hovis (Linnaeus, 1758) H. lincutum ( N'illers, 1789)
3. Oestrus
O. avis Linnaeus, 1758
K. Piophilidae Piophila
P. ciisci (Linnaeus, 17.58)
L. Psychodidae
1. Lntzomijia
L. (uptilonia ( Fairchild and Harwood, 1961) L. cidifornicii ( Fairehild and Hertig, 1957) L. oppidanii ( Dampf , 1944) L. stewarti ( Mangaheira and Galindo, 1944 ) L. vexafor (Coqu"illctt, 1907)
2. Psychodii
P. (dtcrnata Say, 1824
M. Rhagionidae Symphoromyia
S. atripes Bigot, 1887
S. fuhipcs Bigot, 1887
S. hirta Johnson. 1897
S. iiuiuKitor Aldrich, 1915
S. johiisuiii Coquillett, 1894
S. pachyceras Williston, 1886
N. Sarcophagidae
1. Ravinuj
R. acerba (Walker, 1849)
R. derelict (I (Walker, 18.52)
R. erndmnda (Wulp. 1895)
R. latisetosii Parker, 1914
R. Ilicrrninieri ( Robineau-Desvoidv, 1830)
R. plaiiifrons (Aldrich, 1916)
R. pusiola (Wulp, 1895)
2. Sarcophagu
S. argyrostoma ( Robineau-Desvoid) , 1830 )
S. bishoppi .Mdrich, 1916
S. buUata Parker. 1916
S. coolet/i Parker, 1914
S. haemorrlwidalis (Fixllen, 1817)
S. perspicax, .'\ldrich, 1916
S. sarriicenioides Aldrich, 1916
S. slierinani Parker. 1923
S. sinuata Mcigen, 1826
S. utdis Aldrich, 1915
3. Wohlfahrtia
W. vigil opaca Coquillett. 1897
O. Simuliidae
1. Cnephia
C. jeatuie DeFoliart and Peterson, 1960
C. mutata ( Malloch, 1914)
C. villosa DeFohart and Peterson, 1960
2. Prosimuliuin
P. daviesi IVterson and DeFoliart, 1960
P. exigens Dyar and Shannon, 1927
P. flaviantentiiim (Stains and Knowlton,
1940) P. lulviim ( Coiiuillett, 1902) P. longilobuin Peterson and DeFoliart,
1960 P. onyclwdtieti/lum Dyar and Shannon, 1927 P. shewelti Peterson and DeFoliart, 1960 P. travisi Stone, 19.52 P. ttintn Peterson and DeFoliart, 1960 P. i(nici/?(i (Twinn, 1938)
.3. Simidium
S. iircticum Malloch, 1914
S. argus Williston, 1893
S. tiureiim Fries, 1824
S. bicorne Dorogostaj.skij, Rubtzov, and
Vlasenko, 1935 S. bivittatum Malloch, 1914 S. canudcnse Hearle, 1932 S. canonicola ( Dyar and Shannon. 1927) S. corbis Twinn, 1936 S. decorum Walker, 1848 S. defoliurti Stone and Peterson, 1958 S. griseum Coquillett, 1898 S. huntcri Malloch, 1914 S. jacumbae Dvar and Shannon, 1927 S. latipes (Meigen, 1804) S. mediovittatum Knab, 191.5 S. meridionale Riley, 1887 S. nigricoxum Stone, 19.52 S. pctersoni Stone and DeFohart, 1959 S. piperi Dyar ;uid Shannon, 1927 S. pugetense ( Dyar and Shannon, 1927 ) S. rugglesi Nicholson and Mickel, 1950 S. trivittatum Malloch, 1914 S. tuberosum ( Lundstrom, 1911) S. ventitor Dyar and Sh;uinon, 1927 S. venustum Say, 1823 S. virgatum Coquillett. 1902 S. vittafum Zetterstedt, 1838 S. wyomi7igense Stone and DeFoliart, 1959
4. Twinnia
T. nova ( Dyar and Shannon, 1927 )
P. Streblidae 7'ric7io/;iH.s
T. coripiorhini CockereU, 1910 7". major Coquillett, 1899
y. Syrphidae
Eristalis
E. dimidiiitas Wiedemann, 1830 E. tenax (Linnaeus, 1758)
R. Tabanidae
1 . Atylotus
A. i7icisur(dis var. utahensia Rowe Knowlton, 1935
2. Chrysops
C. aestuans Wulp, 1867
C. callidus Osten Sacken, 1875
and
12
Brigham Young Univehsitv Science Bulletin
C. carbonarius Walker, 1848
C. cociuilh'tii Him-, 1904
C. discalin Williston, 1880
C. cxcitans Walker, 1850
C. fri«iclus Osten Sacken, 1877
C. juhiister Osteii Sacken, 1877
C. fiircdiux Walker, 1848
C. i>i(/((,v Osten Sacken, 1875
C. fjiiVi.v Osten Sacken, 1875
C. nigcr Maciinart, 1838
C. nociifcr noctifcr Osten Sacken, 1877
C. noctifcr pcrtinux Williston. 1887
C. pachijccrufi Williston, 1887
C. sackcnii Mine, 1903
C. i(i/c!/(jc Philip, 1955
3. Hacinatopotii
II. (imericimu Osten Sacken, 1875
4. Uijhomitru
II. epistatcs (Osten Sacken, 1878)
H. jmntalh (Walker, 1848)
H. opacii (Coqiiillett, 1904)
H. rhomhicd (Osten Sacken, 1876)
H. rliomhica var. nshurni ( Hine, 1904)
//. nipestris ( MeDonnough, 1921)
//. .•scquiix (Williston, 1887)
H. .wnoiiicnsi.'i var. phaenops ( Osten Sacken,
1877) H. tetrica var. Iiirtida (Bigot, 1892)
5. Pilinms
P. californiciis (Bigot. 1892)
6. Silvius
S. (pimlrhittatus (Say, 1823)
7. Stenotahanus
S. fhwiclus (Hine, 1904)
S. giittdtultis (Townsend, 1893)
8. Tahanus
T. acgrotm Osten Sacken. 1877
T. iitmtus Fabricius, 1775
T. clor.tifcr Walker. 1860
T. gi/rtHi/.v Townsend, 1897
T. laticcps Hine, 1904
T. lineohi Fahricins, 1794
T. proiUictus Hine, 1904
7'. ptimilus Macquart, 1838
T. punctifcr Osten Sacken, 1876
r. .sYoiK'i Philip, 1941
V'. Hemiptera
A. Cimicidae
1. Cimex
C. Iccttilarius Linnaeus. 1758 C. piliKclliLs (Horvath, 1910)
2. OecUicua
O. vicarhis Horvath, 1912
B. R<'duvjidae
1. Rcdiiviu.'i
R. pcrsoiuilu.t ( Linnaeus, 17.58 ) R. minduzvei Wvgodzinskv and Usinger, 1964
2. Triatomii
T. protract,! (Uhler. 1894)
V'l. Hymenoptera A. Apidae 1. Apis
A. mellifera Linnaeus, 1758
Bombus R. ii)>positii.s Cresson, 1878 H. lufarius Cresson, 1878 B. centralis Cresson, 1864 B. edwardsii Crcs.son, 1878 B. jlavifrom Cresson, 1863 B. grlscocollis ( De Geer, 1773) B. hunti Creene, 1860 B. niorri.wni Cresson, 1878 B. nevadcnsis ncvaden.si.f Cresson, 1874 B. occidentalis occidcntali.t Greene, 1858 B. nifocinctus Cres.son, 1863
B. Formicidae
1. Pogonoinipnicx
P. barhatus biirbatiis ( F. Smith, 18.56)
P. californiciis (Buckley. 1867)
P. imberbiculus W.M. Wheeler, 1902
P. occidentalis (Cresson, 1865)
P. rugosus Emerv, 1895
2. Crematngaater
C. mormonum Emery, 1895 C. vcrmiculata Emerv, 1895
3. Solcnopsis
S. rnolcsta mnlesta (Say, 1836) S. rnolcsta validitiscula Emery, 1895 S. salina W.M. Wheeler. 1908 S. xijloni McCixik, 1879
C. Mutillidae
1. Chi/photes
C. cpedaplnis Buzicky, 1941
C. simdis Baker, 190.5
2. Dasijmutilla
D. calif ornica ( Radoszkowski. 1861) D. caneo (Blake, 1879)
D. fulvohirta (Cresson, 1865)
D. gloriosa (Sau.ssure, 1867)
D. khigii (Gary, 1872)
D. nionticola (Cres.son, 1865)
D. phaon phaon (Fox, 1899)
D. phaon var. fimbrialis Miekel, 1928
D. scitula Miekel. 1928
D. Ursula (Cresson, 1875)
D. vesta vesta Cresson, 1865
3. Dilophotopsis
D. concolor cotuolor (Cresson, 1865)
4. Odontophotopsis
O. ercbus ( Melander, 1903)
O. in<-onspicua (Blake, 1886)
O. mclicau.M (Blake. 1871)
O. vcnusta (Blake, 1886)
5. Pscudomcthoca
P. contumax (Cresson. 1865)
P. conturncliosa Miekel, 1935
P. manca Miekel, 1924
P. projiinqua (Cres.son, 1865)
P. toumeiji (Fo.\, 1894)
6. Sphaeropthahna
S. abdomimdis ( Blaker, 1886)
S. ceres (Fox. 1899)
S. dircc (Fox, 1899)
S. marpesia (Blake, 1879)
S. unicolo (Cresson, 1865)
7. Timulhi
r. grotci (Blake, 1871)
Biological Series, Vol. 15, No. 2 Ut.mi Ahthhopods
13
8. Ttjphoctes
T. pccuUaris ( Cresson, 1875)
D. Ponipilidae Pepsis
P. ang^ustimarg^inata Viercck, 1908
P. mihiri Stdl, 1857
P. pallidoUmhata pallidolimlnitii Luca.s, 1895
P. thi.shc Lucas, 1895
E. Sphecidae
1 . Astatii
A. bicolor Say, 1823
A. ncvadicu Cres.son, 1881
A. nubecula Cresson, 1865
A. occidentalis Cre.s.son, 1881
2. Bcnd>ix
B. amcricana comata Parker, 1917
B. americatui spinolae Lepeletier, 1845 B. amoena Handlirsch, 1893 B. occidenttdis W.J. Fox, 1893
B. rugosa Parker, 1917
3. Cerceris
C. conifrons Mickel, 1916
C. coniergeus Viereck and Cockerel], 1904 C finitima Cresson, 1865 C. nigrcsccns Smith, 1856
4. Clypeadon
C. laticinctus (Cresson, 1865)
5. Didineis
D. nodosa Fo.\, 1894
6. Mimcsa
M. cressonii Packard, 1867
7. Philanthus
P. gibliosus (Fabricius, 1775)
8. Priomjz
P. atratus (Lepeletier, 1845)
P. parkeri Bohart and Menke, 1963
F. Vespidae
1. Anchtroccrus
A. antilopc antilopc ( Panzer, 1798 ) A. catskill albopluderatus (Saiissure, 185.5) A. catskill catskill (Saussure, 1853) A. lincativcntris fulvicarpus Cameron, 1908 A. ncocallosus ncocallosus Bequaert, 1943 A. spilogaster Cameron, 1905 A. tigris tigris (Saussure, 1857) A. tuhcrculiccps sutterianus (Saussure, 1875) A. tubcrculiccps tuberculiceps ( Saussure, 1853)
2. EnodijncTus
E. annulatus antiulatus (Say, 1824)
E. annulatus sulphureus (Saussure, 1858)
E. auranus (Cameron, 1906)
E. boscii boscii (Lepeletier, 1841)
E. cxoghjphus alhovittatus ( R. Bohart, 1939)
E. cxogli/phus cxoghjphus (R. Bohart, 1939)
E. joraminatus acqualus (Cameron. 1906)
E. fusus fusus (Cresson, 1872)
E. hidalgo hidalgo (Saussure, 1857)
E. martini ( R. Bohart, 1942)
E. praten-sis pratensis (Saussure, 1870)
£. russatus (R. Bohart, 1942)
3. Eumcnes
E. bollii holla Cresson, 1872
E. iturbide pedalis Fox, 1894
E. sculleni R. Bohart, 1950
E. verticalis tricinctus Isely, 1917
4. Leptochilus
L. eruhesccns ( H. Bohart, 1940)
L. republicanus ( Dalla Torre, 1889)
L. rubicundulus (R. Bohart, 1940)
L. rufinodus (Cresson, 1868)
5. Mischoci/tturus
M, jlavitarsis jliwitarsls (Saussure, 18.54) M. jlavitarsis idahocnsis Bequaert, 1933
6. Odijnerus
O. cinnaharinus R. Bohart, 1939 O. margarctcllus Rohwer, 1915
7. Polistes
P. canadensis var. kaibabcnsis Hayward,
19,32 P. flavus Cresson, 1868 P. fuscatus centralis Hayward, 1933 P. fuscatus utahensis Hayward, 19.33
8. Pseudotnasaris
P. cdwardsii (Cresson, 1872) P. zonalis (Cresson, 1864)
9. Pterocheilus
P. laticeps Cresson, 1872
P. micheneri R. Bohart, 1940
P. pediccllatus R. Bohart, 1940
P. provanchcri (Huard, 1895)
10. Stcnodtjncrus
S. apache R. Bohart, 1949
S. hlamloides blandoides R. Bohart, 1943
S. hlandus blandus (Saussure, 1870)
S. cochisensis (Viereck, 1908)
S. minimoferus R. Bohart, 1949
S. noticeps noticeps R. Boh,art, 1948
S. percampanulatus (Viereck, 1906)
S. toltccus (Saussure, 18.57)
S. valliceps R. Bohart, 1948
11. Symmorphus
S. meridionalis (Viereck, 1903)
12. Vespula
V. artica Rohwer, 1916
V. arenaria (Fabricius, 1775)
V. atropilosa (Sladen, 1918)
V. austriaca (Panzer, 1799)
V. consohrina ( Saussure, 1864 )
V. maculata (Linnaeus, 1763)
V. norvegicoides Sladen, 1918
V. pennsylvanica (Saussure, 1857)
V. vulgaris ( Linnaeus, 17.58 )
VII. Lepidoptera A. Arctiidae
1. Arachnis
A. picta Packard, 1864
2. Arctia
A. caja utahensis (Henry Edwards, 1886)
3. Apantesii
A. ncvadensis (Grote and Robinson, 1866)
A. ornata (Packard, 1864)
A. parthenice ( Kirby, 1837)
A. proxima ( Guerin-Meneville, 1844)
14
Brigham Young University Sgience Bulletin
A. williamsi tooele Barnes and McDun-
noiigli, 1910 A. willi(im.si form determinata ( Neumoegen,
1881)
4. Dkwrisia
D. vagans (Boisduval, 1852)
D. virginica (Fabricius. 1798)
5. Ectijpia
E. clio Jessica (Barnes, 1900)
6. Estigmcnc
E. oregonsis (Stretch, 1873)
7. Iliilijsidotu
II. iiTgentata sulmlpina French. 1890
//. maculata iigassizi Packard, 1864
/-/. oshiri Rothschild, 1909
II. tesi'llaris (J. E. Smith and Abbot, 1797)
8. Ilcrnilu/alcu
H. lahecula ( Grote, 1881)
9. Holomclinii
II. jnigilh (Streckcr, 1878)
10. hia
I. i.sdiiclla (J. E. Smith and Abbott, 1797)
11. Leptarctia
L. calijornkw form dccia (Boisduval, 1869)
12. Nemeophilii
N. plantaginis (Linnaeus, 1758)
B. Lasiocampidae
1. Malacosoma
M. americcimtm (Fabricius, 1793)
iW. adifornicum fragile (Stretch. 1881)
.\/. dis.^tria Hubner, 1822
2. PhijUode.sma
P. americana (Harris, 1841)
3. Tohjpe
T. glenivoodi Barnes, 1900
C. Lymantridae
Dasychira D. vagans grisea (Barnes and McDun- nough, 1913)
D. Nymphalidac
1 . Aghis
A. milherti (Jodart, 1819
2. Argijiitiis
A.' leto Behr, 1862
A. nokomis Edwards, 1862
3. Hasilarchia
B. Inrguini Boisduval, 1852
4. Ntjmphati.'y
N. antiopa (Linnaeus, 1758)
5. Vanessa
V . atahmta Linnaeus, 1758 v. cardui Linnaeus, 1758 V. canje Hubner, 1806
E. Saturniidae 1. Autonwris
A. 10 (Fabricius, 1775)
2. Coloradia
C. pandora Blake, 1863
3. Hemilcuca
H. eleganterina (Boisduval, 1852) H. hcra hera (Harris, 1841) H. nevadcn.'iis Stretch, 1872 H. olivine Cockerel], 1898
4. Platijsaniia
P. eurijalus (Boisduval, 1855) P. gtoveri Strecker, 1872
\111. Mallophaga
A. Gyropidae
Gliricota G. porcelli ( Schank, 1781) G. uvalis Burmeister, 1838
B. Laemobothriidae
Laeinuhothrion L. atrum ( Nitzsch, 1818) L. gliitinans Nitzsch. 1861 L. nuLximum (Scopoli, 1763) L. simile Kellogg, 1896 L. tinnttncidi (Linnaeus, 1758) L. vulturis (J. C. Fabricius, 1775)
C. Menoponidae
1. Actor7iithoplulus
A. laeii.'iiris Cla\, 1962
A. limarius Clay, 1962
A. lumino.sac (Kellogg, 1908)
A. mexicaniis Emerson, 1953
A. ochraeeus (Nitzsch, 1818)
A. paludosiis Clav, 1962
A. patelhtus (Piaget, 1890)
A. piceus lari (Packard. 1870)
A. piceus piceus (Denny, 1842)
A. stictus (Kellogg & Paine, 1911)
A, totani (Schrank, 1803)
A. tindirinus (Burmeister, 1838)
A. uniseriatuni ( Piaget, 1880 )
2. Atni/rsidea
A. inegalosoma (Overgaard, 1943) A. perdicis (Denny, 1842)
3. Ardeiphilus
A. floridae Tuff, 1965
4. Ausfroincnopon
A. aegialilidis ( Durrant, 1906)
A. atrofuhum (Piaget, 1880)
A. durisetosum ( Blagoveshtchenskv, 1948)
A. himantopi Timmermann, 1954
A. Iwu^sae Timmermaiui, 19.54
A. micrandum (Nitzsch, 1866)
A. saehtleheni Timmermann. 1954
A. spenceri Timmermann, 19.56
A. sijuatarolae Timmermann, 1954
A. transversum (Denny, 1842)
5. Bonomiclla
B. columhae Emerson, 1957
6. Ciconiphilus
C. butoridiphagus Carriker, 1964 C. cijgni Price & Beer, 1965
C. decimfaseiaius (Boisduval dsi Lacordaire,
1835) C. pectiniventris (Harrison, 1916)
7. Colpocephalum
C. hraclu/somum Kellogg & Chapman, 1902
Biological Series, Vol. 15, No. 2 Ut.'MI Ahthbopods
15
C. flavescens (do Haan, 1829)
C. fregili Denny, 1842
C. impressum Rudow, 1866
C. keUoggi Osboni, 1902
C. leptopygos Nitzsch, 1874
C. nanum Piaget, 1890
C. napiformc Rudow, 1869
C. pectinatum Oslwni, 1902
C. ifli/si (Ansari, 1951)
C. ttirhinatuin Denny, 1842
C. unciferum Kellogg, 1896
C. zerafae An.sari, 1955
8. Comatomenopon
C. thuhe Tuff. 1967
9. Cuculiphilus
C. altcrnatus ( Osbom. 1902)
10. Dennyus
D. bruneri (Carriker, 1903)
D. spiniger Ewing, 1930
11. Eureum
E. spenccri Emerson & Pratt, 1956
12. Hohorsticlla
H. frontalis Carriker, 1949
13. Holomenopopn
H. clypeilargum Eichler, 1943 H. Icucoxantluim ( Burmeister, 1838) H. setigerum ( Blagoveshtchensky, 1948) H. transvcialcmc (Bedford, 192())
14. Kurodiiia
K. acadicac Priee & Beer, 1963
K. fhimmci Price & Beer, 1963
K. fulvovasciata (Piaget, 1880)
K. haliueti (Denny, 1842)
K. magna Emerson, 1960
K. painci (McGregor, 1912)
K. subpachygastcr (Piaget, 1880)
15. Machaerilaemus
M. amercanus (Ewing, 1930) M. clayac ( Balat, 1966) M. malleus (Burmeister, 1838) M. melospizae Emerson, 1954
16. Menacanthtis
M. ahskcnsis ( Kellogg & Chapman, 1902)
M. annulatiis (Giebel, 1874)
M. chrysophacus (Kellogg, 1896)
M. distinctus ( Kellogg & Chapman, 1899)
A/, eurysternum ( BurmeLster, 1838 )
M. expansus (Osbom, 1896)
M . gonophaeus (Burmeister, 1838)
M. mutahilis Blagoveshtchenskv, 1940
M. perforatiiM (Piaget, 1880)
M. persignatus ( Kellogg & Chapman, 1899)
M. picicola (Packard, 1873)
M. Tohnstm (Kellogg, 1896)
.\/. siramineus (Nitzsch, 1818)
17. Mcnopon
M. pallens Clay, 1949
18. Myrsidea
M. anaspila (Nitzsch, 1866)
M. conspica (Kellogg & Chapman, 1902)
M. culcullans (Nitzsch, 1818)
.\/. dissimilis (Kellogg, 1896)
M. emersoni Clav. 1966
M. incerta (Kellogg, 1896)
M. mtcrrupta (Osbom, 1896)
M. Mifwris (Carriker. 1910)
M. mdanorum (Kellogg, 1896)
M. pallohs (Carriker, 1903)
A/, picae (Linnaeus, 1758)
M. quadrifasciata (Piaget, 1880)
M. quadrimaculata (Carriker, 1902)
M. ridulosa ( Kellogg & Chapman, 1899)
M. nistica (Giebel, 1874)
19. Nosopon
N. lucidum (Rudow 1869)
20. Piagetiella
P. peralis ( Leidy, 1878 )
21. Plegadiphilus
P. plegadis (Dubinin, 1938)
22. Pseudomenopon
P. insolens (Kellogg, 1896) P. par (Kellogg, 1896) P. pilosum (Scopoli, 1763) P. quadrii Eichler. 1952
23. Triniton
T. anserinum (J. C. Fabricius, 1805) T. querqucdulac (Linnaeus, 1758)
D. Philopteridae
1. Acidoproctiis
A. maximus Piaget, 1878
2. Anaticola
A. crassicornis corniccphalus ( Zavaleta,
1946) A. crassicornis crassicornis (Scopoli, 1763) A. crassicornis dajilensis Carriker, 1956 A. crassicornis dcpuratus (Nitzsch, 1866) A. crassicornis hopkinsi Eichler, 19.54 A. crassicornis mcrgiscrrati ( De Gear, 1778)
3. Anatoecus
A. cygni emersoni Keler, 1960
A. dentatus afjinis Keler, 1960
A. dentatus dentatus ( ScopoU, 1763)
A. dentatus ferrugineus (Giebel, 1874)
A. icterodcs bipunctatus (Giebel, 1874)
A. icterodes boschadis Keler, 1960
A. icterodes icterodes (Nitzsch, 1818)
A. icterodes marcui Keler, 1960
A. icterodes simrnillimus Keler, 1960
A. icterodes tcndeiroi Keler, 1960
4. Aquanirmus
A. amercanus (Kellogg & Chapman, 1899)
5. Ardeicola
A. botauri (Osbom, 1896)
A. cruscula Carriker, 1960
A. expallida Blagoveschtchenskv, 1940
A. jlorida nigra Tuff, 1967
A. goisagi Uchida, 1953
A. rhaphidus (Nitzsch, 1866)
6. Bruelia
B. angustifrons (Carriker, 1902) B. arguh (Burmeister, 1838)
B. audax (Kellogg, 1899)
B. biocellata (Piaget, 1880)
B. brachythorax (Giebel, 1874)
B. cedrorum (Piaget, 1880)
B. dcficiens (Piaget, 1885)
R. domestica (Kellogg & Chapman, 1899)
B. ductilis (Kellogg & Chapman, 1899)
B. iliaci brevicolor Ansari, 1956
B. interposita (Kellogg. 1899)
B. limbata (Burmeister, 1838)
B. Innga (Kellogg, 1896)
16
Brigham Young Univehsity Science Bulletin
B. kmgifrons Carriker. 1956
B. iichulosa (Burmeister, 1838)
B. ornalissinui ( Giebel, 1874)
B. pcninsularis (Kellogg, 1899)
B. mtundata (Osboni, 1896)
B. slmminea ( Dcnnv, 1842)
B. siihtilis (Nitzsch,' 1874)
B. icniih ( Bumifister, 1838)
B. xiinllioccplwli (Oshom, 1896)
B. zcrojnmctaiii antiqua Ansari, 1956
B. zcropunctata zcrofmnctata Ansari, 1957
7. Carduiceps
C. cinguhitus cinnidiitus (Denny, 1842) C. cingulatus chujiic Timmermann, 1954 C. zotuirius (Nitzsch. 1866)
8. Chelopistcs
C. mcleiigridis (Linnaeus, 1758)
9. CirrophthiTus
C. testudinarius (Children, 1836)
10. Cnlinicola
C. docophoroides ( Piaget, 1880)
11. Columbicola
C. baculoides Paine, 1912 C. macrourae ( Wilson, 1941 )
12. Craspcdorrhijnchus
C. americanus Emerson, 1960 C. acpiilinus ( Dennv, 1842) C, dihildttis (Rudow, 1869) C. luicmatnpus (Scopoli. 1763) C. liirsutus Carriker, 1956 C. subhaematopus Emerson, 1960
13. Cuclngastcr
C. Iictcroiinirnmicus (Nitzsch. 1866)
14. Cuculicola
C. splcndidus (Kellogg, 1899)
15. Cttculoecus
C. cocajgi (Oshom, 1895)
16. CummingsicUa
C. ambiffta (Burmeister, 1838)
C. longiro.itricola (Wilson, 1937)
17. Degeerhella
D. discocephalus aquilarum Eichler, 1943 D. fidva (Giehel, 1874)
D. fusca (Denny, 1842)
D. nistis nitus (Giehel, 1866)
D. nisuf: vagiins (Giebel, 1874)
D. r(>g(dis (Cliebel, 1866)
D. rufii carrtitlii Emerson, 1953
D. rufn rufa ( BumieLster, 1838)
18. Fulcolipcitrus
I'', margimdis Osbom, 1902 F. suhtralh (Rudow. 1869)
19. Fulicoffula
/•". amcTcana Emerson, 1960
F. comstocki (Kellogg & Paine, 1911)
F. distincta Emerson, 1960
F. longipda (Kellogg, 1896)
20. Coniocotes
G. chn/.mccplialtis Giebel, 1874 G. microlhorcix (.Stephans, 1829)
21. Goniodes
C. bonasus Emerson, 1948 G. ccntrocerci Simon, 1938 G. colchici Dennv. 1842
G. dispuT Burmeister, 1838 G. merriamanus Packard, 1873 G. nebra-ikensut Carriker. 1945 G. stefatii Clay & Hopkins, 1955 C. submamilUitus Emerson, 1950
Ihidoecus I. bisiiitiatus
Nitzsch, 1866)
23. IncidiJTons
I. moiuichus ( Kellogg & Paine, 1911) /. transpositus (Kellogg, 1896)
24 Lagopoecus
L. colcliicus Emerson, 1949
L. gambcli Emerson, 1949
L. gibsoui Hopkins, 1947
L. obscuriis Emerson, 1948
L. perplcxus ( Kellogg & Chapman. 1899)
L. umbellus Emerson, 1950
25. Lipeurus
L. maculosus Clay, 1938
26. Lunaceps
L. holophcicus C(il>enisi Timmermann, 1954 L. limoseUa clinjac Timmermann, 1954 L. numcnii (Denny, 1842)
27. MulcticoUi
M. macwccphalus (Kellogg, 1896)
28. Ornithohiiis
O. goniptcuriis Denny, 1842
O. wcitcrsloiii rccoitditus Timmermann, 1962
29. Oxylipvuru.s
O. corpulcnttis Clav, 1938
O. cllipticus (Keler, 1958)
O. mcsopclios colcliicus Clay, 1938
O. poh/trapczius (Burmeister, 1838)
30. Pcctinoptigus
P. farallonii (Kellogg, 1896)
P. tordoffi, Elbel & Emerson, 1956
31. Pcrcuirmus
P. (ircticus Cariker, 1958
P. auritUK (Scopoli, 1763)
P. gulosus (Nitzsch. 1866)
P. jtingcns (Kellogg, 1896)
P. mirinolatus ( Kellogg & Chapman, 1899)
P. (puidriiiiixluhitus ( Kellogg & Mann, 1912)
32. Philoptcrus
P. (igclaii (Osbom, 1896)
P. (imcriccmuti (Kellogg, 1899)
P. citrincUuc cunirostrac ( Schrank, 1776)
P. corvi ( Limiaeus, 1758)
P. excisus domcsticus (Kellogg, 1896)
P. excisus major (Kellogg, 1896)
P. <:'.v<;'isi/,s microsomaticus Tandan, 1955
P. fringillac (Scopoli, 1772)
P. gurruhic (Piaget, 1880)
P. tuniz^iki Balat. 1955
P. minis ( Kellogg & Chapman. 189^J)
P. occlhitus osborni Edwards, 1952
P. phillipi Emerson, 1953
P. picac (Dennv, 1842)
P. rufus (Kellogg, 1899)
P. ruttcri (Kellogg. 1899)
33. Phi/sconcUoidcs
P. spcnrcri Emerson & Ward, 1958
P. wiscmani Emerson, 1960
P. zcnaidurac (McGregor, 1917)
Biological Series, \ol. 15, No. 2 Utah Arthhopods
17
34. Picicola
P. foedus (Kellogg & Chapman, 1899) P. orphcus (Osbom, 1896) P. snodgrassi (Kellogg, 1896)
35. Quadraceps
Q. alcyonae (Carriker, 1959)
Q. assimilis major (Kellogg, 1899)
Q. canikcri Hopkins iSc Timmermann, 19.54
Q. cunncxus (Kellogg & Mann, 1912)
Q. charadrii hospcs ( Nitzsch, 1866)
Q. falcigcnts (Peters, 1931)
Q. fijnbriatus (Giebel, 1866)
Q. grisctis (Riulow, 1869)
Q. hcmichrous (Nitzsch, 1866)
Q. hiaticuhw Imcphilus (Kellogg, 1896)
Q nigrolimhatu.'i ( Mjoberg, 1910)
Q. phaeonotus (Nitzsch, 1866)
Q. punctatus suhlinguhitiis Timmerman,
1952 Q. mvus (Kellogg, 1899) Q. semifissus rncxicanus Carriker, 1944 Q. similis (Giebel, 1866) Q. zcplnjru (Timmermann, 1954)
36. Rallicola
R. advciuis (Kellogg, 1896)
fl. kelloggi Emerson, 1957
R. riHjsUix (Giebel, 1874)
R. ortygomctrac subporzcmac Emerson, 1957
37. Rlujnonirmus
R. scolopacis ( Denny, 1842 )
38. Rotundiceps
R. cordatiis (O.sbom. 1896)
39. Saemundssonia
S. conica conica (Denny, 1842)
S. conica naumanni (Giebel, 1874)
S. kratochvili Balat, 1950
S. lari congener ( Giebel, 1874 )
S. hbaticcps (Giebel, 1874)
S. pariigcnitalis Ward, 1955
S. plattjgastcr ni(^sc/ii ( Giebel, 1866)
S. platijgastcr phitygaster (Denny, 1842)
S. scolopacisphaeopodis (Schrank, 1803)
S. tricolor Carriker, 1956
S. tringae (O. Fabricius, 1780)
40. Strigiphilus
S. acutifrons Emerson, 1961
S. aitkcni Clay. 1966
S. harhatus (Osbom, 1902)
S. cursor ( Burmeister, 1838)
S. oculatus (Rudow, 1870)
S. otiis Emerson, 1955
S. speotyti (Osbom, 1896)
41. Sturnidoecus
S. simplex (Kellogg, 1896) S. sturni (Schrank, 1776)
E. Ricinidae 1. Ricinns
R. angulatus (Kellogg, 1896)
R. arcnatus (Kellogg & Mann, 1912)
R. homhi/cillac ( Dennv, 1842)
R. diffimis (Kellogg, 1896)
R. inexpeciatm Balat, 1966
R. japonicus ( Uchida. 1915)
R. mcdim Uchida, 1926
R. merulae ( Durrant, 1906)
R. microcephalus (Kellogg. 1896)
R. pictxiratus (Carriker, 1902) R. suhhaslatus (Durrant, 1906) R. sucinaceus (Kellogg, 1896) R. serratus (Durrant, 1906)
2. Trochiloecetes
T. lineatus (Osl)om, 1896)
T. prominens (Kellogg & Chapman, 1899)
T. ochoterenai (Zavaleta, 1943)
F. Trichodectidae
1. Bovicola
B. bovis (Linnaeus, 1758) B. caprae (Gurlt, 1843) B. craasipes (Rudow, 1866) B. equi (Denny, 1842) B. limbatus (Gervais, 1844)
B. ovis (Schrank. 1781)
2. Eutrichophilus
E. setosus (Giebel, 1861)
3. Felicola
F. subrostrata (Burmeister, 1838)
4. Geomydoecus
G. californicus (Chapman, 1897)
5. Neotrichodcctes
N. osborni Keler, 1944
1\. Siphonaptera
A. Amphipsyllidae
Amphisylki A. sibirica washingtoni Hubbard, 1954
B. Ceratophylhdae
1. Amphalius
A. necopinus (Jordan, 1925)
2. Ceratophyllus
C. affinis ncglectus Smit, 1958 C. celsus celsus Jordan, 1926 C. garei Rotlischild, 1902
C. nigcr C. Fo.x. 1908
C. petrochelidoni Wagner, 1936
3. DactylopsylUi
D. (Foxella) ignota apachina (C. Fo.x,
1941) D. (Foxella) ignota arizonensis (Hubbard.
1947) D. (Foxella) igtwta comis Jordan, 1929 D. ( Foxella ) ignota ignota B;iker, 1895 D. ( Foxella ) ignota rccula ( Jordan and
Rothschild, 1915) D. ( Foxella) ignota iitahensis (Wagner,
1931) D. (Foxclloidcs) minidoka Prince and Stark,
1951 D. (Spicata) rara I. Fo.x, 1940
4. Diamanus
D. montanus (Baker, 1895)
5. Malaraeus
M. bitterrootensis (Dunn, 1923) M.euphorbi ( Rotlvschild. 1905) M. xinomus (Jordan, 1925) M. telchimtm ( Rotkschild, 1905) M. vonfintelis Prince, 1959
6. Megabothris
M. abantis (Rotkschild, 1905)
7. Monopsyllu.t
M. ciliatus kincaidi Hubbard, 1947
18
Brigham Young UNivERSiTk- Science Bulletin
M. cyrturus (Jorilan, 1929)
M. eumolpi (irncriciinus Hubbard, 1950
M. eumulpi eumolpi (Rothschild, 1905)
M. exilis (Jordan. 1937)
M. vison (Baker, 1904)
A/, tvagneri (Baker, 1904)
8. NosojKyltus-
N. fo.icialus ( Bosc d'Aiitic, 1801 )
9. Opisocrosiis
O. hirsutus (Baker, 1895) O. Iiilm (Jordan and RothscMd, 1922) O. tubercuhitus cipwmuris Jellison, 1939 O. tuhcrculatu.s tiihcrctdatus ( Baker, 1904 )
10. 0/)i.so(/rt.vi/.v
O. kccii'i keeni (Baker, 1896) O. pseudarctomys (Baker, 1904)
1 1 . Orchopcu.s
O. caedens caedens ( Jordan, 1925 )
O. howardii (Baker. 1895)
O. Icucopus (Baker, 1904)
O. neotomac Augiiston, 1943
O. nepos ( Rotli.schild, 1905)
O. sexdcntiitus agdis ( Rotlischild. 1905)
O. sexdrntutus ncvndcnsis (Jordan, 1929)
1 2. Oropstilla
O. idahocmis (Baker. 1904)
13. Thrassis
T. acarmmtis medius Stark, 1970
T. acatjuwtis ulcdiensis (Wagner, 1936)
T. aridis campestris Prince, 1944
T. aridis hoffmcmi ( Hvihbard, 1949)
T. arizoncnsis ( Baker, 1898 )
T. hacclii hacchi (Rothschild, 1905)
r. bacchi caducus (Jordan, 1930)
T. hacchi con.similis Stark, 1957
T. hacchi gladioU.i (Jordan, 1925)
T. francisi harncsi (Stark, 1970)
T. francisi francisi ( C. Fox. 1927 )
T. pandorae pandorae Jellison, 1937
T.stanfordi (Wagner. 1936)
C. Hystrichtipsi/Uidue
1. Anomiopsi/lhis
A. ampliihohis Wagner, 1936 A. nudniiis (Baker, 1898)
2. AtypJdocfriis
A. echis cchis Jordan and Rothschild, 1915 A. multidcntatus mtdtidentatus (C. ¥ok, 1909)
3. Callisloim/Uus
C. tcrinus (Rotlischild, 1905)
4. Catallagia
C. dccipiens Rothschild, 1915
C. neweyi Holland and Loshbaiigh, 1958
5. Carteretta
C. carteri clavata C.ood. 1942
6. Conorliiuopsr/lla
C. standfardi Stewart, 19.30
7. Corrodopsijlhi
C. curvuta curvata (Rotlischild. 1915) C. curvata ohtusata (Wagner, 1929)
8. Clenophthahnus
C. pseiidtigi/rtes pseudagyrtes ( Baker, 1904 )
9. Delotelis
D. tehgoni Rothschild, 1905
10. Epitedia
E. scapani (Wagner, 1936) £. stanfordi Traub, 1944
E. testor (Rothschild, 1915)
E. wcnnuinni icen77janni (Rothschild, 1904)
11. IlyslrichopsyHti
II. dij)j)ici trnncata Holland, 19.57 //. hnsdidci Holland, 19.57
12. Jttrdanopsylla
}. allrcdi Traub and 'ripton, 1951
13. Megarthroglossus
M. becki Tipton and AUred, 1951 M. divisus divisus (Baker, 1898) M. procus Jordan and Rotlischild. 1915 M. smiti Mcndez, 1956
14. Meringis
M. dipodomys Kohls, 1938 M. huhhiirdi Kohls, 1938 A/, jamcsoni Hubbard, 1943 M. jewclli Hubbard, 1940 M. parkcri (Jordan, 1937)
15. Nearctopsylla
N. hrook'si (Rotlischild, 1904) N. hyrtaci (Rothschild, 1904)
16. Ncopsylla
N. inopina Rolkschild, 1915
17. Phalacropsyllti
P. alios \Vagner, 1936
18. Rhadinopsylla
R. hciscri (McCoy. 1911)
R. sectilis goodi (Hubbard, 1941)
R. sectilis sccldis (Jordan and Rothschild.
1923) R. fraterna (Baker, 1895)
19. Stcrmtomcra
S. alpina (Baker, 1895) ,S. huhlnirdi Egoscue, 1968 S. macroducUjla (Good, 1942)
D. Ischnopsyllidae
1. Myodopsylhi
it. gcntilis (Jordan and Rothschild, 1921)
2. Stcrnopsylhi
S. distincta lexana (C. Fox, 1914)
E. LeptosyUidae
1. Ctcnophyllus
C. armatus trrrihilis ( Rotkschild. 1903)
2. Odontopsyllus
O. dentatus (Baker, 1904)
3. Ornithophaga
O. nearctica Holland and Loshbaugh, 1958
4. Pcromyscopsylhi
P. hainifcr ligcns (Jordan, 1937) P. hcspcromys iidclpha (Rothschild. 1915) P. hespcromys ravallirn.sis (Dunn. 1923) P. sclcni-s- (Rothschild. 1906)
F. Puhcidae
1 . Ccdiopsylla
C. inaequalis inacqualis (Baker, 1895) C. intcrrupta Jordan, 1925
BiuLOGiCAL Sekies, \'()l. 15, No. 2 Utah Ahihhopods
19
2. Ctenocephalides
C. jclis fdis (Bouche, 1835)
3. Echidnophaga
E. gaUinacea ( Westwood, 1875)
4. lloplopsi/llus
H. (Euhoplopst/llus) ghicUdh iiffiiiis (Baker,
1895) H. (HopIo})syUus) anonudus (Baker, 1904)
5. Pulex
P. irritans Linnaeus, 1758
6. Xenopsi/lla
X. cheopis ( Rotlischild, 1903)
G. Vermipsyllidae Chaetopsylla C. stewarti Johnson, 1955
BIBLIOGRAPHY
ACARINA
Allred, D. M. 1954a. Mites found on mice of the genus Peromyscus in Utah. Unpuh. dissertation. Univ. of Utah, Salt Lake Cit>-, Utali. 163 p. + maps, graplis, ;md illustrations.
Allred, D. M. 19,54b. Mites ;w intermediate hosts of tapeworms. Pro*.-. Utiili Acad. Sci., Arts and Letters 31:44-51.
Allred, D. M. 1954c. Obser\ations on the stylosome (feeding tube) of some Utixli chiggers. Proc. Utiili Acad. Sci., Arts and Letters 32:61-63.
Allred, D. M. 1954d. Mites found on mice of the genus Peromi/scus in LUah. 1. General infestation. Great B;isin Nat. 16:23-31.
Allred, D. M. 1957a. Notes on the life history and bionomics of the wood rat mite, Brevisterna utah- ensUi (Aearina). Trans. Amer. Micro. See. 76:72-78.
Allred, D. M. 1957b. Mites found on mice of the genus Peronu/scus in LIt;ih. II. Family Haemogama- •sidae. Proc. Entomol. Soc. Wash. 59(l):31-39.
Allred, D. M. 1957c. Mites found on mice of the genus Peromyscus in Utah. 111. Family Dermanys- .sidae. Amer. Midland Nat. 57:450-460.
Allred, D. M. 1957d. Mites found on mice of the genus Peromyscus in Utah. V. Trombiculidae and miscellaneous families. Great B;isin Nat. 17:95-102.
Allred, D. M. 1957e. The male, deutonymph, and protonvmph of the mite Eubracht/laelaps circularis (Ewing) (.^carina: Laelaptidae) with notes on morphological variations. Ann. Entomol. Soc. Amer. .50:206-2TO.
Allred, D. M. 1957f. A new species of mite, Uirs- tiontissus hisetosus, from the nests of the desert wood rat, Ncotoma lepida lepida Thomas. Proc. Entomol. Soc. Wash. 59:83-89.
Allred, D. M. 1957g. Setal variations on mites of the .species Brevisterna utahensis (Ewing) (Aear- ina). Proc. Utah Acad. Sci., Arts and Letters 34: 51-54.
Allred. D. M. 1958. Mites found on mice of the genus Peromyscus in Utah. IV. Families Laelapti- dae and Phytoseiidae. Pan-Pacific Entomol. 34: 17-32.
Allred, D. M. 1961. Parasitic mites on marmots in Utah. J. Parasitol. 47:124.
Allred, D. M. 1965. Clarification of the type data for Ischyropoda fnrmani Keegan. J. Parasitol. 51: 604.
Allred, D. M. 1966. Unusual records of mites in Utah. Great Basin Nat. 26:34.
Allred, D. M., and D E. Beck. 1953a. Mite fauna of wood rat nests in Utah. Proc. Utah Acad. Sci., Arts and Letters 30:53-56.
Allred, D. M., and D E. Beck. 1953b. A new spe-
cies of Acomatacurus (Aearina, Trombicuhdae ) from Utah. Great Basin Nat. 13( 3-4 ) :87-90.
Allred, D. M., and D E. Beck. 1966. Mites of Utali mammals. Brigham Young Univ. Sci. BuU., Biol. Series 8(1):1-123.
Allred, D. M., and N. J. Marchett. 1957. Experi- mental feeding of the mite, Brevlstcrnu utahensis (Aearina: Haemogamasidae ) . J. Parasitol. 43:51-.54.
Allred, D. M., and E. ]. Roscoe. 1956. Life history of the tick Dermaccntor parumupertus in Utah. J. Parasitol. 42:516-522.
Allred, D. M., and E. J. Roscoe. 1957. Parasitic mites in desert wood rat nests, with notes on free living forms. Trans. Amer. Micro. Soc. 76:389-403.
Allred, D. M., D E. Beck, and L. D. White. 1960. Ticks of the genus Ixodes in Utah. Brigham Young Univ. Sci. Bull., Biol. Series l(4):l-42.
Ash, D. B. 1963. A .seasonal study of mites on deer mice (Peromyscus municukitus) in a chaparral community. L'npub. master's thesis, Brigham Young Univ., Provo, Utali. .33 p.
Bacha, W. J.. Jr. 1957. The life history of Otohius lagophdus. J. Panisitol. 43:560-565.
Beck, D E. 1955. Some unusual distributional rec- ords of ticks in Utah. J. Parasitol. 41:198-201.
Beck, D E., and D. M. Allred. 1955. Seasonal study of the tick Ornithodoros hermsi, found in the nest of the desert wood rat, Neotoma lepida lepida in Utah. Proc. Ut;ih Acad. Sci., .\rts and Letters (1954-19.55), 32:131-135.
BisHOPP, F. C. 1911. The distribution of the Rocky Mountain spotted fever tick. U. S. Dept. Agr. Bur. Ent. Circ. 136. 4 p.
BisHOPP, F. C., AND H. L. Trembly. 194.5. Distribu- tion and hosts of certain North American ticks. J. Para.sitol. 31(l):l-54.
Bhennan, J. M., AND D E. Beck. 195.5. The chig- gers of Utah (Aearina: Trombiculidae). Great Basin Nat. 15( 1-4) : 1-21.
Brinton, E. p. 1965. Identification of the adults, nymphs, and larvae of ticks of the genus Derma- ccntor Koch (I.xodidae) in the western United States. Unpub. master's thesis, Brigham Young Univ., Provo, Utah. 63 p.
Brinton, E. P., and D E. Beck. 1963a. Hard-bodied ticks of the western United States. Brigham Young Univ. Sci. Bull. Biol. Series 2(3) part 1:1-28.
Brinton, E. P., and D E. Beck. 196.3b. Hard-bodied ticks of the western United States. Brigham Young Univ. Sci. Bull., Biol. Series 2(3) parts 2 and 3:1-21.
Brinton, E. P., and G. M. Kohls. 1963. New dis- tributional and host data for the tick Dermacentor huntcri Bishopp. Great B:Lsin Nat. 23:166.
20
Bhigham Vounc University Science Bulletin
Bbinton, E. p., D E. Beck, and D. M. Allhed. 1965. Identification of the adults, nymplis, ;ind larvae of the genus Dcnnaccntor Koch ( Ixodidae ) in the western United States. Brigham Young Univ. Sci. Bull., Biol. Series 5(4): 1-44.
BnowN, J. H. 1945. The rabbit tick, Haemaplujsalis leporis-paluslri.s Packard, a,s an ectoparasite of man. Can. Entomol. 77:176.
CooLEY, R. A. 1938. The genera Dermacentor and Octocentor (l.\odidae) in the United States witli studies in variation. Nat. Inst. Health Bull. 171. 89 p.
CoOLEY, R. A. 1945. The genera Boophilus, Rhipi- cephalis, and Hacimipln/salis ( l.vodidae ) of the New World. Nat. Inst. Health Bull. 187. 54 p.
Cooley, R. a., and G. M. Kohls. 1944. The Arga- sidae of North America, Central America, and Cuba. Amer. Mid. Nat. Mono. 1. 151 p.
Cooley, R. A., and G. M. Kohls. 1945. The genus Ixodes in North America. Nat. Inst. Health Bull. 184. 246 p.
Davis, G. E. 1939. Omithodoros parkerii Distribu- tion and host data: spontaneous infection with relapsing fever spirochetes. Public Health Rpts. 54:. 345-349.
Despain, W. J. 1968. Elevational occurrence of the ticks Dermacentor (imlersoni and Dermacentor pa- rumapertus in Utah County, Utah. Unpub. master's thesis, Brigham Young Univ.. Provo, Utah. 36 p.
Edmunds, L. R. 1951. A checklist of the ticks of Utah. Pan-Pacific Entomol. 27:23-26.
Elzinga, R. J., and D. M. Rees. 1964. Abihty of immature ticks of Ixodes kingi and Dermacentor IHirumapertus to survive low temperatures. Proc. Utah Acad. Sci., Arts and Letters 41:213-216.
Fremling, C, and a. Gastfriend. 19.55. Seasonal abundance of the tick Dermacentor panimapcrtus. Ecology 36:162-163.
FuRMAN, D. p., AND V. J. TiPTON. 1955. The genus Myonyssus Tiraboschi (Acarina: Dermanyssidae ) including a new species from pika. J. Parasitol. 41:179-184.
Bastfriend, a. 19.55. New host records for immature stages of the tick Dermacentor paritmapertus. J. Parasitol. 41:63-65.
Herrin, C. S. 1970. A systematic revision of the genus Hirstionyssus (Acari: Mesostigmata ) of the Nearctic region. ]. Med. Entomol. 7:391-437.
Herrin, C. S., andD E. Beck. 1965. Observations on the biology, anatomy, and morphology of Oto- /him- lagophilus Coolev iuid Kohls. Brigham Young Univ. Sci. Bull., Biol. Series 6(2): 1-19.
HiccANs, H. G., and T. a. Wooley. 1962. A new species of Pcjssolozctes from Utah with notes on the genus (Acarina: Oribatidei). Great Basin Nat. 22:93-96.
Hooker, W. A. 1909. Geographical ihstribution of American ticks. J. Ec-on. Entomol. 2:403-428.
Howell, J. F., D. M. Allred, and D E. Beck. 19.57 Seasonal population fluctuations of mites in desert wood rat nests in central Ut;ili. Ecologv 38:82-88.
Howell, J. F., and R. J. Elzinga. 1962. A new Radfordm (Acarina: Myobiidae) from the kangaroo rat and a key to the known species. Ann. Entomol. Soc. Amer. 55:. 547-555.
Johnson, E. D. 1966. Ticks of the Dugway Proving Grounds and vicinity and their host associations. Proc. Utah Acad. Sci.'. Arts and Letters 43:49-66.
Jorcensen, C. D. 19.57a. Oviposition habits of the tick Dermacentor parumapertus Neumann and
factors influencing egg development. Great Basin Nat 17:42-51.
Jorcensen, C. D. 1957b. The Ord kanganm rat and its physical resistance to the tick Dermacentor parumapertus Neumann. Proc. Utah Acad. Sci., Arts and Letters 34:18.3-184 (Ab.stract).
Keegan, H. L. 19.53. Collections of parasitic mites from Utah. Great Basin Nat. 13:3.5-42.
Kohls, G. M. 19,52. A record of the occurrence of the tick Ixodes muris Bishopp and Smith on musk- rats in Utah. Great Basin Nat. 12:65-66.
Kohls, G. M., and C. M. Clifford. 1963. Omitho- doros sparnus n. sp., a parasite of wood rats, Neo- tovm spp. and deer mice, Peromyscus spp. in Utah and Arizona (Acarina: Arg;isidae). J. Parasitol. 49:857-861.
Kohls, G. M., and C. M. Clifford. 1968. The sub- genus Persicarf^as ( Ixodoidea, Argasidae, Argas). 6. A. (P.) giganteus, n. sp., from w ild birds in western United States and Sonora, Mexico. Ann. Entomol. Soc. Amer. 61:1113-1116.
Kohls, G. M., and H. Hoogstraal. 1960. Observa- tions on the subgenus Argas ( Ixodoidea, Argasidae, Argas ) . 2. A. cooleyi, new species, from western North American birds, Ann. Entomol. Soc. Amer. 53:62.5-631.
Kohls, G. M., and R. R. Parker. 1948. Occurrence of the brown dog tick in the western states. J. Econ. Entomol. 41:102.
Kohls, G. M., H. Hoogstraal, C. M. Clifford, and M. N. Kaiser. 1970. The subgenus Persicargas ( Ixodoidea, Argasidae, Argas ) . 9. Redescription and new world records of Argas ( P. ) persicus (Oken). and resurrection, redescription, and rec- ords of A. (P.) radiatus Raillie-t, A. (P.) sanchezi Duges, and A. (P.) minUitus Koch, new world ticks misidentified as A. (P.) persicus. Ann. En- tomol. Soc. Amer. 63:590-606.
Merino, J. M. 1967. Ticks of the genus Hacma- physalis in the western U, S. Unpub. master's thesis, Brigham Young Univ., Provo, Utidi. 116 p.
Parker, R. R., C. B. Philip, G. E. Davis, and R. A. Cooley. 1937. Ticks of the United States in relation to disease in man. y. Econ. Entomol. 30: 51-69.
Radovsky, F. J. 1967. The Macronyssidae and Lae- lapidae (Acarina: Mesostigmata) parasitic on bats. Univ. Calif., Pub. Entomol. 46:1-288. 48 pis.
Robertson, R. G. 1961. A morphological study of some larval ticks of the germs Dermacentor with emphasis on stnictures of possible taxonomic im- portance. LInpub. master's thesis, Brigham Young Univ., Provo, Utah. 30 p.
Rosasco, M. E. 1957. Seasonal abundance of the tick, Dermacentor panimapcrtus. on black-tailed jackrabbit, with notes on other ectoparasites. J. Mammal. 38:485-490.
Talley, G. M. 1957. The incidence of nasal mites in over-wintering red-wing blackbirds in the vi- cinity of Utiih Lake, Utah. Unpub. master's thesis, Brigham Young Univ.. Provo, Utah. 13 p.
White, L. D. 1959. Identification of hmal ticks of the genus Ixodes known to occur in Utah. Unpub. master's thesis, Brigham Young Univ., Provo, Utah. 64 p.
ANOPLURA
Bell, I. L. 1950. A study of sucking lice (Anoplura) on rodents in Utali. L'npub. master's thesis, Univ. of Utah, Salt Lake City, Utah. 55 p.
BiouiGicAL Series, Vol. 15, No. 2 Utah Arthropods
21
Emerson, K. C. 1970. Persomil communication.
IcNOFFO, C. M. 1956. Notes on lou.se-ho.st associa- tions of the Great Salt Lake Desert with keys to the hce. Great Basin Nat. 16( 1-4 ) :9-17.
IcNOFFO, C. M. 1957. New records of mammal-lice associations. Entomol. News 68:162.
Johnson, P. T. 1971. Personal communication.
ARANEIDA
Chamberlin, R. V. 1928. Notes on spiders from
the La Sal Mountains of Utali. Can. Entomol. 60:
93-95. Chamberlin, R. V., and W. Ivie. 1933. Spiders of
the Raft River Mountains of Utah. Bull. Univ.
Utah 23(4): 1-53. Chamberlin, R. V., and \V. Ivie. 1935. The hlack
widow spider and its varieties in the United States.
Bull. Univ. Utah Biol. Ser. 3(1): 1-29. Gertsch. \V. J. 1958. The spider genus Loxosceles
in Nortli America, Central America, and the West
Indies. Amer. Mus. Nov. 1977:1-46. Gertsch, W. J. 1970. Personal communication. L.\fferty, K. M. 1949. A prehminarv study of the
spiders of Red Butte Canyon. Unpub. master's
thesis, Univ. of Utah. Sdt Lake City, Utah. 152 p. Levi, H. \V. 1959. The spider genus Latrodectus
(Araneae: Thaeridiidae). Trans, Amer. Micro. Soc.
78:7-43. Levi, H. W., and A. Spielman. 1964. The biology
and control of the South American browTi spider,
Loxosceles laeta ( Nicolet ) , in a Nortli American
focus. Amer. J. Trop. Med. Hyg. 13:1.32-1.36.
COLEOPTERA
Arnet, R. H., Jr. 1960. The beetles of the United States. Cath. Univ. Press, Washington, D. C. 1112 p.
Gould, D. J., .and R. S. Real, Jr. 1952. New records of Leptinus testaccus from North America. Pan- Pacific Entomol. 28:193.
Parks, J. J., and J. W. B.'^.rnes. 1955. Notes on the fiunilv Leptinidae including a new record of Lep- tinilhis lalidus (Horn) in North America (Coleop- tera). Ann. Entomol. Soc. .\Tner. 48:417-421.
Selandeh, R. B. 1951. The bhster beetles ( Meloi- dae) of Utah. Unpnb. master's thesis, Univ. of Utah, Salt Lake City. Utiih. 140 p.
Selander, R. B. 1970. Personal communication.
Werner, F. G., and R. L. Edwards. 1948. Leptinus amcricanus Lee. taken on a shrew ( Coleoptera- Leptinidae), Psyche 55 (2):51-54.
Werner, F. C. W. R. Enns, and F. H. Parker. 1966. The Meloidae of Arizona. Univ. of Arizona Agr. Expt. Sta. Tech. Bull. 175. 96 p.
Wood, D. M. 1965. Studies on the beetles LeptinUlus validus ( Horn ) and Platiipst/Uus castoris Ritsema from beaver. Proc. Entomol. Soc. Ontario 95:33-63.
DIPTERA
Aldrich, J. NL 1905. A catalogue of North American Diptera. Smithsonian Misc. Collection 46:195-209.
.\rnell, J. H. 1968. The taxonomy, distribution, and biology of the Aedes varipalpis mosciuito complex in Utah. Unpub. master's thesis, Univ. of Utah, Salt Lake City. Utah. 73 p.
Arnell, J. H., AND L. T. Nielsen. 1967. Notes on the distribution and biology of tree hole mosquitoes in Ut;ih. Proc. Utah Mosq. Abate. Assoc. 20:28-29.
Arnold, F. D. 1966. An evaluation of the housefly problem in the major populated counties in Utah. Unpub. Ph.D. dissertation, Univ. of Utiih, Salt Lake City, Utah. 81 p.
Baird, C. R. 1970. Lal«>ratory studies of tvvo species of jackrabbit bot flies. Unpub. master's thesis, Utah State Univ., Logan, Utah. 40 p.
Baird, C. R., and K. J. Capelle. 1969. Successful lalxjratorv mating of two species of jackrabbit bot flies. J. Med. Entomol. 6:196.
Beadle, L. D. 19.55. Man-biting habits of Culex tarsalis and associated mosquitoes in northern Utah. Proc. Utah Mosq. Abate. As,soc. 8:3.
Beadle, L. D. 19.59. Field observations on the biting habits of Culex tarsalis at Mitchell, Nebraska, and Logan, Utah. Amer. J. Trop. Med. Hyg. 8:134-140.
Beck, D E. 1960. Mostjuito survey of central Utah Vallev, Utah. Proc. Utah Mosq. Abate. Assoc. 13: 18-23.
Beck, D E. 1961. Central Utah County, Utah, mos- quito survey studies. Mosq. News 21:6-11.
Bennett, G. F., and C. W. Sabrosky. 1962. The Nearctic species of the genu.s Cephcncmyia ( Dip- tera, Oestridae). Can. Zool. 40:431-448.
Bequaert, J. C. 19.54-19.57. The Hippoboscidae or louse-fhes (Diptera) of mammiils and birds. Part II. Taxonomy, evolution and revision of American genera and species. Entomol. Amer. 24-26:1-611.
Bullock, H. R. 1952, Cuhcoid gnats of Salt Lake County. Unpub. master's thesis. Univ. of Utah, Salt Lake City, Utah. 69 p.
Capelle, K. J. 1968. Seasonal incidence and species composition of blowflies at Bear River, Utah, 1963- 1965. Paper presented at Entomol. Soc. Amer, Meetings, Dallas, Texas, Dec, 2, 1968. Unpublished.
Capelle, K, J. 1970. Studies on the life history and development of Cuterebra polita (Diptera: Cuter- bridae) in four species of rodents. J. Med. Ento- mol. 7:320-327.
Capelle, K. J. 1970. Personal communication.
Carpenter, M. J. 1965. A study of ovarian cycles and longevity in univoltine and multivoltine species of Rocky Mountain Aedes mosquitoes. Unpub. master's thesis, Univ. of Utah, Salt Lake City, Utah. 53 p.
CoLLETT, G. C, and D. M. Rees. 1957. A com- parison of mosquito collections made with portable (C02) bait traps and New Jersey type light traps. Proc. Utah Mosq. Abate. Assoc. 10:6-7.
CoLi.ETT, G. C.. AND D. M. Rees. 1959. Mosquito abatement programs in Utah in 19.58. Proc. 46th Ann, Meeting New Jersey Mosf], Exterm. Assoc. p. 61-63.
Collett, G. C, and D. M. Rees. 1961. Recent changes in Aedes niammaculk (Ludlow) popula- tions in the Salt Lake City Mostpiito Abatement District. Proc. and Papers 29th Ann. Conf. Cailf. Mosq. Control Assoc, p. 115.
Defoliart, G. R., and B. V. Peterson. 1960. New North American Simuliidae of the genus Cnephia Enderlein (Diptera), Ann. Entomol. Soc. Amer. 53:213-219.
Elbel, R. E., G. T. Crane, L. E. Stipe, G. B. Van Nosdol, and K. L. Smart. 1969. Mostpiitoes and arbovinises from Callo, Utah, for the years 1966 and 1967 (Abstract). Proc, Utah Mosq, Abate. Assoc. 22:.39.
Bricham Young University Science Bulletin
Graham, C. L. 1962. Cuterbru of Ut^ili and neigh- boring; states. Unpiib. master's thesis, Utah State Univ., Logan, Utah. 35 p.
Graham, C. L. 1964. Biological studies of three species of Cutehra from nortlieni Utah. Unpub. Ph.D. dissertation, Utali State Univ., Logan, Utah. 82 p.
Graham, J. E. 1950. A preliminarv stud)' of the gnats or midges of the tribe Tendipedini in north- em Utah. Unpiil). master's thesis, Univ. of Utah, Salt Lake City, Utah. 48 p.
Graham, J. E. 1957. The need for a professional basis for pest control work in Utah. Proc. Utah Acad. Sci., Arts and Letters 34:31-33.
Graham, J. E., and D. M. Rees. 1955. Granular insecticide carriers used in Utah in Moscjuito abatement operations. Proc. and Papers 23rd Ann. Conf. Calif. Mosc). Control As.soc. p. 106-108.
Graham, y. E., and D. M. Rees. 1956. Progress re- port on the field testing in L'tah of .some organic phosphorus insecticides for mosquito control. Proc. Utah Mos(i. Abate. Assoc. 9:23-25.
Graham, J. E., and D. M. Rees. 1958. Chemical control of mosquitoes in Utah. Proc. and Papers 26tli Ann. Conf. Calif. Mosq. Control Assoc, p. 16-19.
Graham, J. E., D. M. Rees, and G. F. Edmunds. 1953. A season of mosquito control with hepta- chlor. Proc. and Papers 22nd Ann. Conf. Calif. Mosq. Control Asscx:. p. 21.
Graham, J. E., D. M. Rees, and L. T. Nielsen. 1958. Trends in mosquito populations in Salt Lake County. Mosq. News 18:98-100.
Graham, J. E., and I. E. Bradley. 1962a. The ef- fects of species on density of mosquito larval popu- lations in Salt Lake Count\' Utah. Mosq. News 22:239-247.
Graham, J. E. and 1. E. Bradley. 1962b. An evaluation of some techniques used to measure mosquito populations. LIt:ili Acad. Sci., Arts and Letters .39:77-83.
Grundmann, a. W. 1949. Mosquitoes as pests and vectors of disease in Utah. Proc. Utah Mosq. Abate. Assoc. 2:58-63.
Ghundmann, a. W., and J. M, Butler. 1953. Dip- terous lar\ae infesting tlie intestinal tract of the covote, CaiiK latrans Sav 1823. J. Kansas Entomol. Soe. 26:129-131.
Grundmann, A. W., and ]. R. Crook. 19.59. Endo- parasites of rodents found in Glen Can von. Univ. Utah Anthropol. Papers No. 40:105-106.
Harmstron, F. C, and A. D. Hess. 1964. Mosfjui- toes as actual and potential vectors of disease as now known for Utah. Proc. L'tah Mosq. Abate. A.ssoc. 17:2.5-27.
HoPLA, C. E. 1947. A ta.\onomic sttidy of the adult mosquitoes of Utah County, with notes on the biology and distribution of the more common species. Unpub. master's thesis, Brigham Young Univ., Provo, Utah. 97 p.
loNOFFo, C. M. 1961. Biology of Cutcrchra jclliwni on Lcpus caUfornicus deserticoki. Ann. Entoninl, Soc. Amer. .54:,509-512.
James, M. T. 1948. The flies that cause mviasis in man. U. S. Dcpt. Agr. Publ. 631. 175 p. ( 1947).
JoBLiNG, B. 1938. A revision of the genus Trirhohius Diptera, Acalvpterae, Streblidae). Parasitology 30:3.58-587.
Jone-S, D. T. 1941-1943. Utah house flies. Proc. Utah Acad. Sci., Arts and Letters 18-20:10.5-108.
Knowlton, G. F. 1931. Notes on Ut;ili Diptera. Can. Entomol. 63:1,52-157.
Knowlton, G. F. 1936. Some Utah flesh flies. Proc. Utah Acad. Sci., Arts and Letters 13:237-239.
Knowlton, G. F., and L. E. Fronk. 1950. Some blood-sucking Diotera of Utah. Utah Agr. Expt. Sta. Mimeo. Ser. 369.
Knowlton, G. F., and F. C. Harmston. 1937. Utah Diptera. Proc. Utah Acad. Sci., Arts and Letters 14:141-149.
Knowlton, G. F., and D. E. Madsen. 1935a. The sheep "tick." Ut;\h Agr. Expt. Sta. Leaflet 67.
Knowlton, G. F., and D. E. Madsen. 193.5b. Mos- quitoes annoy sheep. ]. Ecxin. Entomol. 28:934-935.
Knowlton, G. F., and D. R. Maddock. 1944. Snipe flies in Utah. J. Econ. Entomol. 37:119.
Kn(jwlton. G. F., and J. A. Rowe. 19.34. New blood-sucking flies from Utah ( Simuliidae, Dip- tera). Ann, Entomol, Soc. Amer. 27:.580-584.
Knowlton, G. F., and T. O. Thatcher. 1934. Utah horseflies. Proc. Utah Acad. Sci., Arts and Letters 11:291-294.
Knowlton, G. F., F. C. Harm.ston, and D. E. Hard- ing. 1938. Blood-sucking Utah Diptera. Proc. Utah Acad. Sci., Arts and Letters, 15:103-105.
Knowlton, G. F., F. C. Harmston, and G. S. Stains. 1939. Insects of Ut;ih, Diptera. Utah Agr. Expt. Sta. Mono. Ser. 200 (Tech) part 5. 22 p.
Knowlton, G. F., M. }. Janes, and W. L. Thomas. 1933. Some Utah Diptera. Proc. Utah Acad. Sci., Arts and Letters 10:155-157.
Knudsen, a. B., and D. M. Rees. 1967. The emerg- ence of Chnjsops discalis WiUiston on the shore of the Great Salt Lake, Utah. Proc. Utah Mosq. Abate. Assoc. 20:29-33.
Lewis, W. 1949. The taxonomy and notes on bio- nomics and control of the Tabanidae of Salt Lake Countv, Utah. Unpub. master's thesis, Univ. of Utah. Salt Lake City. Utah, 81 p.
Linam, |, H. 1961, A mosquito sur\ey of Skull Vallev, Tooele County, Utah. Proc. Utah Mosq. Abate. Assoc. 14:26-27.
Linam, J. H., and L. T. Nielsen. 1962. Notes on the taxonomic separation of adult females of Culex pipiens L. and Culex quinquejasciatus Sav. Mosq. News 22:290-293.
Linam, J. H,, and L. T. Nielsen. 1963a. L'tah mos- quitoes— their publi.shed lustory, Proc, Utah Mosq, Abate. Assoc. 16:16-23.
Linam, J. H., and L. T, Nielsen. 1963b. Notes on the identification of some western Culex larvae. Proc. .50th .i^nn. Meeting New Jersev Mosci, Exterm, Assoc, p, 411-415,
Linam, J, H,, and L, T, Nielsen. 1964. Utah mos- quitoes— their published histor\', Suppkment 1. Proc. Utah Mosq. Abate. Assoc'. 17:22-23.
Linam, J. H., and L. T. Nielsen. 1966. Notes on the distribution, ecology, and overwintering habits of Ci//ex pipiens Adams in Utali, Proc. Entomol. Soc, Wash, 68(2): 1.36-1.38,
Mahlor, R, L,, and D, M, Rees, 1957, The influence of climatological factors on the biting activities of the mosquito Ac(/c,s dorsalus (.Meigen), Proc. Utah Mosq. Abate. Assoc. 10:19-22.
MiKHERjEE, A. B. 1964. A comparative study of the karyotypes of four genera and nineteen species of mosquitoes present in Utah. Unpub. master's thesis, Univ. of Utah, Salt Lake City, Utah. .52 p.
Biological Series, Vol. 15, No. 2 Utah Arthropods
23
Neilsen, L. T. 1957. Notes on tlie flight nmges of Rocky Mountain moscjiiitoes of the genus Aecles. Proc.'Utah Acad. Sci., Arts and Letters 34:27-29.
Nielsen, L. T. 1958. Control of snow mosquitoes in the mountains of Utah. Proc. Utah Mosq. Abate. Assoc. 11:33-34.
Nielsen, L. T. 1959a. The mosquitoes of Utah — a revised list. Mosq. News 19:45-47.
Nielsen, L. T. 1959b. Seasonal distribution and longevit)' of Rocky Mountain snow mosquitoes of the genus Aedes. Proc. Utah Acad. Sci., Arts imd Letters 36:83-87.
Neilsen L. T. 1962. The Utah Mosquito Abate- ment Association — a brief history of mosquito con- trol in Utah. Proc. Utah Mosq. Abate Assoc. 15: 12-15.
Nielsen, L. T. 1968. A current list of mosquitoes known to occur in Utali with report of new rec- ords. Proc. Utali Mosq. Abate. Assoc. 21:34-37.
Nielsen, L. T., and J. H. Linam. 1963. New dis- tributional records of the mosquitoes of Utah. Proc. Utah Acad. Sci., Arts and Letters 40:19.3-196.
Nielsen, L. T., and ]. H. Linam. 1964, Additional distribution records for Utah mos<]uitoes with notes on biology. Proc. Utah Mosq. Abate. Assoc. 17:29-31.
Nielsen, L. T., and J. H. Linam. 1966. Utah mos- quitoes— tlieir published history: Supplement II. Proc. Utah Mosq. Abate. Assoc. 19:21-23.
Nielsen, L. T., and D. M. Rees. 1961. An identi- fication guide to the mosquitoes of Utah. Bull. Univ. Utah Biol. Series 12(3): 1-58.
Nielsen, L, T., J. H. Arnell, and J. H. Linam. 1967. A report on the distribution and biology of treehole mo.stiuitoes in the western United States. Proc. 35th Ann, Conf, Calif. Mos(]. Cont. As.soc, Inc and Amer, Mosq, Cont, Assoc, p. 72-76,
Nielsen, L. T„ J, H, Linam, and D, M, Rees, 1963, New distributional records for mosquitoes in the Rocky Mountain States. Proc, 50th Ann, Meeting New Jersey Mosq, Exterm. Assoc, p, 424-428,
Nielsen, L, T., ]. H, Linam, J, H, Arnell, and T, J. Zavortink, 1968. Distributional and biological notes on the treehole mosquitoes of the western United States. Mosq. News 28:.361-365.
OcDEN, L, J„ AND J, W. KiLPATRiCK, 1958, Control of Fannia catiicularK ( L, ) in Utah dairy bams, J. Econ. Ent. 51:611-612.
Peterson, B, V, 1953, The black flies (Diptera: Simuliidae) in the canyons in the vicinity of Salt Lake City with notes on their biology. Unpub. master's thesis, Univ. of Utah, Salt Lake City, Utah. 67 p.
Peterson, B. V. 19.55. A preliminary list of the black flies (Diptera: Simuliidae) of Utah. Proc. Utah Acad. Sci., Arts and Letters 32:113-115.
Peterson, B. V. 19.56. Observations on the biology of Utah black flies (Diptera: Simuliidae). Can. Entomol. 88:496-507.
Peterson, B, \'. 1958a. The taxonomy and biology of Utah species of black flies (Diptera: Simuli- idae). Unpub. Ph.D. dissertation, Univ. of LTtah, Salt Lake City, Utah. 336 p,
Peterson, B, V, 1958b, A redescription of the fe- male and first description of the male, pupa, and larva of Prosimnlium flaviantennus (S, and K, ) with notes on the biologv and distribution. Can. Entomol. 90:469-473.
Peterson, B, \', 19.59a, New distributional and host records for bat fhes and a key to the North Am-
erican species of Basilui Hibciro (Diptera: Nycteri- biidae). Proc, Entomol, Soc, Ontario 90:30-37.
Peterson, B. V. 1959b. Three new black fly records from Utah (Diptera: Simuliidae). Proc. Entomol. Soc. Wa.sh. 61:21.
Peterson, B. V. 1959c. Notes on the biology of some species of Utah black flies (Diptera: Simuhidae). Mosq. News 19:86-90,
Peterson, B, V. 1959d. Observations on mating, feeding, and oviposition of some Utah species of black flies (Diptera: Simuhidae), Can, Entomol. 91:147-155.
Peterson, B. V. 1960a. Notes on .some natural ene- mies of Utah black fhes (Diptera: Simuhidae). Can. Entomol. 92:266-274.
Peterson, B. V. 1960b. The Simuliidae (Diptera) of Utah, Part I. Keys, original citations, types and distribution. Great Basin Nat. 20:81-104.
Peterson, B. V, 1970a. The Prosimnlium of Canada and Alaska (Diptera: Simuliidae). Mem. Entomol. Soc. Can. No. 69:1-216,
Peterson, B, V, 1970b, The identities of three closely related western species of Prosimulium (Diptera: Simuliidae). C;m. Entomol. 102:118-128.
Peterson, B, V., and G. R, Defoliart, 1960, Four new species of Prosimulium (Diptera: Simuliidae) from western United States, Can, Entomol. 92: 85-102.
Rees, D. M. 1934a. Mosquito records from Utah, Pan-Pacific Entomol, 10:161-165,
Rees, D. M, 1934b. Notes on mosquito fish in Utah, Gamhusia afjinis ( Baird and Girard). Copeia 1934 (4):157-159.
Rees, D. M. 1935. Observarions on a mosquito flight in Sak Lake City. Bull. Univ. Utah 25(5), Biol. Series 2(6): 106.
Rees, D. M, 1939a, Origin of mosquito producing waters in the vicinitv of Salt Lake City, Utah, Bull, Univ. Utah 29(5), Biol. Series 3(9): 1-13.
Rees, D. M. 1939b. Progress in mosquito control in the vicinity of Salt Lake City. Proc. 26th Ann. Meeting New Jersey Mosq. E.\term. Assoc. 155-160.
Rees, D, M, 1942a, The mosquitoes of Utah. Bull. Univ, Utah Biol, Series .33(7):l-99,
Rees, D, M. 1942b, Supplementary list of mosqui- toes from Utah, Pan-Pacific Entomol, 18:77-82,
Rees D. M. 1944. A new mosquito record from Utah (Diptera: Culicidae). Pan-Pacific Entomol. 20:19.
Rees, D, M, 1945a, The utilization of fish by a mosquito abatement district; their effectiveness and limitation, Proc, 32nd Ann, Meeting New Jersey Mosq, E.\term, Assoc, p, 211-216,
Rees, D, M, 1945b, Supplemental notes on mosquito fish in Utah, Gamhusia affinis (Baird and Girard). Copeia 1945(4):236.
Rees, D. M, 1949, Advances in mosquito work in Utah in 1948, Proc. 36th Ann. Meeting New Jersey Mosq. Exterm. Assoc, p. 58-59,
Rees, D, M. 1952, The effectiveness of mosquito control in Salt Lake City, Utah, as determined by light trap collections. Proc. 39th Ann. Meeting New Jersey Mosq. Exterm. Assoc, p. 105-109.
Rees, D. M, 1954, The Acdes nigromticulis mosquito situation in Utah, Proc, Utah Mosq, Abate, A,s.soc. 7:23-24.
Rees, D. M. 1955. Recent advances in mosquito control in the Rocky Mountain region. Proc. and Papers 23rd Ann. Conf. Calif. Mosq. Control As- .soc. and 11th Ann. Meeting Amer. Mosq. Control Assoc, p. 6-9.
24
Bricham Young University Science Bulletin
Rees, D. M. 1957. Cooperative measures applied in mosquito abatement in Utah during 19.56. Proc. and Papers 25th Ann. Conf. Calif. Mosq. Control Assoc, p. 15-17.
Rees, D. M. 1958. Report on the results of control methods applied to biting gnats in the vicinity of Salt Lake Cit\', Utah, (Diptera: Ceratopogoni- dae). Proc. 10th intern. Congr. Entomol. 3:741-744.
Rees, D. M. 1964. Mo.squitoe.s — importance in Utah. Utah Water News 12(3): 14.
Rees, D. M., and F. D. Ahnolo. 1953. An outbreak of blowflies in Salt Lake County in 1962. Proc. Utah Acad. Sci., Arts and Letters 30:60-62.
Rees, D. M., and H. R. Bullock. 1954. Culicoid gnats of Salt Lake County, Utah (Diptera: Cera- topogonidae ( = Heleidae) ). Proc. Utah Acad. Sci., Arts and Letters 31:42-43.
Rees, D. M., and G. C. Collett. 1954. The biology of Aedes niphadopsis Dyar and Knab (Diptera: Cuhcidae). Proc. Entomol. Soc. Wash. 56:207-214.
Rees, D. M., and C. C. Collett. 1955. An air- propelled boat used in mosquito abatement work in Utah. Mosq. News 15:193-195.
Rees, D. M., and G. C. Collett. 1956. Mosquito abatement activities in Utah during 1955. Proc. and Papers 24th Ann. Conf. Calif. Mosq. Control Assoc, p. 6-8.
Rees, D. M.. and G. C. Collett. 1959. Factors influencing the encephalitis outbreak in Utah in 1958. Proc. and Papers 27th Ann. Conf. Calif. Mosq. Control Assoc, p. 88-97.
Rees, D. M., and J. E. Graham. 1953. 1952 field tests of heptachlor for mosquito control. Proc. and Papers 21st Ann. Conf. Calif. Mosq. Control As- soc, p. 4-6.
Rees, D. M., and F. C. Harmston. 1946. Observa- tions on the habits of Anopheles freeborni in north- em Utah and southern Idaho (Diptera: Cuhcidae). Mosq. News 6:73-75.
Rees, D. M., and L. T. Nielsen. 1947. On the biology and control of Aedes dorsalis { Meig. ) in Utah. Proc. 34th Ann. Meeting New Jersey Mosq. E.xterm. Assoc, p. 160-165.
Rees, D. M.. and L. T. Nielsen. 1951. Four new mosquito records from Utah. Pan-Pacific Entomol. 27:11-12.
Rees, D. M., and L. T. Nielsen. 1955. Additional mosciuito records from Utah. Pan-Pacific Entomol. 31:31-33.
Rees, D. M.. and K. Onishi. 1951. Morphology of the terminalia and internal reproductive organs, and copulation in the mosquito, CuUseta inornata (Williston) (Diptera: Culicidae). Proc. Entomol. Soc. Wa.sh. 53:233-246.
Rees, D. M., and B. V. Peterson. 1953. The black flies (Diptera: Simuhidae) in the canyons near Salt Lake City, Utah. Proc. Utah Acad. Sci., Arts and Letters 30:57-59.
Rees, D. M., and J. V. Smith. 1950. Effective con- trol methods used on biting gnats in Utah during 1949 (Diptera: Ceratopogonidae ) . Mosq. News 10:9-15.
Rees D. M., and J. V. Smith. 1952. Control of biting gnats in North Salt Lake City, Utah, (Dip- tera: Heleidae). Mosq. News 12:49-52.
Rees, D. M., G. C. Collett, and R. N. Winget. 1969. The Leptoconops problem in Utah. Proc. Utah .Mosq. Abate. Assoc. 22:16-18.
Rees, D. M., G. F. Edmunds, and L. T. Nielsen. 1953. Additional uses of granular larvicides in
mosquito abatement. Proc. and Papers 22nd Ann. Conf. Calif. Mosq. Control Assoc, p. 20-21.
Richards, C. S., L. T. Nielsen, and D. M. Rees. 1956. Mosquito records from the Great Basin and the drainage of the lower Colorado River. Mosq. News 16:10-17.
Richens, V. B. 1965. Larvae of bot fUes in the northern pocket gopher. J. Mammal. 46:689-690.
Rowe, J. A. 1953. Mosquitoes and health. Proc. Utah Moscj. Abate. Assoc. 6:12-14 ( Mimeo).
Rowe, J. A., and G. F. Knowlton. 1935. The genus Tahamts in Utah. Can. Entomol. 67:238-244.
Rowe, J. A., and G. F. Knowlton. 1936. Pangoni- inae of Utah (Tabanidae: Diptera). Ohio J. Sci. 36:253-259.
Sabrosky, C. W. 1941. The Hippelates flies or eye gnats: Prehminary notes. Can. Entomol. 73:23-27.
Sedman, Y. S. 1952; The Syqjhidae of Utah. Unpub. master's thesis, Univ. of Utah, Salt Lake City, Utah. 174 p.
Smith, J. V. 1950. A study of the control of the gnat, Leptoconops kerteszi var. americanus Carter in North Salt Lake Citv. Utah. Unpub. master's thesis, Univ. of Utah, Salt' Lake City, Utah. 51 p.
Stains, G. S.. and G. F. Knowlton. 1940. Three new western Simuliidae (Diptera). Ann. Entomol. Soc. Amer. 33:77-80.
Stains, G. S., and G. F. Knowlton. 1943. A ta.xo- nomic and distributional study of the Simuhidae of the western United States. Ann. Entomol. Soc. Amer. 36:259-280.
Stone, A., and B. V. Peterson. 1958. Simulium defoliarti, a new black fly from the western United States (Diptera: Simuliidae). Bull. Brooklyn En- tomol. Soc. 53:1-6.
Stone, A., C. W. Sabrosky, W. W. Wirth, R. H. Foote, and J. R. Coulson. 1965. A catalog of the Diptera of America north of Mexico. U. S. Dept. Agr. Handbook 276. 1696 p.
Taylor, S. K. 1959. Bionomics of Culex tarsalis Coquillett in Ut;ih County. Unpub. master's thesis, Brigham Young Univ., Provo, Utah. 36 p.
Trent, D. W. 1960. Observations on the hiberna- tion of Ctdex tarscdis Coquillett in Utah Vallev. Unpub. master's thesis, Brigham Young Univ., Provo, Utah. 30 p.
TwiNN, C. R. 1938. Blackfhes from Utah and Idaho, with descriptions of new .species ( Simuhidae, Dip- tera). Can. Entomol. 70:48-55.
Wirth, W. W., and F. S. Blanton. 1967. The North American Ctilicoides of the guttipcnnis group (Diptera; Ceratopogonidae). Florida Ento- mol. .50:207-243.
Wirth, W. W., and F. S. Blanton. 1969a. North American Culicoides of the ptdicaris group ( Dip- tera: Ceratopogonidae). Florida Entomol 52: 207-243.
WwTH, W. W., and F. S. Blanton. 1969b. New species and records of Culicoides from western North America. (Diptera: Ceratopogonidae). Proc. Entomol. Soc. Wash. 71:556-.567.
HEMIPTERA
Knowlton, G. F. 1955. Hemiptera of Utah — records.
Utah Agr. Expt. Sta. Mimeo. Series 140. Knowlton, G. F., and E. J. Taylor. 1&48. Some
predaceous Hemiptera largely from Utah. Proc.
Utah Acad. Sci., Arts and Letters 26:135.
Biological Series, Vol. 15, No, 2 Utah Arthropods
25
Nielsen, G. L. 1955. A study of the familie.s of Utali Hemiptera. Unpub. ma.ster's thesi.s, Brigham Young Univ., Provo. Utiih. 46 p.
Ryckman, R. E. 1954. Reduvius senilus Van Duzee from the lodge.s of Neotoma in San Juan County, Utah. Bull. So, Calif. Acad. Sci. 53(2):88.
UsiNCEH, R. L. 1944. The Triatominae of North and Central America and the West Indies and their pubhc he;ilth significance. U. S. Pubhc Health Ser. Public Health Bull. 283, 83 p.
UsiNCER, R. L. 1966. Monograph of the Cimicidae ( Hemiptera-Heteroptera ) Thomas Say Foundation Pub. 7. 585 p. + xi.
WVCODZINSKY, P., AND R. L. UsiNCER. 1964. The
genus Reduvius Fabricius in western North Ameri- ca ( Reduviidae; Hemiptera, Insecta). Amer. Mus. Nov. 2175:1-15.
HYMENOPTERA
BoHART, R. M. 1966. A review of Aphilanthops and related genera. Proc. Entomol. Soc. Wash. 68(2): 158-167.
BoHART, R. M., AND A. S. Menke. 1963. A reclassi- fication of the Sphecinae with a revision of the Nearctic species of the tribes Sceliphronini and Sphecini. Univ. Calif. Publ. Entomol. 30(2) :91-182.
BuREN, W. F. 1968. A review of the species of Crcmatogaster, sensu stricto, in North America ( Hvmenoptera: Formicidae). J. Georgia Entomol. Soc. 3:91-121.
Cole, A. C. 1942. The ants of Utah. Amer. Mid- land Nat. 28:3.58-388.
Cole, A. C. 1968, Pogonomyrmex harvester ants, a stiidy of the genus in North America. Univ. Tennessee Press, Kno.wille, Tennessee. 222 p.
Evans, H. E., and R, W, Matthews, 1968, North American Bembix, a revised key and suggested grouping. Ann. Entomol. Soc. Amer, 61:1284-1299.
FuRGUsoN, W. E. 1967. Male sphaerophthalmine mutillid wasps of the Nevada test site. Brigham Young Univ. Sci. Bull. Biol. Series 8(4):l-26.
Hayward, C. L. 1932. The paper wasps of Utah, including a description of a new varietv of Polistes canadensis L. Proc. Utah Acad. Sci. 9:8.5-101.
Ingham, C. O. 1959. Ants of the Virgin River Basin, southwestern Utah. Unpub. master's thesis, Univ. of Utah. Salt Lake City, Utah. 140 p.
Klieweb, J. W. 1952. Biology and control of paper wasps of Salt Lake City and vicinity. Unpub. master's thesis, Univ. of Utah, Salt Lake Citv, Utali. .36 p.
Knowlton, G. F. 1939. Utah Hymenoptera. Utah Agr. E.xpt, Sta, Mono. Series 200 (Tech.) part 4. 8 p.
Kbombein, K. V. 1967. Hymenoptera of America north of Mexico: Synoptic Catalog. USDA Agr. Mono. 2. Supplement 2. 584 p.
Krombein, K. v., et al. 1958. Hymenoptera of America north of Mexico: Synoptic Catalog. USDA Agr. Mono. 2. Supplement 1. 305 p.
Miller, C. 1961. Taxonomv and distribution of Nearctic Vespula. Can. Entomol. Supplement 22. 52 p.
Muesebeck, C. F, W., K. \'. Kbombein, H. K. Townes, et al. 1951. HvTnenoptera of America north of Mexico: SvTioptic catalog. USDA Agr. Mono. 2. 1420 p.
Rees, D. M., and A. W. Grundman. 1940. A pre-
liminary list of the ants of Utah. Univ. of Utah
Bull. 31:1-11. Rees, D. M., and J. W. Kliewer. 1953. Notes on
the biology ;ind control of wasps in Salt Lake City
and vicinity. Proc. Utah Acad. Sci., Arts and
Letters .30:114-115. Schuster, R. M. 19.58. A revision of the sphaerop-
thalmine Mutilidae of America north of Mexico II.
Entomol. Amer. 37:1-30. ScuLLEN, H. 1965. Review of the genus Cerceris in
America north of Mexico. Proc. U.S. Nat. Mus.
116:3.33-.548. ScuLLEN, H. 1968. A revision of the genus Eucer-
ceris Cresson. Bull. U. S. Nat. Mus. 268:1-97.
LEPIDOPTERA
Knowlton, G. F. 1939. Lepidoptera. Utah Agr.
Expt. Sta. Mono. Series 200 (Tech.) part 2. 14 p. SucDEN, J. W. 1937. Notes on the migrational
flights of Vanessa cardui in Utah. Pan-Pacific
Entomol. 13:109-110.
MALLOPHAGA
Behle, William E. 1971. Checkhst of the birds of
Utah. Unpublished manuscript. Elbel, R. E., and K. C. Emerson. 1956. A new
mallophagan from the White Pelican. Entomol.
News 67(7): 173-176. Emerson, K. C. 1950. The genus Lagopoecus (Phil-
opteridae: Mallophaga) in North America. J. Kan- sas Entomol. Soc. 23( 3) :97-101. Emerson, K. C. 1961. The genus Panenirmus (Mal- lophaga) found on North American woodpeckers.
J. Kansas Entomol. Soc. 34( 1 ) ; 34-43. Emerson, K. C. 1962. A tentative list of Mallophaga
for North American birds ( north of Mexico ) .
Dugway Proving Ground: 1-217. Emerson, K, C. 1964a. CheckUst of the Mallophaga
of North America ( north of Mexico ) . Part I.
Suborder Amblvcera. Dugway, Utah, (Mimeo).
171 p. Emerson, K. C. 1964b. Checkhst of the Mallophaga
of North America (north of Mexico). Part II.
Suborder Amblycera. Dugway, Utah, (Mimeo).
104 p. Emerson, K. C. 1968. The genera and species of the
Mallophaga of North America (north of Mexico).
Part I. Suborder Ischnocera. Mimeo. 194 p.
ORTHOPTERA
Barnum, a. H. 1952. The taxonomy of Utah Orth- optera with notes on distribution. Unpub. master's thesis, Brigham Young Univ., Provo. Utah. 236 p.
Barnum, A. H. 1954. The taxonomy of Utah Orth- optera. Great Ba.sin Nat. 14:39-60.
Hansen, H. O. 1963. The cockroaches of Salt Lake Countv, Utah. Unpub. master's thesis, Univ. of Utah. Salt Lake City, Utah. 65 p.
Knowlton, G. F. 1935. Bed bugs and cockroaches. Utah Agr. Expt. Sta. Leaflet 68.
SCORPIONIDA
Gertsch, W. ]., and D. M. Allred. 196.5a. Scor- pions of the Nevada test site. Brigham Young Univ. Sci. Bull., Biol. Series 6(4):1-15.
26
BniGHAM Young Univebsitv Science Bulletin
Gertsch, \V. J., AND D. M. Allred. 1965b. The scorpions of the Vejovis boreus group (subgenus Parucoctonus) in North America ( Scorpionida, Vejovidae). Amer. Mvis. Nov. 2278:1-54.
Johnson, J. D. 1968. Scorpions of Utah. Unpub. master's thesis, Brigham Young Univ., Provo, Utah. 36 p.
Wood, H. C. 1863. Description of a new species of scorpion in the Utah territory: Centrums phaeo- ckcttflus Wood. Proc. Aca. Nat. Sci. Philadelphia 2:372.
SIPHONAPTERA
Allred, D. M. 1951. A preliminary study of the distribution of fleas in Utah known to be capable and potential vectors of plague. Unpub. master's thesis, Brigham Young Univ., Provo, Utah. 141 p.
Amoureaux, R. L. 1964. Ta.xonomic study of fleas of the genus Foxella (Wagner) in Utah ( Siphonap- tera: Dohchopsylhdae ) . Unpub. master's thesis, Brigham Young Univ.. Provo, Utah. 27 p.
Beck, D E., and D. M. Allred. 19.50-1951. Fur- ther distributional data on Utah Siphonaptera. Proc. Utah Acad. Sci., Arts and Letters 28:113 ( Abstract ) .
Ecoscue, H. J. 1966a. New and additional host-flea associations and distributional records of fle;i.s from Utah. Great Basin Nat. 26:71-75.
Ecoscue, H. J. 1966b. A new species of the genus Stenistomera (Siphonaptera: Hystichopsvllidae). Bull. So. Calif. Acad. Sci. 67(3) :138-142.
Ewing, H. E., and I. Fox. 1943. The fleas of North America. Classification, identification, and geo- graphical distribution of these injurious and dis- ease-spreading insects. USDA Misc. Publ. 500. 142 p.
Howell, ]. F. 1954. A study of the a.spectional variation of Siphonaptera associated with the nests of the Thomas wood rat, Neotoma lepicla lepida Thomas. Unpub. master's thesis, Brigham Young Univ., Provo, Utah. 38 p.
Howell, J. F. 1957. Fleas associated with nests of the desert wood rat. }. Parasitol. 43:566-573.
Hubbard, C. A. 1947. The fleas of western North America. Iowa State College Press, Ames, Iowa. 533 p.
Johnson, P. T. 1961. A revision of the species of Monopsi/llus Kolenati in North America. Agr. Res. Ser. Tech. BuU. 1227. 69 p.
Johnson, P. T. 1971. Personal communication.
Parker, D. D. 1958. Sea.sonal occurrence of fleas on antelope ground squirrels in the Great Salt Lake Desert. J. Econ. Entomol. 51:. 32-36.
Parker, D. D., and J. F. Howell. 1959. Host-flea relationships in the Great Salt Lake Desert. J. Parasitol. 45(6) :. 597-604.
Smit, F. G. a. M. 1958. A new North American bird-flea. Proc. U. S. Nat. Mus. 108(3394) :51-57.
Stanford, J. S. 1931. A preliminary list of Utah Siphonaptera. Proc. Utah Acad. Sci. 8:1.53-1.54.
Stanford, J. S. 1944. More Utah Siphonaptera. Proc. Utah Acad. Sci., Arts and Letters 19-20: 173-178.
Stark, H. E. 1958. The Siphonaptera of Utah. U. S. Dept. HEW Comm. Disease Center, Atlanta, Georgia. 239 p.
Stark, H. E. 1970. A revision of the flea genus Tlirassii Jordan 1933 (Siphonaptera: Ceratoplivl- lidae) with ob.servations on ecology and relation-
ship to plague. Univ. Calif. Pub. Entomol. 53:vii -I- 184 p.
Tipton \'. J. 1949. A preliminary study of the siphonapterous ectoparasites found on the mam- mals of the families Cricetidae and Muridae in Utah County. Unpub. master's thesis. Brigham Young Univ.. Provo, Utah. 87 p.
Tipton, V. J. 1950. New distributional records for Utiih Siphonaptera. Great Basin Nat. 10:62-65.
Tipton, V. J., and D. M. Allred. 1952. New dis- tributional records of Utah Siphonaptera with the description of a new species of Meourthroglosstis Jordan and Rothschild 1915. Great Basin Nat. 11:105-114.
Traub, R., and V. J. Tipton. 1951. Jordanopsijlla allred:, a new genus and species of flea from Utah (Siphonaptera). J. Wash. Acad. Sci. 41:264-270.
COLORADO TICK FEVER
Anonymous. 1970. Colorado tick fever. RML Cir- cular 2. Rocky Mountain Laboratory, Hamilton, Montana. 1 p.
Becker, F. E. 1930. Tick-bome infections in Colo- rado. II. A survey of the occurrence of infections transmitted by the wood tick. Colorado Med. 27: 87-95.
Eklund, C. M., R. C. Kennedy, and Mary Casey. 1961. Colorado tick fever. Rockv Mountain Med. J. 58(6):21-2,5.
Eklund, C. M., G. M. Kohls, and J. M. Brennan. 1955. Distribution of Colorado tick fever and virus carrying ticks. J. Amer. Med. As.soc. 157: 335-337.
Topping, N. H., J. S. Cullvkord, and G. F. Davis. 1940. Colorado tick fever. Public Health Rpts. 55:2224-22,37.
ENCEPHALITIS
Brookman. B. 1955. The current status of encepha- litis in the west. Proc. Utah Mosq. Abate. Assoc. 8:5.
Crane, G. T., R. E. Elbel, D. E. Klimstra, and K. L. Smart. 1969. Isolation of Hart Park-like and California encephalitis group from mos(|uitoes of Cedar and Utah valleys, Utah, in 1967 (Abstract). Proc. Utah. Mosq. Abate. Assoc. 22:39-40.
Crane, G. T., R. E. Elbel, and K. L. Smart. 1970. Arbovirus isolations from mosquitoes collected in West Central Utah, 1969. Proc. Ut;ili Mosq. Abate. A.SSOC. 23:21 (Ab.stract).
Crane, G. T., R. E. Elbel, and K. L. Smart. 1970. Arbovirus isolations from mosquitoes collected in Central Utah in 1967. Amer. J. Trop. Med. Hyg. 19:540-543.
Elbel, R. E., G. T. Crane, L. E. Stipe, G. B. Van NosDOL, AND K. L. Smart. 1971. Arbovirus isolations from mosquitoes collected at Callao, Utah, 1966 and 1967. Mosq. News 31:61-68.
Gebhart, L. p., G. J. Stanton, and S. de St. Jeer. 1968. Ecology of western equine encephaUtis virus in nature. Proc. Utah Moscj. .\bate. Assoc. 21:13-15.
Graham, J. E., and G. C. Collett. 1964. Surveil- lance for western equine encephalitis in Utah. Mosq. News 24:149-153.
Graham, J. E.. I. E. Bradley, and G. C. Collett. 1960. Some factors influencing larval populations
Biological Series, Vol. 15, No, 2 Utah Arthropods
27
of Culex tarsalis and western equine encephalitis in Utali. Mosq. News 20:100-103.
Grundmann, a. W. 1950. Preliminari,' results of an investigation in Utah of mosquito borne encepha- litis. Proc. Utah Mosq. Abate. Assoc. 3:43-45.
Grundmann, A. W. 1954. The encephalitis problem in Utah. Proc. Utah Mosq. Abate. Assoc. 7:19-21. (Mimeo ).
Grundmann, A. W., and G. R. Leymaster. 1954. Equine encephalomvelitis in Utah, a survey of horses and man utilizing serum-neutralization tests. ]. Amer. Med. Assoc. 124:40-42.
Hess, A. D., and R. O. Hayes. 1967. Seasonal dy- namics of western encephalitis vims. Amer. J. Med. Sci. 253:333-348.
Hess, A. D., and P. Holden. 1957. The natural history of the arthropod-borne encephalitides in the U. S. Ann N. Y. Acad. Sci. 70:294-311.
Holden, P.. and A. D. Hess. 1959. Cache Valley virus, a previously imdescribed moscjuito-bome agent. Science 130(3383) : 1187-1188.
Jenkins, A. A., and R. Donath. 19.59. The 1958 encephalitis outbreak in northern Utah. 1. Human aspects. Mosq. News 19:221-222.
Knowlton, G. F. 1934. Insect transmission of equine encephalomvelitis studies. Utah Agr. E.\pt. Sta. Bull, 250:47 (Biennial Report).
Knowlton, G, F, 1936. Insect transmission of equine encephalom\elitis. Report of Twelfth Rocky Mt. Conf. of Entomologists, Pingree Park, Colorado, p. 9.
Knowlton, G. F., and J. A. Rowe. 1935. Handling mosquitoes on equine encephalomveUtis investi- gations, J, Econ, Entomol. 28:824-829.
Love, G. J., and D. B. Francv. 1963. Summary of activities of the encephaHtis section during 1962. Proc. Utah Mo.sq. Abate. Assoc. 16:8-11.
Madsen, D. E. 19.34. Equine encephalomyelitis. Proc. Utah Acad. Sci., Arts and Letters 11:95-99.
Madsen. D. E., and G. F. Knowlton. 1935. Mos- quito transmission of equine encephalomyelitis. ]. Amer. Vet. Med. Assoc. 86:662-666.
M.^DSEN, D. E., G. F. Knowlton, and J. A. Rowe. 1936. Further studies on transmission of equine encephahtis bv mosquitoes, J. Amer. Vet. Med. Assoc. 89:187-196,
Rees, D, ^L, AND G. C. Collett. 1959. Factors influencing the encephalitis outbreak in Utah in 1958. Proc. and Papers 27th Ann, Conf. Calif. Mosq. Control Assoc, p. 88-97.
Rees, D. M., L. J. Ocden, G. C. Collett, and J. E. GRAHA>f. 19.59. The 1958 encephalitis outbreak in northern L'tah. 3. Mosquito populations in rela- tion to the outbreak. Mosq. News 19:227-231.
Spendlove, G. a., J. A. Rowe, and C. F. Knowlton. 1953. Mosquitoes and encephalitis. Utah State Agr. Coll. Ext. Circ. 199:1-2.
Thomas, L. A., and J. V. Smith. 19.59. The 1958 encephalitis outbreak in northern Utah. 2. Infection rates in birds, mammals and mosquitoes. Mosq. News 19:223-226.
MALARIA
Grundmann, A. W., R. G, Warneck, and H. D. New- son. 1952. Nonhuman malaria in mosquitoes and the English sparrow. Passer domesiicus Linn, in the vicinitv of Salt Lake City, Utah. Mosq. News
12:.53-.57, '
Knowlton, G. F. 1948. Malariology, mosquito con- trol, and aviation. The Sanitarian 10(5): 185, 187, 189-190.
Marshall, H. L., and D. M. Rees. 1948. Malaria in Utah. Rocky Mt. Med. ]. 45:469-472.
Rees, D. M., and H. D. Newson, 1950. Mosquitoes of Salt Lake Countv, Utah, infected with nonhuman malaria. Proc. Utah Acad. Sci., Arts and Letters 27:71 (Abstract).
PLAGUE
Allred, D. M. 1952. Plague important fleas and
mammals in Utah and the western United States.
Great Ba.sin Nat. 12:67-75. Anonymous. 1936. Human and rodent plague in
California, Idaho, and Utah. Public Health Rpts.
51:1019. Anonymous. 1936. Rodent plague in California and
Utah. Public Hedth Rpts, 51:1138. Anonymous. 1936. Plague infection cUscovered in
fleas and hce taken from marmots in Montana and
in a marmot from Utah. Public Health Rpts. 51:
1159-1160. Anonymous. 1936. Rodent plague in Beaver County,
Utah, PubUc Healtli Rpts, 51:1207. Anonymous. 1936. Plague infection in California
and Utah. Public Health Rpts. 51:1257, .\nonymous. 1936. Plague in prarie dogs in Garfield
County, Uttdi. Public Health Rpts, 51:1287, Anonymous. 1937. Plague infection in fleas in Cali- fornia, Nevada, and Utah. Public Health Rpts.
52:1184-1185. Anonymous. 1937. Plague infection in Wasatch
County, Utah, and Madison County, Michigan.
PubUc Health Rpts. 52:1.321. Anonymous. 1938. Plague infection found in fleas
from desert wood rats in Utah. Public Health
Rpts. .53:1042. Anonymous. 1938. Plague infection found in ground
squirrel and fleas from ground squirrels in Utah.
Public Health Rpts. .53:1318. Anonymous, 1938, Plague infection in fleas from
ground squirrels in Rich Countv, Utah, Public
Health Rpts, 53:14.36. Anonymous. 1940. Bubonic plague in Utah. PubUc
Healdi Rpts. .55:2.58. Anonymous. 1948. Plague infection in Salt Lake
County, Utah. Public Health Rpts. 63:1478. Anonymous. 1949. Plague infection in Utah and
Washington. Public Health Rpts. 64:496. Anonymous. 1949. Plague infection in San Juan
County, New Mexico, and Sevier Countv, Utah.
Public Health Rpts. C>4:810. Beck, D E,, 1955. Distributional studies of parasitic
arthropods in Utah, determined as actual and
potential vectors of Rocky Mountain spotted fever
and plague, with notes on vector-host relationships.
Brigham Young L'niv, Sci. Bull. Biol. Series 1(1):
1-64. Byington, L. B. 1940. Two epizootics of plague in- fections in wild rodents in the western United
States in 1938, Public Health Rpts, 55:1496-1501, Hampton, B. C. 1940. Plague in the United States.
Public Health Rpts. 55:1143-1158. Hampton B. C. 1945. Plague infection reported in
the United States during 1944 and summary of
human cases, 1900-1944. PubUc Health Rpts. 60:
1361-1365.
28
BuicHAM Young University Science Bulletin
Marchette, N. J. 1963. PtisteurelUi /)C.s(i.v of low vinilcnce for mice and guinea pigs isolated from Peromyscus plague focus in Utah. Amer. J. 'I'rop. Med. Hyg. 12:215-218.
McCoy, G. W. 1911. Studies upon plague in ground squirrel. 11. A plague-like disease of rodents. Public Health Bull. 43:53-71.
MoHR, C. O. 1951. Entomological hackground on the distribution of murine typhus and murine plague in the United States. Amer. J. Trop. Med. Hyg. 31:355-372.
Mykleuust, R. J., F. C. Hahmston, and D. R, Mad- dock. 1949. Sylvatic plague investigations in Utah. CDC Publ. Fed. Sec. Agency, Atlanta, Georgia ( Mimeo ) .
Parkeh, D. D. 1971. Personal communication.
Univ. of utah. 1963. Summary status report on Paiteurella pestis. 1952-1962. Ecol. and Epizool. Ser. 92. Ecol. and Epizool. Res. Dugwav, Utah. 110 p.
Wayson, N. E. 1947. Plague-field sur\eys in western United States during ten years (1936-1945). Pub- lic Health Rpts. 62:780-791.
ROCKY MOUNTAIN SPOTTED FEVER
Beck, D E. 1955. Distributional studies of para- sitic artlxropods in Utah, determined as actual and potential vectors of Rocky Mountain spotted fever and plague, witli notes on vector host relation- ships. Brigham Young Univ. Sci. Bull. Biol. Series l(l):l-64.
Becker, F. E. 1926. Investigations on Rocky Moun- tain spotted fever in Colorado. J. Infect. Dis. 39: 81-88.
CoFFEV, M. D. 1953. Some preliminary studies of Rocky Mountain spotted fever vectors in Utah. Unpub. master's thesis, Brigham Young Univ., Provo, Utah. 84 p.
Coffey, M. D. 1954. A study of some Rock\ Moun- tain spotted fever vectors and their hosts in Utah. Great Biisin Nat. 14:31-37.
Hampton, B. C, and H. G. Eubank. 1938. Rocky Mountain spotted fever. Pubhc Health Rpts. 53: 984-990.
Herrin, C. S. 1956. Dermacentor andersoni and Rocky Mountain spotted fever in national forest recreational sites of Utah. Unpub. master's thesis, Brigham Young Univ., Provo, Utah. 51 p.
Jellison, W. L. 1945. The geographical distribu- tion of Rocky Mountain spotted fever and Nuttall's cottontail in the western United States. Public Health Rpts. 60:958-961.
LuNDCREN, D. L., B. D. Thorpe, and C. D. Haskell. 1966. Infectious di.seases in wild animals in Utah VI. E.xperimental infection of birds with RickettsUi rickcttsii. J. Bact. 91:963-966.
Parker, R. R., C. B. Philip, and W. L. Jellison. 1933. Rocky Mountain spotted fever. Potentialities of tick transmission in relation to geographical occurrence in the United States. Amer J. Trop. Med. 13:341-379.
RucKER, W. C. 1912. Rockv Mountain spotted fever. Public Health Rpts. 27:1465-1482.
Rumreich, a., R. E. Dyer, and L. F. Badceh. 1931. The typhus-rocky mountain spotted fever group. Public Health Rpts. 46:470-480.
TULAREMIA
Allhed, D. M., G. N. Stacg, and ]. F. Lavender.
1956. E.xperlmental transmission of Paiteurella
tultircnsis by the tick, Dermacentor parumapertus.
j. Infect. Dis. 99:143-145. Anonymous. 1927. Se;isonal incidence and sources of
infection of tularemia. Public Health Rpts. 42:
2948-2951. Anonymous. 1940. Tularemia (rabbit fe\er). Public
Health Rpts. 55:667-670. Barnes, W. C. 1928. Rabbit fever or tularemia.
Sci. Monthly 27:463-469. Burroughs, A. L., R. Holdenhied, D. S. Loncanecker,
AND K. F. Meyer. 1945. A field study of latent
tularemia in rodents with a list of all known nat- urally infected vertebrates. ]. Infect. Dis. 76:115-
119. Byfield, G. v., L. Bheslow, R. R. Cross, Jn., and
N. J. Hersey. 1945. Tick-borne tularemia. Re- port of 15 cases. J. Amer. Med. Assoc. 127:191-196. Clement, R. 1927. Tularemia: contamination and
transmission by rabbits. Presse med. 35:1608-1610. Cox, K. B. 1964. Bibliography of Tularemia. Mimeo.
83 p. Co.x, K. B. 1965a. Tularemia and deer flies in the
environs of Utah Lake, Utah. Unpub. master's
thesis, Brigham Young Univ., Provo, Utah. 35 p. Co.x, K. B. 196.5b. Tularemia and deer flies in the
environs of Utah Lake, Utah. Great Basin Nat.
25(1-2): 13-29. Crawford, M. 1932. Tularemia from ingestion of
insufficiently cooked rabbit. J. Amer. Med. Assoc.
99:1497-1498. Francis, E. 1919. Deer-fly fever, or Pahvant Valley
plague. A disease of man of hitherto imknown
etiology. Public Health Rpts. .34:2061-2062. Francis, E. 1921. Tularemia Francis 1921. I. The
occurrence of tularemia in natxire as a disease of
man. Public Health Rpts. .36:1731-1738. Francis, E. 1922. Tularemia Francis 1921: A new
disease of man. J. Amer. Med. Assoc. 78:1015-1018. Francis, E. 1925. Tularemia. J. Amer. Med. Assoc.
84:1243-1250. Francis, E. 1926. Tularemia. Hektoen Lecture. Proc.
Inst. Med. Chicago. 6:27-37. Francis, E. 1928a. Symptoms, diagnosis and pathol- ogy of tularemia. J. Amer. Med. Assoc. 91:1155-
1161. Francis, E. 1928b. History of Tularemia. DeLamar
Lecture, Johns Hopkins Univ., Sch. Hvg. p. 94-115. Francis, E. 1928c. A summary of present knowledge
of tularemia. Medicine 7:411-432. Francis, E. 1928d. Summary of present knowledge
of tularemia. Harvev Lecture. Harvey Soc. N. Y.
Series 1927-1928:2.5-48. Francis, E. 1929. Arthropods in the transmission
of tularemia. Trans. Fourth Int. Cong. Entomol.
2:929-944. Francis, E. 1937. Sources of infection and seasonal
incidence of tularemia in man. Public Health Rpts.
.52:103-113. Francis, E., and G. R. Callender. 1927. Tularemia,
the microscopic ch;uiges of the lesions in man.
Arch. Path. Lab. Med. 3:.577-607. Francis, E., et al. 1922. Tularemia Francis 1921:
A new disease of man. USPIIS Hvg. Lab. Bull.
130. 87 p. Francis, E., and G. C. Lake. 1921. Tularemia
Francis 1921. III. E.\perimental transmission of
tularemia in rabbits by the rabbit louse Haemodip-
Biological Series, Vol. 15, No. 2 Utah Ahthhopods
29
sus ventricosus (Dennv). Public Health Rpts. 36:1747-1753.
Francis, E., and G. C. Lake. 1922a. Tularemia Francis 1921. IV. Tran.smission of hilaremia by the bedbug, Cinicx lectiihiritis. Public Health Rpts. 37:83-95.
Francis, E., and G. C. Lake. 1922b. Tularemia Francis 1921. V. Transmission of tularemia by the mouse louse Poh/plax scrmtus (Burm.). Pubhc Health Rpts. 37:96-101.
Francis. E., and B. Mavne. 1921. Tularemia Francis 1921. 11. Experimental transmission of tularemia bv flies of the species Chrysops di.icalis. Pubhc Health Rpts. 36:1738-1746.
Francis, E., and D. Moore. 1926. Identity of Ohara's disease and tularemia. J. Amer. Med. As.soc. 86:1329-1.332.
Gebhardt, L. P., and B. D. Thorpe. 1962. A review of tularemia. L'niv. Utah Manuscript. 87 p.
Hillman, C. C., and M. T. Morgan. 1937. Tula- remia: report of a fulminant epidemic transmitted by the deer fly. ]. Amer. Med. Assoc. 108:538-.540.
Jellison, \V. L. 1950. Tularemia: Geographical dis- tribution of "Deer-fly fever" and the biting fly, Chn/sops discalis Williston. Public Health Rpts. 63:1321-1329.
Jellison, W. L. 1970. Tularemia in Montana. Mon- tana WildUfe: 1-24.
Jellison, W. L. 1971. Personal communication.
Jellison, W. L., and G. M. Kohls. 1955. Tularemia in sheep and in sheep industry workers in western United States. Public Health Mono. 28. 17 p.
Jellison, W. L., G. M. Kohls, and C. B. Philip. 1951. Tularemia: muskrats as a source of human infecHon in Utah. Rocky Mt. Med. J. 48:594-597.
Jellison, W. L.. and R. R. Parker. 1945. Rodents, rabbits, and tularemia in North America: Some zoological and epidemiological considerations. Amer. J. Trop. Med. 25:349-362.
JuNKiN, H. D. 1927. Two unusual cases of tularemia. Northwest Med. 26:415.
Knudsen, A. B., D. M. Rees, and G. C. Collett. 1968. Tularemia in Salt Lake County, Utah. Proc. Utiih Mosq. Abate. Assoc. 21:15-16.
Larson, C. L.. \V. Wicht, and W. L. Jellison. 1955. A new organism resembling P. tularcn.m isolated from water. Public Health Rpts. 70:253-258.
LuNDGREN, D. L.. N. J. Marchette, and p. S. Nich- ols. 1961. Certain immunological responses of wild rodents and laboratory- animals following chal- lenge with PastcureUa tularensis. Ecol. and Epizool. Series 60, June 30, 1961. Inst. Environ. Res. Univ. Utah.
LuNDCREN, D. L., N. J. Marchette, and K. L. Smart. 1957. Tularemia in the coyote, Ctinis latrans
lestes Merriam. J. Infect. Dis. 101: 154-157.
Marchette, N. J., D. L. Lundcren, P. S. Nicholes, and E. D. Ve.st. 1961. Studies on infectious diseases in wild animals in Utah. I. Susceptibihty of wild animals to experimental tularemia. Zoonoses Res. 1:49-73.
McCoy, G. W.. and C. W. Chapin. 1912. Further observations on a plague-like disease of rodents with a preliminary note on the causative agent. Bacterium tularense. J. Infect. Dis. 10:61-72. Parker, D. D. 1957. Attempted transmission of Pasteurella tularensis bv three species of fleas. J. Econ. Entomol. 50:724-726. Parker, D. D., and D. E. Johnson. 1957. Experi- mental transmission of Pasteurella ttihrensis by the
flea, Orchopeas leucopus (Baker). J. Infect. Dis. 101:69-72.
Parker, R. R., and E. Francis. 1926. Susceptibihty of the cx)yote (Canis lestes) to tularemia. Public Health Rpts. 41:1407-1410.
Pearse, R. A. 1911. Insect bites. Northwest Medi- cine, N. S. 3:81.
St.\gg, G., W. S. Tanner, and J. Lavender. 1959. Experimental infections of native animals with Pasteurella tularensh. J. Infect. Dis. 99:34-37.
Thorpe, B. D. 1962. Review of tularemia in arthro- pods and birds. Dept. Bact. Univ. Utah. Unpub. Manuscript 64 p.
Thorpe, B, D., R. W. Sidwell, D. E. Johnston, K. L. Smart, and D. D. Parker. 1965. Tularemia in the wildhfe and livestock of the Great Salt Lake Desert Region, 1951 through 1964. Amer. J. Trop. Med. Hyg. 14:622-637.
Vest, E. D., and N. J. Marchette. 1958. Trans- mission of Pasteurella tularensis among desert ro- dents through infective carcas.ses. Science 128 (3320): .36.3-364.
Woodbury, A. M. 1955. Ecological studies of tula- remia in Utah. In Symposium on ecology and dis- ease transmission in native animals. U. S. Army Chemical Corps, Dugway. Utah. 112 p.
MISCELLANEOUS
Allred, D. M. 1956. Ticks and mites as potential vectors. In Symposium on ecology of disease trans- mission in native animals, p. 66-69. Special Publ. Ecol. Res., Univ. Utah and Army Chemical Corps, Dugway, Utah.
Anonymous. 1958. Morbidity: Utah 1958. Utah State Dept. Health, Bureau of Vital Statistics, Salt Lake City, Utah. 94 p.
Anonymous. 1968. UtaJi Vital Statistics, 1968. An- nual Rpt. State of Utah, Dept. Soc. Services, Divi- sion of Health. Salt Lake City, Utah. 126 p.
Armstrong, C. 1922. Typhus fever on the San Juan Indian Reservation, 1920 and 1921. Public Health Rpts. 37:685-693.
Ashley, R. F. 1968. The distribution of overwinter- ing arthropods in a sheep pasture in Utah County, Utah. Unpub. master's thesis, Brigham Young Univ., Provo, Utah. 19 p.
Beck, D E. 1953. A study of some consortes found at a nesting site of the northern Cliff Swallow Petrochelidon alhifrons alhifrons ( Rafinesque ) . Proc. Utah Acad. Sci., Arts and Letters 30:39-42.
Beck, D E., A. H. Barnum, and L. Moore. 1953. Arthropod consortes found in nests of Neotoma cinerca acraia ( Ord. ) and Neotoma lepida lepida Thomas. Proc. Utah Acad. Sci.. Arts and Letters 30:43-52.
Berge, T. O., and E. H. Lennette. 1953. World distribution of Q fever: Human, animal, and ar- thropod infection. Amer. J. Hyg. 57:12.5-143.
CoTT, E. H. 1956. Ecological observations on the insect fauna of the Great Salt Lake Desert and their epizootic implications. In Symposium on ecol- ogy of disease transmission in native animals, p. 69-73. Special publ. Ecol. Res., Univ. Utah and Army Chemical Corps, Dugway. Utah.
CoTT, E. H., D. M. Allred, and E. J. Roscoe. 1956. The ectoparasite rearing program. In Symposium on ecology of disea.se transmi.ssion in native animals, p. 59-61.' Special publ., Ecol. Res., Univ. Utah and Army Chemical Corps, Dugway, Utah.
30
Bhigham Vounc. University Science Bulletin
Davidoff, E. B. 1956. Ectoparasites of birds and mammals, /ii .SvTnposiiim on ecolog\' of disease transmission in native animals, p. 61-66. Special pnbl. Ec'ol. Res., Univ. Utah and Army Chemical Coqis, Dugway, Utah.
Davis, G. E. 1939. Ornithodows purkcri: Distribution and host data; spontaneous infection with relapsing fever spirochetes. Public Health Rpts. 54; 1345- 1349.
Davis, G. E. 1940. Ticks and relapsing fever in the United States. Pui)lic Health Rpts. 55:2347-2351.
Davis, G. E. 1941. Ornithodorn.i parkvri and relaps- ing fever .spirochetes in Ut;ih. Public Health Rpts. 56:2464-2468.
Elzinga, R. J., and D. M. Rees. 1964. Comparative rates of ectoparasite infestation on deer and harvest mice. Proc. Utah Acad. Sci., Arts and Letters 41: 217-220.
Felsenfeld, O. 1965. Borreliae, human relapsing fever, and parasite-vector relationships. Bac. Rev. 29:46-74.
Ho, Benc Chaun, and D. M. Rees. 1963. Ectopara- site-host associations in three areas in Utah and Wyoming. Proc. Utah Acad. Sci.. Arts and Letters 40:115.
HoRSFALL, F. L., AND I. TA^l^I ( eds. ) . 1965. Viral and Rickettsial infections of man. Philadelphia, Lippincott. 1282 p.
Howell, J. F. 1960. Arthropod consortes of a kit fox den. Great Basin Nat. 20:71-77.
James, M. T., and R. F. Harward. 1969. Herm's Medical Entomology (sixth edition). MacMillan Co., New York, 484 p. -|- viii.
Jellison, W. L. 1956. Rodent disease and parasites in Washington County, Utiih. with particular ref- erence to coccidioidomycosis. Proc. 11th Ann. Meeting INCDNCM, Salt Lake City, Utah. Aug- ust 30, 31, and September 1, 1956. p. 96-104.
Knowlton, G. F. 1960. Insect conditions in Utah — 1959 (Abstract). Proc. Utah Acad. Sci., Arts and Letters 37:151-152.
Knowlton, G. F. 1966. Entomology — observations. Proc. Utah Acad. Sci., Arts and Letters 43:1.58.
Lundgren, D. L., N. J. Marchette, and P. A. Nich- OLES. 1961. Studies on the ecology and epi- zoolog)' of the native fauna of the Great Salt Lake Desert. Ecol. and Epizool. Series 60. June 30, 1961. Inst. Environmental Res. Univ. Utah. '
MATHE.SON, R. 1950. Medical Entomology. Comstock Publishing Associates, Ithaca, New York. 612 p.
MvKLEBUsT, R. J. 1951. Some studies on the con- sortes of the Norway rat in Utah County, Utah. Unpub. master's thesis, Brigham Young Univ., Pro\o. Ut:di. 73 p.
Pahrish, H. M. 1959. Deaths from bites and stings of venomous animals and insects in the United States. Arch. Int. Med. 104:198-207.
Philip, C. B. 1969. Tick paralysis. RML Circ. 4. Rocky Mount;un L;iboratorv, Hamilton, Montana. 2p. •
Pratt. H. D.. and P. L. Rice. 1969. Current status of some important arthropod-borne diseases in the United States. Dept. HEW. PHS 30 p.
Rees, D. M., and B. C. Ho. 1961. Arthropods found in Flaming Gorge Reservoir Basin, 1959. Ihiiv. Utah Anthropol. Papers No. 48:100-110, November 1960, pub. 1961.
Roberts, R. S. 1954. Moscjuitoes and the disejLse svTiovitis ( Staphyloc(xx:osis ) in turkeys. Proc. Utah Mosq. .Abate. Assoc. 7:17-19.
SiDWELL, R. \V'., D. L. Lundgren, J. B. Bushman, and B. D. TiioRi'E. 1964. The occurrence of possible epizootic of Q fever in fauna of the Great Salt Lake Desert of Ut;ili. Amer. J. Trop. Med. Hyg. 13:754-762.
Smart, J. 1948. A handlxKik for the identification of insects of mediciil imjxjrtance. British Museum (Niitural History). London. 295 p. + XllI pis.
Stanford, J. S. 1934. Some ectoparasites of Utah birds and mammals. Proc. Utah Acad. Sci., Arts and Letters, 11:247.
Stoenner, H. G., R. Holdenried, D. Lackman, and J. S. Osborn, Jr. 1959. The occurrence of Coxiella burnetii. Brucella, and otiier pathogens among the faima of the Great Salt Lake Desert in Utah. Amer. J. Trop. Med. Hyg. 8:590-596.
Univ. of Utah. 19.55a. Ecolog)' of disease trans- mission in native animals. Ecol. Res. Dugway, Utah. 16 p.
Univ. of Utah. 1955b. Symposium on ecology of disease transmi.ssion in native animals. Ecol. Res. Dugway, Utah. 112 p.
Univ. of Utah. 1955-1956. A study of the ecology of infectious cUseases in the native fauna. Ecol. Res. Dug\y;iy, Utah. 78 p.
Univ. of Utah. 1957. A study of the ecology of infectious dLseiises in the native fauna of the Great Salt Liike Desert. Ecol. Res. Dugway. Utah. 49 p.
Univ. of L'tah. 1959a. Studies on the ecology and epizoology of the nati\e fauna of the Great Salt Lake Desert. Ecol. and Epizool. Series 41. Ecol. and Epizool. Res. Group, Dugway, Utah. 6.3 p.
L'niv. of Utah. 1959b. Epizoological survey of certain endemic diseases in the southern part of the Great Salt Lake Desert. Ecol. and Epizool. Series 42. Ecol. and Epizool. Res. Group, Dug- way, Utah. 18 p.
Univ. of Utah. 1960. Studies on the ecoIog\- and epizoology' of the native fauna of the Great Salt Lake Desert. Ecol. and Epizool. Series 44. Ecol. and Epizool. Res. Group Dugway, Ut;di. 67 p.
Univ. of Utah. 1961. Studies on the ecology ;md epizoologN' of the native faima of the Great Salt L:ike BiLsin. Ecol. and Epiz(«)l. Series 63. Ecol. and Epizool. Res. Group, Dugway. L'tah. 59 p.
L'niv. of L'tah. 1962. A study of the ecology- and epizoolog\- of the native faima of the Gre;it Salt Lake Desert. Ecol. and Epizool. Series 70. Ecol. and Epizool. Res. Group, Dugway, Utah. 103 p.
Univ. of Utah. 1963. A study of the ecology and epizoology of the native fauna of the Great Salt Lake Desert. Ecol. and Epizool. Series 100. Ecol. and Epizool. Res. Group, Dugway, Utah. 127 p. + 29 figs.
Univ. of Utah. 1964. A studs of the ec'ologN- and epizoology of the nati\e faiuia of the Gre;it S;dt Lake Desert. Ecol. and Epizool. Series 108. Ecol. and Epizool. Res. Group, Dugway. Ut;ih. 192 p.
Univ. of L'tah. 1965. A studv of the ecolog\ and epizoology of the native fauna of the Great Salt Lake Desert, 1964. Ecol. and Epiz(K)l. Series 117. Ecol. and Epizool. Res. Group, Dugw;i\, Ut;ih. 180 p.
Univ. of Utah. 1966. A study of the ecology ;md epizoology of the native fauna of the Gre;it Salt L;ike Desert. 1965. Ecol. ;ind Epizool. Series 123. Ecol. and Epizool. Res. Group, Dug\va\. Utah. 171 p.
Unb-. of I'taii. 1967. ,\ studv of the ecology^ and epizoology of the nati\e fauna of the Great Salt Lake Desert. 1966. Ecol. and Epizool. Series 141.
Biological Series, Vol. 15, No. 2 Utah Arthhopods
31
Ecol. and Epizool. Res. Group, Diigwa\, Utah.
313 p. Univ. of Utah. 1968. A study of the ecology and
epizcx>loi;N- of the native fauna of the Great Salt
Lake Desert, 1967. Ecol. and Epiztxil. Series 144.
Ecol. and Epizool. Res. Group. Dugwav, Ut;ili.
234 p. Univ. of Utah. 1969. A study of the ecology and
epizoology of the native fauna of the Great Salt
Lake Desert, 1968. Ecol. and Epiz(x>I. Series 145.
Ecol. and Epizool. Res. Group, Dugway, Utaii.
244 p. Vest, E. D., et al. 1961. Studies on ecology of Q
fever in native fauna in the Great Salt Lake Des-
ert. Ecol. and Epizool. Series 66. Summary pro- gress report. Inst. Environ. Biol. Res. Univ. Utah.
Vest, E. D., D. L. Lundcren, D. D. Parker, D. E. Johnston, E. L. Morse, J. B. Bushman, R. W. SiDWELL, AND B. D. Thorpe. 1965. Results of a five-year survey for certain enzootic disea.se in the fauna of western Utali. Amer. J. Trop. Med. Hyg. 14:124-135.
Woodbury, A. M. (ed. ). 1956. Ecological check lists: The Great Salt Lake series. Ecological Res. Univ. Utah. 125 p. (Mimeo).
Woodbury, A. M. 1964. Disease di.s,scmination among biotic communities of the Great Salt Lake Desert. Univ. Utah. Div. Biol. Sci. Misc. Papers 4. 282 p.
Brigham Young University Science Bulletin
MUS. COMP. ZOOL LIDRARY
FEB 1 8 IQ72
HARVARD UNiVERSITY
NEW RECORDS AND SPECIES OF
NEOTROPICAL BARK BEETLES
(SCOLYTIDAE: COLEOPTERA)
PART V
by Stephen L. Wood
BIOLOGICAL SERIES — VOLUME XV, NUMBER 3 DECEMBER 1971
BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN BIOLOGICAL SERIES
Editor: Stanley L. Welsh, Department of Botany,
Brigham Young University, Prove, Utah
Members of the Editorial Board:
Vernon J. Tipton, Zoology Ferron L. Anderson, Zoology Joseph R. Murdock, Botany Wilmer W. Tanner, Zoology
Ex officio Members:
A. Lester Allen, Dean, College of Biological and Agricultural Sciences Ernest L. Olson, Chairman, University Publications
The Brigham Young University Science Bulletin, Biological Series, publishes acceptable papers, particularly large manuscripts, on all phases of biology.
Separate numbers and back volumes can be purchased from Publication Sales, Brigliam Young University, Provo, Utah. All remittances should be made payable to Brigham Young University.
Orders and materials for library exchange should be directed to the Division of Gifts and Exchange, Brigliam Young University Library, Provo, Utah 84601.
Brigham Young University Science Bulletin
NEW RECORDS AND SPECIES OF
NEOTROPICAL BARK BEETLES
(SCOLYTIDAE: COLEOPTERA)
PART V
by Stephen L. Wood
BIOLOGICAL SERIES — VOLUME XV, NUMBER 3 DECEMBER 1971
TABLE OF CONTENTS
ABSTRACT 1
INTRODUCTION 1
SYSTEMATIC SECTION 2
Phrixnsama crehmm, n. sp 2
Phrixosoma viriosum, n. sp 2
Phrixosonui frustratum, n. sp 3
Chmtncfius macrocornis, n. sp 3
Chramesus simplicis. n. sp 3
Chramesus orinocetisis, n. sp 4
Chramesus strigilis, n. sp 4
Clirarnesus imporcatus, n. sp 5
Chramesus impolitus, n. sp 6
Chramesus parcus, n. sp 6
Chramesus denticulatus, n. sp 6
Chramesus prisons, n. sp 7
Chramesus vinealis, n. sp 7
Chramesus solicitatus, n. sp 8
Chramesus pcnicuhis, n. sp 8
Chaetophlocus atidinus, n. sp 9
Liparthrum carapae, n. sp 9
Liparthrum mcridensis. n. sp 10
Cladoctonus holiviae (Wood) _ 10
Ptjcnarthrum inornatum, n. sp 11
Pycnarthrum funerium, n. sp 11
Pycnarthrum fici, n. sp II
Pycnarthrum pcrditum. n. sp 12
Pycnarthrum hicidum, n. sp 12
Pycrmrthrum carinatum, n. sp 13
Pycnarthrum suhcarinatum, n. sp 13
Pycnarthrum brosniii, n. sp 14
Gymnochihis alni, n. sp 14
Scolytodes ommateus, n. sp 15
Scolytodes punctifer, n. sp 15
Scohjtodes hirsuiits, n. sp 16
Scolytodes crassus, n. sp 16
Scohjtodes pannuceus, n. sp 16
Scolytodes micidus, n. sp 17
Microborus lectus, n. sp 17
Scolytus harinensis, n. sp 18
Scolytopsis orinocanus, n. sp 18
Carphodicticus, n. gen 19
Carphodicticus cristatus, n. sp 19
Pseudotht/sanoes querneus, n. sp 20
Pseudothtjsanoes jurvescerxs, n. sp 21
Pseudothysanoes junereus, n. sp 21
Pseudothi/sanoes graniticus, n. sp 21
Pseudothysanoes verticillus, n. sp 22
Pseudothysanoes cuspidis, n. sp - 22
Pseudothysanoes tenellus, n. sp 23
Pseudothysanoes yaccavonts, n. sp 23
Thysanoes inornatus, n. sp 24
Micracisella adnata, n. sp - 24
Micracisella scrjaniae, n. sp 25
Micracis amplinwi, n. sp 25
Micracis incertus, n. sp 26
Micracis torus, n. sp 26
Micracis exilis, n. sp 27
Micracis vitulus, n. sp 27
Micracis sentus, n. sp 28
Ilylociirus torosus, n. sp 28
Ili/locuriis verrucosus, n. sp 28
Ili/locurus (lilulus, n. sp 29
Hylacurus dispurilis, n. sp 29
Htjlocurus dissidens, n. sp 30
Hyhcurus villifrotts, n. sp 30
Hylocurus singularis, n. sp 31
Hylocurus fktgellutus, n. sp 32
Stegnmerus mirandus, n. sp 32
Phacrylus pruni, n. sp 33
Periocn/pludus. n. gen 33
Periacryphalus pullus, n. sp 33
Hypothenemus nanellus, n. sp 34
Hypothenemus ascitus, n. sp 35
Hypothcncmus teretis, n. sp 35
Cryptocarinus coroiuitus, n. sp _ 36
Cryptocarinus lepidus, n. sp 36
Dendrocranulus tardulus, n. sp 37
Dendrocranulus divcrsus, n. sp 37
Dendrocranulus pumilus, n. sp 38
Dendrocranulus lityius, n. sp _ 38
Xyleborus ehenus, n. sp 39
Mimips mimicus Schedl 39
Mimips analogus, n. sp _ 40
Mimips fortis, n. sp - 40
Mimips bidens, n. sp 41
Mimips uncinatus, n. sp 41
Mimips ocularis, n. sp 42
Styplilosoma granuhitum Blandford 43
Styphlosoma suhulatum, n. sp 43
Dentroterus eximius, n. sp 44
Dentroterus parilis, n. sp 44
Dentroterus sndalis, n. sp 44
Dentroterus resolutus, n. sp 45
Dentroterus defcctus, n. sp 45
Dentroterus cognatus, n. sp 46
Phelloterus, n. gen 46
Phelloterus tcrsus, n. sp 47
Phelloterus anaxeus, n. sp 47
Phelloterus atrocis, n. sp 48
Pityophthorus arceuthobii, n. sp 48
Pityohorus hondurensis, n. sp 49
Pityohorus frontalis, n. sp 49
Pseudopityophthorus singularis, n. sp 50
Pseudopityophtliorus virilis, n. sp 50
Pseudo pitt/ophthorus declivis, n. sp 50
Pseudopityophthorus colombianus, n. sp 51
Ciuithophthorus ctenmtis, n, sp 51
Gnathophthonis cracens, n. sp _ 53
Cruithophthorus rallus, n. sp 53
Gnathophthorus pertusus, n. sp 53
Sphenoceros aztecus, n. sp 54
NEW RECORDS AND SPECIES OF NEOTROPICAL BARK BEETLES (SCOLYTIDAE: COLEOPTERA), PART V
by
Stephen L. Wood-
ABSTRACT
The tribe Carphodicticini, the genera Car- ]>hodicticus, Pcnocnjphalus, and Phelloterus, and 101 species are described as new to science. Notes on the habits of Cladoctonus boliviae (Wood), Mimips mimicus Schedl, and Styphlo- soma Qramilatum Blandford are included; and the female of Slijphlosoma granulatum Bland- ford is described. The species new to science include the following: Phrixosonm crehnim, P. jrustratum (Colombia), P. viriosum, Chramesus macrocornis, C. orinocensis, C. striQiUs, C. im- porcatus, C. impolitus, C. parens, C. denticu- latus, C. priscus, C. vinealis, C. solicitatus, C. penicuhts, ChaetophJoeus andinus, Liparthrum carapae, L. inerideiisis (Venezuela), Pijcnarth- riim inornatum, Pijc. funerium, Pijc. perditum (Honduras), Pijc. fici (Honduras, Venezuela), Pijc. Jucidum (Costa Rica), Pijc. cariimtum (Peru), Pyc. suhcarinatum (Venezuela), Pijc. brosmii (Venezuela, Colombia), Gijinnochdus alni (Mexico), Scohjtodes ommateus (Colom- bia), S. punctifer, S. Jiirsnhis (Costa Rica), S. crassus (Panama), S. pannuceus (Honduras), S. micidtis (Mexico, Guatemala), Microhorus lectus, Scohjttis barinensis, Scohjtopsis orinocan- us. Carpliodicticus cmtatits (Venezuela), Pseu- dothifsanoes cptenietis, Ps. ftirvescens, Ps. funer- eus, Ps. graniticus, Ps. verticillus, Ps. cuspidis, Ps. tenellus, Ps. tjuccavorus, Thijsanoes inornat- us. MicracuseUa adnata (Mexico), M. serjaniae
(Honduras), Micracis amplinis, Mic. incertus, Mic. torus (Mexico), Mic. exilk, Mic. vitulus, Mic. sentus (Venezuela), Htjlocurus torosus (South Carolina), H. dilutus, H. dissidens (Mexico), H. disparilis (Honduras), H. verruco- sus, H. viUijrons, H. singularis, H. flageUatus, Stegomerus mirandus, Phacrylus pruni, Periocry- plwlus puUtis (Venezuela), Hypothenemus nan- ellus, H. ascitus (Costa Rica), H. teretis (Costa Rica, Venezuela), Cn/ptocarenus coronatus (Venezuela), C lepidus (Costa Rica, Guate- mala), Dendrocranulus tarduhis, D. diversus, D. pumilus, D. limus (Costa Rica), Xyleborus ebe- nus (Costa Rica, Panama), Mimips analogus, Mini, iincinatus (Venezuela), Mini, fortis (Costa Rica, Panama), Mim. bidens, Mim. ocu- laris (Colombia), Styphlosoma subulatum (Ven- ezuela), Dendrotcrus exiniius. Den. sodalis (Guatemala), Den. parilis. Den. resolutus (Costa Rica), Den. defectus (Costa Rica, Pana- ma), Den. cognatus (Mexico), Phelloterus tersus (Venezuela), Ph. anaxeus, Ph. atrocis (Colombia), Pityophthorus arceuthobii (Mexi- co), Pityoborus hondurensis (Honduras), Pit. frontalis (Mexico), Pseudopityophthorus singu- laris, Pd. virUus, Pd. declivis (Mexico), Pd. co- lombianus (Colombia), Gnathophthorus clema- tis (Mexico), G. cracens (Honduras), G. rallus, G. pertusus (Venezuela), and Splienoceros az- tecus (Mexico).
INTRODUCTION
While preparing a taxonomic monograph of the Scolytidae of North and Central America a large number of species new to science were discovered. Because it will be several years before that work will be concluded, the new names are being published in order to stabilize nomenclature and to facilitate identification.
On the following pages 101 species and three genera are described as new to science; a new tribe Carphodicticini is also proposed for the genera Carphodicticus Wood, described below, and Craniodicticus Blandford. The new species represent the following genera: Phrixosoma (3), Liparthrum (2), Chaetophloeus (1), Chramesus
*Most of the field work that led to the discovery of these insects was sponsored by the National Science Foundation. n^epartment of Zoology, Brigham Young University. Provo. Utah. Scolytoidea contribution number 42.
Bhigham Young University Science Bulletin
(12), Pycnarthrum (8), Gymnochilus (1), Sco- It/todes (6), Micwhoms (1), Scohjtus (1), Sco- lytopsis (1), Carphodicticus (1), Pseudothij- sanoes (8), Thijsanoes (1), Micracisella (2), MicracK (6), Hylocurus (8), Sfegowienw (1), Phacrylus (1), Hyjiotheneinus (3), Periocry- phalus (1), Cryptocarenus (2), Deiidrocmnulus (3), Xyleborus (1), Miinips (5), Styphlosoma (1), Dendroterus (6), PheUoteru.s (3), PUyoph- thortis (1), PHijobonis (2), Pseiidopityophthor- tts (4), Gnathoplithorus (4), and Sjilienoceros
The new species are from the following countries: United States (1), Mexico (25), Guatemala (2), Honduras (8), Costa Rica
(12), Panama (1), Colombia (8), Venezuela ( 38 ) , and Peru ( 1 ) . One species in each of the following combinations of countries is also in- cluded: Mexico-Honduras-Costa Rica; Hondur- as-Venezuela; Honduras-Costa Rica; Guatemala- Costa Rica; and (Colombia-Venezuela.
Except as noted below, the type series were tentatively deposited in my collection presently housed at the Brigham Young University, in order to facilitate preparation of a monograph of the Scolytidac of the Western Hemisphere. Upon completion of that monograph paratypes will be distributed to several major cooperating museums.
SYSTEMATIC SECTION
Phrixosoma crehrum, n. sp.
This species is closely allied to minor Wood, but it is distinguished by the much more weakly impressed transverse line on the female vertex and by the finely punctured pronotum with granules restricted to the margins.
Female.— Length 1.8 mm (paratypes 1.5-1.9 mm), 1.9 times as long as wide; color almost black.
Frons as in minor except more coarsely re- ticulate-granulate, line on vertex and laterally, of same pattern but much less strongly im- pressed. Pronotum as in minor, except surface smooth, shining, punctures irregularly shaped, distinctly, shallowly impressed, granules indica- ted only near base and lateral margins. Elytra as in minor except surface more strongly reticu- late-granulate, interstrial granules evidently slightly smaller.
Male.— Similar to female except devoid of impressed and sinuate line on vertex.
Type Locality.— Eight km south of Colonia (near Buenaventura), Valle de Cauca, Colombia.
Type Matekial.— The female holotype, male allotype, and 49 paratypes were collected on 9 July 1970, .30 m elevation. No. 640, from Rheedia miidrufw, by S. L. Wood. The biramose parental galleries were more or less longitudinal.
The holotype, allotype, and paratypes are in my collection.
Phrixosoma viriostim, n. sp.
This species is remotely allied to obesum Blackman, but it is distinguished by the much
larger size, by the pronotal sculpture, and by the much finer, more numerous interstrial gran- ules.
Female.— Length 2.7 mm (paratypes 2.4- 2.7 mm), 1.8 times as long as wide; color almost black.
Frons as in crebrum Wood, except very slightly less strongly convex, and line on vertex not at all modified or impressed; vestiture very short, moderately abundant.
Pronotum outline much as in crebrum except median area extended posteriorly more than in other species; surface densely, strongly, finely reticulate-granulate, intermixed with abundant, shining, small, irregular, somewhat elongate, rounded granules evidently derived from mar- gins of punctures. V^estiture of rather abundant short, stout recumbent bristles.
Elytra 1.1 times as long as wide, 1.7 times as long as pronotum; sides weakly arcuate on basal two-thirds, rather narrowly rounded be- hind; striae narrowly, very deeply impressed, longitudinally strigose, punctures very smidl, rather deep, rather widely spaced; interstriae about four times as wide as striae, strongly re- ticulate-granulate, with very numerous, con- fused, narrow crenulate granules, each granule about one-fourth as wide as an interstriae on basal half. Declivity beginning about middle of elytra, gradual, convex; interstriae becoming about half as wide as on disc, granules largely obsolete before middle of declivity. Vestiture of abundant, slender, curved scales of equal length, confused.
Male.— Indistinguishable from female by external characters.
BioLCx;iCAL Series, Vol. 15, No. 3 New Neotropical Scolytidae
Type Locality.— Forty km .southeast of So- copo, Barinas, Venezuela.
Type M.\terl\l.— The female holotype, male allotype, and 96 paratypes were taken at the type locality on 25 January 1970, 150 m eleva- tion. No. 254, from Rheedia madruho, by S. L. Wood. The biramose parental galleries were longitudinal.
The holotype, allotype, and paratypes are in my collection.
Phrixosoma frustraium, n. sp.
This species may be remotely allied to virios- um Wood, but it is unique in having the striae much more v^'eakly, irregularly impressed, and in having, in the female, a large, subquadrate extension of the anterior margin of the pronotum fitting snugly against the head behind the eye.
Female.— Length 2.S mm (paratypes 2.7-3.0 mm), L6 times as long as wide; color almost black.
Frons as in viriostim except granules finer. Pronotimi as in viriostim except posteromedian area less strongly produced, and surface with very indistinct small punctures indicated and granules smaller, closer. Vestiture of fine and coarse short hair. Anterolateral margin of pro- notum with a large, subquadrate extension pro- jecting forward flush with side of head between upper and lower halves of eye and equal in area to upper half of eye.
Elytra as in viriosum except striae much less strongly impressed, granules slightly smaller, vestiture much more slender, a central row on each interstriae slightly stouter and very slightly longer on declivity.
Male.— Similar to female except process on anterolateral margin of pronotum absent.
Type Locality.— Eight km south of Colonia (near Buenaventura), Valle de Cauca, Colombia.
Type Material.— The female holotype, male allotype, and 93 paratypes were taken at the type locality on 9 July 1970, 30 m elevation, from Rlieedia madruho, by S. L. Wood. The biramose parental galleries were longitudinal.
The holotype, allotype, and paratypes are in mv collection.
Chramesus macrocornis, n. sp.
This species evidently is allied to hylurgoides Schedl, but it is smaller, it has much smaller strial punctures, and the male frons is armed by a pair of very large tubercles.
Male.— Length 2.3 mm (paratypes L9-2.3 mm), 2.1 times as long as wide; color very dark brown, elytra lighter.
Frons deeply concave from epistoma to upper level of eyes; lower lateral margin at level of antennal insertion armed by a pair of very large, triangular, almost hornlike processes; surface minutely reticulate, dull; concavity im- punctate, fine, sparse, shallow punctures above upper level of eyes; vestiture short, fine, sparse. Antennal club typical of genus; rather large; apex rounded.
Pronotum 0.85 times as long as wide; widest at base, sides arcuately converging to slight constriction just before broadly rounded, sub- emarginate, anterior margin; surface subreticu- late, shining, entirely devoid of granules and asperities; punctures rather small to moderately large intennixed, rather deep, moderately close except very close near base. Vestiture of very fine, short, inconspicuous hair.
Elytra 1.3 times i\s long as wide; sides al- most straight and parallel on basal two-thirds, broadly rounded behind; striae 1 moderately, others feebly or not at all impressed, punctures moderately large at base, gradually decreasing in size until minute at base of declivity; inter- striae smooth, twice as wide as striae at base, punctures mostly uniseriate, finely granulate on basal half. Declivity rather steep, broadly con- vex; striae not impressed, punctures very fine; interstriae smooth, punctures minute, confused. Vestiture of rather abundant, fine, moderately long hair.
Female.— Similar to male except frons weakly concave, a transverse callus just below middle, surface reticulate, finely, sparsely punc- tured, tubercles absent; lateral areas of prono- tum finely subasperate; strial punctures decrease in size only slightly toward declivity; interstrial granules moderately large to declivity; strial punctures on declivity slightiy larger, granules obsolete.
Type Locality.— Merida, Venezuela.
Type Material.- The male holotype, female allotype, and 56 paratypes were taken at the type locality on 22 October 1969, 1700 m eleva- tion, from rather old stumps of cut Japanese bamboo, by S. L. Wood. The habits were as in gracilis Wood.
Chramesus simplicis, n. sp.
This species evidently is similar to hylur- goides Schedl, but it is distinguished by the smaller size, by the more widely separated eyes
BwGHAM Young University Science Bulletin
in the male, by the absence of pronotal punc- tures, and by the very fine strial punctures.
Male.— Length 2.2 mm (paratypes 1.9-2.5 mm), 2.4 times as long as wide; color black.
Frons deeply, broadly concave from epi- stoma to upper level of eyes, lower fourth of lateral margin acute, not strongly elevated or dentate; surface minutely rugulose-rcticulate, not punctured; vestiture on lateral margins moder- ately abundant, fine, very long, in concavity very short, sparse. Antennal club tvpical of genus; moderately large; apex rounded.
Pronotum 0.91 times as long as wide; widest one-third length from base, sides rather strongly arcuate, moderately constricted just before rather broadly rounded anterior margin; surface strongly reticulate; all punctures replaced by narrow, small asperities over entire surface. Vestiture of rather abundant, fine, moderately long hair.
Elytra 1.7 times as long as wide, 2.0 times as long as pronotum; sides almost straight and parallel on basal two-thirds, rather broadly rounded behind; striae very feebly, broadly im- pressed, punctures minute; interstriae smooth, dull, four or more times wider than striae, punc- tures minute, a few on basal half finely granu- late, subasperate near base on interstriae 2-4. Declivity steep, rather narrowly convex; sculp- ture about as on posterior half of disc. Vestiture of moderately abundant, fine, rather long hair.
Female.— Similar to male except frons mod- erately convex, vestiture shorter, more evenly distributed; strial punctures larger, rather small; interstriae except 1, each with a row of rather fine granules to base of declivity; punctures of declivital striae smaller than on disc; interstrial granules largely obsolete.
Type Locality.— Carbonera Experimental Forest, about 50 km (airline) northwest of Mer- ida, Venezuela.
Type Material.— The male holotype, female allotype, and 3.3 paratypes were taken at the type locality on 12 January 1970, 2.500 m eleva- tion. No. 21.3, from native bamboo, by S. L. Wood. Tlie habits were as in gracilis Wood.
Male.— Length 1.7 mm (paratypes L6-L8 mm), 1.6 times as long as wide; color very dark brown, vestiture somewhat lighter.
Frons rather deeply concave from epistomal margin to just below upper level of eyes, then flattened from there to vertex; lateral margins acutely, rather strongly elevated from epistomal area to half distance between level of antennal insertion and upper margin of eye; entire sur- face strongly reticulate, punctures ver\' small, not clearly indicated; vestiture of fine, long hair, moderately abundant near marginal areas of upper half.
Pronotum 0.80 times as long as wide; es- sentially as in subopacus.
Elytra 1.0 times as long as wide; sides al- most straight and parallel on slightly less than basal half, broadly rounded behind; striae slight- ly impressed, punctures moderately large, dis- tinctly, shallowlv impressed; interstriae twice as wide as striae, weakly convex, almost smooth but not shining, punctures fine, abundant, con- fused. Declivity as in other species; sculpture as on disc. Vestiture of abundant, short, inter- strial scales in ground cover, each slightly longer than wide, some with acute points; and inter- strial rows of longer, erect scales, each about two and one-half times as long as ground cover, about six times as long as wide.
Female.— Similar to male except frons weak- ly convex, lateral margins not modified, vestiture short, coarse, unifonnlv distributed; anterolateral areas of pronotum moderately asperate; basal half of each discal interstriae armed by a row of small granules.
Type Locality.— Campamento Rio Grande, .30 km east of Palmar, Bolivar, Venezuela.
Type Material.— The male holotype, female allotype, and 71 paratypes were taken at the type locality on 12 June 1970, 200 m elevation. No. 568, from Bejuco Trinitario, by S. L. Wood. The host appeared very similar to, if not iden- tical with. CeJtis iguanae. Specimens were taken from a small, broken branch, from transverse parental tunnels.
The holotype, allot)'pe, and parat)'pes are in mv collection.
Chramcsus orinocensis, n. sp.
This species is allied to subopacus Schaeffer, but it is distinguished by the smaller size, by the less strongly, less extensively impressed male frons with the lateral margins less well devel- oped, and by the larger strial punctures.
Chramesus strigilis, n. sp.
This species evidently is allied to orinocensis Wood, but it is distinguished by the larger average size, by the lighter color, by the coarser vestiture and by the strongly, closely serrate lateral margias of the male frons.
Biological Series, Vol. 15, No. 3 New Neotropical Scolytidae
Male.— Length 2.0 mm (paratypes 1.9-2.1 mm), 1.5 times as long as wide; color brown.
Frons deeply, broadly concave from episto- mal margin to well above eyes; lateral margins acutely elevated from epistoma to level of an- tennal insertion, much more strongly elevated and closely, stronglv serrate from there to two- thirds distance to upper level of eyes; surface reticulate, a few fine, obscure punctures in- dicated; vestiture of fine, sparse, short hair.
Pronotum 0.74 times as long as wide; widest at base, sides arcuate and strongly converging to slight constriction just before broadly round- ed anterior margin; surface reticulate; postero- median area rather sparsely, coarsely, shallowly punctured, anteriorly and laterally from this point anterior margins of punctures become in- creasinglv asperate as punctures become smaller and more obscure, a few nonasperate punctures interspersed. Vestiture of rather long, slender scales.
Elytra 1.1 times as long as wide; sides al- most straight and parallel on basal half, broadly rounded behind; striae deeply, rather narrowly impressed, punctures rather small, moderately deep; interstriae about three times as wide as striae, rather strongly convex, almost smooth, punctures very fine, confused; a few pointed granules at bases of large, erect scales. Declivity moderately steep, convex; sculpture about as on disc; interstriae 1 weakly elevated. Vestiture of rather abundant interstrial ground scales, each slightly longer than wide; and interstrial rows of longer, erect scales, each twice as long as ground scales and four times as long as wide, spaced within a row by two or more times length of a scale.
Female.— Similar to male except frons weakly convex, shallowly foveate at center, lat- eral margins not elevated or otherwise modified; pronotal asperities larger but more nearly re- stricted to anterolateral areas; interstrial tuber- cles larger, verv widely separated posteriorly but extending to declivity at bases of major setae.
Type Locality.— Carbonera Experimental Forest, 50 km (airline) northwest of Merida, Venezuela.
Type Material— The male holotype, female allotv'pe, and 33 paratypes were taken at the type locality on 27 October 1969, 2.500 m eleva- tion, No. 89, from an unidentified cut seedling by S. L. Wood.
The holotype, allotype, and paratypes are in my collection.
Chramesus imporcatus, n. sp.
This species represents a species group new to me. The male frons bears the pair of tuber- cles on the lateral margins near the antennal insertions as in many other species, but it also bears a pair of sharply elevated carinae on the upper fourth of the lateral margins, their upper limits ending just below the upper level of the eyes. In this species the upper carinae are on the crest of the lateral margins.
Male.— Length 2.0 mm (paratypes 1.9-2.1 mm), 1.5 times as long as wide; color almost black.
Frons deeply, broadly excavated from epi- stomal margin to well above eyes; lateral mar- gins below level of antennal insertion strongly, acutely elevated, not pointed, margins above antennal insertion subacutely, weakly elevated then upper third strongly, acutely elevated to just below upper level of eyes; surface deeply reticulate, punctures sparse, small, obscure; vestiture of very fine, short, sparse hair. Anten- nal club large, typical of genus.
Pronotum 0.79 times as long as wide; widest one-third from base; sides strongly, arcuately converging to moderate constriction just before broadly rounded anterior margin; surface finely reticulate, punctures small, rather shallow, some in anterolateral area finely asperate on anterior margins. Vestiture of elongate, slender scales of moderate abundance.
Elytra 1.0 times as long as wide; sides almost straight and parallel on basal half, broadly rounded behind; striae strongly impressed, punctures rather small, deep; interstriae moder- ately convex, smooth, almost three times as wide as striae, a median row of narrow, rather high crenulations on each except confused on disc of 2 and part of 3, those in lateral areas more like flattened nodules, minute supplemental punc- tures also moderately abundant. Declivity rather steep, convex; sculpture about as on disc, gran- ules slightly smaller. Vestiture of very small ground scales, each about twice as long as wide, and longer, erect interstrial scales in rows ex- cept confused on 2 and 3 and toward base, each scale about three times as long as ground scales, about six times as long as wide, spaced within a row by length of a scale.
Female.— Similar to male except frons weak- ly convex, margins simple.
Type Locality'.— Seven km northwest of So- copo, Barinas, Venezuela.
Type Material.- The male holotype, female allotype, and nine paratypes were taken at the
Bricham Young University Science Bulletin
type locality on 13 February 1970, 200 m eleva- tion, No. 323, from a shrub known as Palito de Cruz, by S. L. Wood.
The holotype, allotype, and paratypes arc in my collection.
Chramesus impolitus, n. sp.
This species is very closely related to im- porcatus Wood, but it is distinguished by the smaller size, by the larger strial punctures, by the absence of ground vestiture on the elytra, and by the slight median displacement of the upper ridges on the male frons.
Male.— Length 1.8 mm (paratypes 1.6-2.0 mm), 1.6 times as long as wide; color almost black.
Frons as in imporcatus except upper carinae dLsplaced medially very slightly. Pronotum as in imporcatus except punctures more shallow.
Elytra as in imporcatus except striae less strongly impressed, punctures larger; interstriae twice as wide as striae except 2 wider; ground vestiture entirely absent, small interstrial punc- tures also absent.
Female.— Similar to male except frons weakly convex, margins simple.
Type Locality.— Campamento Rio Grande, 30 km east of Palmar, Bolivar, Venezuela.
Type Material.— The male holotype, female allotype, and 54 paratypes were taken at the type locality on 12 June 1970, 200 m elevation. No. 581 (some paratypes No. 567), from the shrub Rosa de Montana (presumably Brownia sp. ), by S. L. Wood.
The holotype, allotype, and paratypes are in my collection.
granulate; subglabrous. Antenna! club rather small for this genus.
Pronotum 0.85 times as long as wide; widest (me-third length from base, sides moderately arcuate on basal half, a slight constriction just before rather broadly rounded anterior margin; surface reticulate; punctures coarse, shallow, not close. Vestiture restricted to a few coarse setae in lateral and anterior areas.
Elvtra 1.2 times as long as wide, 1.5 times as long as pronotum; sides almost straight and parallel on slightly more than basal half, broadly rounded behind; striae not impressed, punctures coarse, rather deep; interstriae smooth, shining, slightly wider than striae, punctures replaced by moderately large, uniseriate, setiferous granules. Declivity rather steep, convex; striae and inter- striae narrower than on disc, strial punctures and interstrial granules distinctly smaller. Vesti- ture of interstrial rows of erect bristles; each bristle slightly wider near its tip, spaced within a row by distances slightly less than length of bristle, between rows by distances slightly greater than length of a bristle.
Female.— Similar to male except frons flat- tened on lower third, foveate at center, lateral margins unmodified.
Type Locality'.— Rancho Grande, Aragua, Venezuela.
Type Material.- The male holotype, female allotype, and 27 paratypes were taken at the type locality on 9 April 1970, 1100 m elevation. No. 426, from a Tahehuia twig, by S. L. Wood. The biramose parental tunnels were broadly V-shaped, with both egg tunnels diagonal.
The holotvpe, allotype, and paratypes are in my collection.
Chramesus parens, n. sp.
This species differs from all other represen- tatives of the genus known to me by the small size, by the total absence of ground vestiture on the elytra, by the absence of pronotal as- perities, and by the presence of a pair of tu- bercles arming the male frons as in most other species of the genus.
Male.— Length 1.4 mm (paratypes 1.3-1.4 mm), 1.9 times as long as wide; color rather dark brown, elytra slightly lighter.
Froas rather strongly concave from epistoma to just below upper level of eyes; lateral margins subacutely elevated on lower half, armed at level of antennal insertion by a pair of coarse, pointed denticles; surface very finely reticulatc-
Chramesus denticulatus, n. sp.
This species is unique in the genus. It is distinguished from all known species by the small size, by the unamied male frons, by the finely asperate anterolateral areas of the prono- tum, and by the finely dentate declivital inter- striae.
Male.— Length 1.3 mm (paratypes 1.2-1.3 mm), 1.9 times as long as wide; color yellowish brown.
Frons rather deeply concave on a subcircular area from epistoma to upper level of eyes, lateral margins subacute, unarmed; epistomal margin carinatelv elevated, more strongly on median third; surface reticulate, punctures minute, ob- scure; vestiture uniformly short, coarse, uni-
Biological Series, V'ol. 15, No. 3 New Neotropical Scolytidae
formly distributed setae. Antennal club rather small for this genus.
Pronotum 0.77 times as long as wide; widest near base, sides convergently arcuate to broadly rounded anterior margin; surface reticulate, al- most unifonnly covered by isolated, rather wide- ly spaced, small subasperate tubercles. Vesti- ture short, of almost equal numbers of very fine hair and rather broad scales.
Elytra 1.3 times as long as wide, 1.7 times as long as pronotum; sides almost straight and parallel on slightly more than basal half, broad- ly rounded behind, posterior outline interrupted by declivital teeth; striae moderately impressed, punctures rather coarse, shallow; interstriae sub- reticulate, weakly convex, as wide as striae; punctures replaced by rows of granules, very fine at base, becoming coarse toward declivity. Declivity rather steep, broadly convex; striae narrower and deeper than on disc; interstriae more narrowly con\'ex, each armed by a row of close, rather small pointed teeth, except al- most obsolete on lower half of 2. Vestiture of ground cover formed by two indefinite rows of short, stout setae on margins of each interstriae, and a central row of short, erect scales; each scale about twice as long as wide, spaced within and between rows by slightly more than twice length of a scale; scales about twice as long as ground setae.
Female.— Similar to male except frons weak- ly concave, epistoma weakly elevated, frontal vestiture of short scales; pronotal asperities larger in anterolateral areas; interstrial teeth on declivity slightly smaller, regularly present on 2.
Type LocALiTi.— Twenty km southwest of El Vigia, Merida, Venezuela.
Type Material.— The male holotype, female allotype, and three paratypes were taken at the type locality on 21 November 1969, 50 m eleva- tion, No. 150, from a small stem of a large, cut Bignoniaceae vine (liana), by S. L. Wood.
The holotvpe, allotype, and paratypes are in my collection.
Chramesus prisons, n. sp.
This species remotely resembles denticulatus Wood, but the relationship is remote. It is dis- tinguished by the unanned male frons that has the epistomal margin transversely carinate, by the feebly asperate anterolateral areas of the pronotum, by the different vestiture, and by the minute dechvital granules.
Male.— Length 2.2 mm (paratypes 2.0-2.2 mm), 1.9 times as long as wide; color brown,
elytra slightly lighter.
Frons rather deeply, broadly concave from epistoma to slightly above eyes; median half of epistoma acutely, transversely carinate, a pair of minute denticles at ends of carina; lateral mar- gins subacute, the fourth immediately above level of antennal insertion thicker, more strongly elevated, not dentate; surface reticulate, punc- tures very fine, sparse; vestiture fine, long, rather sparse. Antennal club more slender and more sparsely pubescent than in most species.
Pronotum 0.73 times as long as wide; widest at base, sides rather strongly, arcuately converg- ing to slight constriction just before broadly rounded anterior margin; surface strongly reticu- late, punctures replaced by fine, sparse granules, finely asperate in lateral areas. Vestiture of fine and stout hairlike setae of moderate abundance.
Elytra 1.3 times as long as wide, 2.0 times as long as pronotum; sides almost straight on basal half, slightly wider near base of declivity, broadly rounded behind; striae weakly im- pressed, punctures moderately large, rather deep; interstriae twice as wide as striae, weakly convex, minutely irregular, shining, punctures replaced by rows of fine granules, punctures giving rise to ground vestiture mostly too small to see at 80 diameters magnification. Declivity rather steep, convex; striae slightly deeper, inter- striae more convex than on disc; interstrial granules rather small, close. Vestiture short, rather coarse, moderately abundant; central row on each interstriae slightly longer and coarser.
Female.— Similar to male except frons weak- ly convex, weakly, transversely impressed above epistoma, a central fovea present; anterolateral pronotal asperities much larger.
Type Locality.— Carbonera Experimental Forest, about 50 km (airline) northwest of Merida, Venezuela.
Type Material.— The male holotype, female allotype, and nine paratypes were taken at the type locality on 9 December 1969, 2500 m eleva- tion. No. 175, from dead mistletoe (possibly Phoradendwn) , by S. L. Wood.
The holotype, allotype, and paratypes are in my collection.
Chramesus vinealis, n. sp.
This species is very closely allied to orino- censis Wood, but it is distinguished as indicated below in the description.
Male.— Length 1.9 mm (paratypes 1.7-2.0 mm), 1.6 times as long as wide; color very dark brown, almost black.
Bricham Young Univlhsitv Science Bulletin
Frons as in orinocensis except acutely ele- vated part of lateral margin more strongly ele- vated on its lower half and less strongly elevated on its upper half. Pronotuni as in orinocensis except punctures distinctly larger, slightly deep- er, asperities in lateral areas larger, more numerous.
Elytra as in orinocensis except striae slightly more strongly impressed, punctures on 1-3 obso- lete, others smaller, not as deep; declivital inter- striae each with a row of widely spaced fine granules; interstrial ground scales much smaller; erect scales as in orinocensis.
Female.— Similar to male except frons con- vex, margins simple; striae 1-3 with punctures clearly indicated; interstriae each with a row of fine granules on both disc and declivity.
Type Locality.— Thirty km north of Caiion Zancudo, Zulia, Venezuela.
Type Material.— The male holotype, female allotype, and 136 paratypes were collected at the type locality on 4 June 1970, 10 m eleva- tion, No. 522, from an unidentified vine (liana), by S. L. Wood.
The holotype, allotype, and paratypes are in my collection.
Chraniesus soUcitatus, n. sp.
This species might possibly be allied to rotundatus Chapuis, but it is distinguished by the smaller size, by the larger tuft of hair on the scape, by the much more deeply excavated male frons, and by other characters.
Male.— Length 1.6 mm (paratypes 1.4-1.6 mm), 1.6 times as long as wide; color brown.
Frons shallowly, broadly concave from epi- stomal margin to well below upper level of eyes; middle third of lateral margins distinctly, subacutely, not strongly elevated, indistinctly serrate; surface deeply reticulate above, irregu- larly, indistinctly rugose-reticulate below; vesti- ture inconspicuous. Antennal scape elongate, apex slightly produced and ornamented by about a dozen hairlike setae equal in length to scape; club mmlerately small for this genus.
Pronotum 0.73 times as long as wide; widest at base, sides rather strongly, arcuately conver- gent to broadly rounded anterior margin; surface reticulate, fine and moderately coarse, shallow punctures intermixed; anterolateral areas with a few irregularities, not clearly asperate. Vesti- ture of moderately abundant, slender scales.
Elytra 1.0 times as long as wide; sides straight and parallel on less than basal half, broadly rounded behind; striae distinctly im-
pressed, punctures moderately coarse, rather shallow; interstriae weakly convex, almost smooth, dull, almost twice as wide as striae, <>ach with a row of rather coarse granules and minute, confused punctures. Declivity rather steep, broadly convex; striae more narrowly impressed, punctures smaller, less distinct; interstriae nar- rower, less convex, granules smaller and more widely spaced. Vestiture of small, abundant ground scales, each slightly longer than wide, and interstrial rows of long scales, each almost three times as long as ground scales, six to eight times as long as wide, about half as long as dis- tance between scales within a row or between rows.
Female.— Similar to male except frons con- vex, a transverse callus at level of antennal inser- tion, central fovea indistinct; scape not orna- mented by hair; four or five rather coarse asperi- ties in anterolateral area of pronotum; scales in ground cover of interstriae slender, up to four times as long as wide.
Type Locality.— Campamento Rio Grande, 30 km east of Palmar, Bolivar, Venezuela.
Type Materlvl.— The male holotype, female allotype, and 13 paratypes were taken at the type locality on 12 June 1970, 200 m elevation. No. 569, from an unidentified vine (liana), by S. L. Wood. The parental galleries were oblique to longitudinal.
The holotvpe, allotype, and paratypes are in my collection.
Chramesus peniculus, n. sp.
This species is allied to rotundatus Chapuis, but it is distinguished by the much smaller size, and by the very long interstrial bristles.
Male.— Length 1.2 mm (paratypes 1.2-1.3 mm), 1.5 times as long as wide; color dark brown with pale scales.
Frons rather deeplv, broadiv concave from epistomal margin to slightly above eyes; lateral margins subacutely elevated, with a rather stronglv elevated, subcjuadrate denticle on mar- gin slightly above level of antennal insertion; surface strongly reticulate, punctures not evi- dent; vestiture sparse, inconspicuous. Antennal scape ornamented at apex by a tuft of perhaps a dozen long hairlike setae as long as scape; club rather small for this genus.
Pronotum 0.70 times as long as wide; widest near base, sides strongly arcuate, converging to moderate constriction just before rather broadly rounded anterior margin; surface strongly reticu- late, punctures moderately close, small and
Biological Series, Vol. 15, No. 3 New Neotoopic.al Scolythme
rather large inteniiixcd, becoming finely then rather coarsely a.sperate toward anterolateral areas. Vestiture of moderately abundant, slen- der and stout, short, scmirccumbent bristles.
Elytra 0.90 times as long as wide; sides straight and parallel on less than basal half, verv broadly rounded behind; 13 crenulations on basal margin, submarginal row extending to striae 5; striae rather strongly impressed, punc- tures small, rather shallow; interstriae slightly wider than striae, evidently weakly convex, de- void of granules, punctures very small. De- clivitv' beginning at middle, moderately steep, broadly con\ex; sculpture about as on disc ex- cept striae and interstriae somewhat narrower. Wstiture of ground cover of abundant, short, interstrial scales, each scale about as long as wide; and erect, flattened bristles about four times as long as ground cover, spaced between rows bv distances about equal to length of a bristle, within a row h\ one to three times length of a bristle. Broad strial grooves smooth, except for punctures, and glabrous.
Female.— Similar to male except frons weak- ly convex, a weak, transverse callus just above level of antennal insertion, an obscure fovea at center; scape not ornamented by hair; antero- lateral crenulations on pronotum slightly larger; interstrial bristles wider, more nearly scalelike on basal half of elytra.
Type Locality.— Thirty km north of Canon Zancudo, Zulia, Venezuela.
Type Materlvl.- The male holotype, female allotype, and 14 paratypes were taken at the type locality on 4 June 1970, 10 m elevation. No. 522, from the same unidentified vine ( liana ) that contained vinealis, by S. L. Wood.
The holotype, allotype, and paratypes are in mv collection.
Chaetophloeus andinus, n. sp.
This is the second species of Chaetophloeus known to occur in South America. It is allied to brasilicmis Blackman, but it may be distin- guLshed bv the larger size, by the more deeply emarginate epistoma, by the more deeply exca- vated and elaborately ornamented male frons, and by the longer, erect, interstrial setae.
Male.— Length 1.9 mm (paratypes 1..3-1.8 mm), 1.5 times as long as wide; color very dark bro\\ii, with pale and light brown vestiture.
Frons deeply concave to inner margin of eye from epistomal margin to vertex; epistomal mar- gin ver\' deeply, broadly emarginate, less than three times as wide as deep, a pair of marginal
tubercles directed orad near median line; lateral and upper margins ornamented by a dense brush of long hau-, tips of some setae on vertex extend beyond epistomal margin. Antennal club large, 1.9 times as long as wide; three sutures weakly procurved, suture 3 two-thirds club length from base.
Pronotum as in hra.siUensis, with surface sculpture finer, one paired group of three tu- bercles in anterolateral areas; setae of two types, some slender, short, others scales one and one-half to two times as long as wide except three times as long on anterior margin and in small area near scutellum.
Elytra as in brasiliensis except strial punc- tures slightly smaller; interstriae about three times as wide as striae; erect interstrial scales more numerous, in less definite rows, each erect scale about twice as long as wide on disc, up to five times as long as wide on declivity.
Female.— Similar to male except frons weak- ly convex; epistomal emargination more than four times as wide as deep; frontal vestiture much shorter, unifonnly distributed; pronotum with two paired groups of tubercles.
Type Locality.— Three km east of Lagunil- las, Merida, Venezuela.
Type Material.— The male holotype, female allotype, and 26 paratypes were taken at the type locality on 12 January 1970, 1000 m eleva- tion. No. Z37, from Mimosa twigs and No. 236 and 2.39 from an unidentified vine, by S. L. Wood.
The holotype, allotype, and paratypes are in my collection.
Liparthriim carapae, n. sp.
This is the first record of this genus from South America. As in americamim Wood the male frons of this species is broadly concave, but it differs by the restriction of frontal pubes- cence to the vertex, by the much finer, less abundant vestiture on pronotum and elytra, by the obsolescent strial punctures, and by the larger size.
Male.— Length 1.1 mm (paratypes 0.8-1.2 mm), 2.4 times as long as wide; color dark red- dish brown.
Frons rather deeply concave from eye to eye from epistomal margin to vertex; surface shin- ing, punctures very minute, rather abundant; concavity apparently glabrous, upper margin ornamented by a brush of long hair, tips of some setae reaching middle of concavity. An-
10
Bricham Young University Science Bulletin
tennal club rather largp, broadly oval, devoid of sutures.
FronoUiin 1.0 times as long as wide; sides subparallel, feebly arcuate on basal half, rather broadly rounded in front; moderately declivous on anterior fourth; surface subreticulate, with minute, isolated asperities on median area al- most to base, punctures not evident. Vestiture of fine, recumbent hair and a few erect, slender scales.
Elytra 1.4 times as long as wide, 1.4 times as long as pronotum; sides almost straight and parallel on basal two-thirds, rather broadly rounded behind; each basal margin armed by six crenulations; striae not impressed, punctures almost totally obsolete, a few feebly impressed near center of disc; interstriae minutely irregu- lar, uniseriately granulate. Declivity steep, con- vex; as on disc. Vestiture consisting of very fine, rather short, recumbent, strial hair; and rows of erect, interstrial scales, each scale about half as long as distance between rows, each about three times as hmg as wide on disc, on declivity al- most as wide as long.
Female.— Similar to male except frons con- vex, minutely reticulate-granulate, glabrous; an- terior margin of pronotum armed bv eight teeth; pronotal asperities much larger; striae more distinctly impressed on declivity.
Type Locality.— Campamento Rio Grande, 30 km east of Palmar, Bolivar, Venezuela.
Type Material.— The male holotype, female allotype, and 230 paratypes were collected at the type locality on 12 June 1970, 200 m eleva- tion. No. 572, Carapa gtiiatiemis, S. L. Wood. The beetles infested the phloem of branches and bole of the host. The galleries were typical of the genus.
The holotype, allotype, and paratypes are in my collection.
Liparthniin meridensis, n. sp.
This species has the concave male frons of americanum Wood, but it may be distinguished from other species of this group by the elevated interstriae 3 and by the absence of scales on declivital interstriae 2.
MALE.-Length 0.8 mm (paratypes 0.75-0.85 mm), 2.2 times as long as wide; color very dark brown.
Frons concave from eye to eye, from cpi- stoma to upper level of eyes; surface reticulate, punctures minute, rather abundant; vestiture moderately abundant laterally and above, rather abundant and apparently part of setae remark- ably spatulate on their distal halves.
Pronotum 0.86 times as long as wide; widest near base, sides rather strongly arcuate on basal half, narrowly rounded in front; surface subre- ticulate, with rather abundant, isolated granules or minute asperities on median two-thirds to base.
Elytra 1.4 times as long as wide, 1.6 times as long as pronotum; basal margins armed by six pairs of crenulations; striae not impressed, punc- tures small, moderately deep; interstriae as wide as striae, smooth, uniseriate punctures feebly vulcanate. Declivity steep, convex; interstriae 1 and 3 distinctly elevated, 2 flat. Vestiture of rather fine, short, recumbent strial and inter- strial hair, and interstrial rows of erect scales; each scale about twice as long as wide; declivital interstriae 2 and lower half of 4 devoid of scales.
Female.— Similar to male except frons con- vex, reticulate, with a few obscure punctures; anterior margin of pronotum armed by four teeth; anterior slope of pronotum armed by about a dozen coarse asperities; strial punctures about twice as large, deep; declivital interstriae 1 and 3 more strongly elevated, about a fourth as high as wide.
Type Locality.— Five km east Lagunillas, Merida, Venezuela.
Type Material.— The male holotype, female allotype, and 71 paratypes were collected at the type locality on 12 January 1970, 1000 m eleva- tion. No. 238, from a composite shrub, by S. L. Wood,
The holotype, allotype, and paratypes are in my collection.
Cladoctonus boliviae (Wood)
Hoplitophthonis boliviae Wood, 1961, Gt. Ba.sin Nat. 21:106 (Holotype, female; Route liehveen Boyiiilbe and Charagiia via Ciieva, Ingri, etc., Bolivia; U.S.
Nat. Mus.).
A long series of this species was taken at Campo Capote, 27 km northeast of Montoya, Santander, Colombia, 2-VII-70, 150 m elevation. No. 601, from PseucholmecUa, by me. The ap- parently monogamous beetles entered the bark through tunnels made by an equally small Phloeotribtis. In the cambium region they ap- propriated one or both of the Phloeotribtis egg galleries and extended or branched from it in constructing their owii tunnels that engraved both phloem and xylem. The beetles worked more or less in pairs along one branch of the tunnel while other pairs constructed their tun- nels as branches of the tunnel of their com- p;mions. Usually about two to four branches were fomied from one original Cladoctonus
BtOLOCiCAL Series, Vol. 15, No. 3 New Neotoopical Scolytidae
H
entry. Larval mines were in the phloem, but were completely confused with those of the Phlocotribits. These tunnels were almost identi- cal to those of Carpliodicticus crwtatus Wood.
Pycnarthrum iiionuitmn, n. sp.
This species, at least superficially, is very similar to hisj)ic1tnn (Ferrari) except that all lontj setae or bristles are absent from the entire liodv; the vestiture consists only of the very short ground cover of scales.
Female.— Length 1.6 mm, about 2.2 times as long as wide (elytra spread slightly); color ver)' dark brown, elytra lighter.
Frons weaklv convex except indistinctly con- cave on central third; eyes separated above by distance greater than width of an eye; surface reticulate, punctures moderately coarse, rather close, sharply, not deeply impressed: vestiture of minute hair and scales, each not longer than twice diameter of a puncture. Antennal club and flagellum missing on type.
Pronotiim about ecjual in length and width; widest at middle, sides weakly arcuate, broadly rounded in front; surface shining, subreticulate on anterior surface, with minute points mod- erately abundant, punctures moderately coarse, deep, close, interspaces not wider than distance equal to diameter of a puncture. Vestiture in- conspicuous, consisting of short, slender hair and on anterior half some equally short, stout setae intennixed.
Elytra about 1.4 times as long as wide; sides almost straight and parallel on basal two-thirds, rather broadlv rounded behind; striae 1 mod- eratelv, others not impressed, punctures coarse, deep, rather close; interstriae as wide as striae, subshining, very obscurely subreticulate, punc- tures small, moderately abundant, confused. De- clivitv steep, broadly convex; interstriae 2 some- what impressed, narrower than others. Vestiture consisting of minute strial hair on disc only, and short, confused, subplumose, narrow, interstrial scales equally abundant on disc and declivity; rows of erect bristles found in other species of this genus absent.
Type Locality.— Olanchito, Yoro, Honduras.
Type Material.— The female holotype was collected at the tvpe locality on 7 October 1949, at light, by E. C. Becker.
The holotype is in my collection.
Pycnarthrum funerium. n. sp.
This species is allied to hispidum (Ferrari) but may be distinguished by the very small
size, by the black color, by the small, shallowly impressed stri;d punctures, and by the more slender, less plumose setae in the clytral ground vestiture.
Female.— Length 1.4 mm (paratypes 1.. 3-1.4 mm), 2.2 times as long as wide; color black.
Frons rather strongly, evenly convex to epi- stoma; surface reticulate, becoming subrugulose below, punctures fine, shallow, obscure; eyes separated by distance greater than width of an eye; vestiture of rather sparse, stout, erect, moderately long, white bristles. Antennal club 2.0 times as long as wide, two sutures indicated.
Pronotum 0.9.3 times as long as wide; widest at or slightly in front of middle, sides weakly arcuate, very broadly rounded in front; surface strongly reticulate on anterior third, obscurely reticulate behind, subshining, punctures small, close, moderately deep. Vestiture of fine hair and stout bristles intermixed over entire surface, moderately long.
Elytra about 1.4 times as long as wide (elytra spread slightly); sides almost straight and paral- lel on basal tsvo-thirds, rather broadly rounded behind; striae not impressed, punctures rather small, distinct, very shallow; interstriae almost twice as wide as striae, evidently obscurely sub- reticulate, median row of punctures uniseriate, very small but almost twice as large as confused supplementary punctures. Declivity rather steep, rather narrowly convex, not at all im- pressed; essentially as on disc, except strial punc- tures very slightly deeper. Vestiture consisting of short, interstrial ground vestiture of rather fine (anteriorly) to stout (posteriorly) semi- recumbent hairlike bristles, a few exhibit indi- cations of being subplumose, and longer, erect, uniseriate rows of stout, almost scalelike setae, each erect bristle slightly less than twice as long as ground vestiture and about four to six times as long as wide; not longer on declivity.
Type Locality.— La Ceiba, Atlantida, Hon- duras.
Type Material.— The female holotype was taken at the type locality 10 June 1949, at light, by E. C. Becker. Three female paratypes bear identical data except one was taken on 27 May 1949 and two on 20 May 1949.
The holotype and paratypes are in my collec- tion.
Pycnarthrum fici, n. sp.
This species superficially resembles hispidum (Ferrari) but may be distinguished by the deeply concave male frons, by the pale yellow mature body color, by the more narrowly convex
12
Bhicham Young Univebsity Science Bulletin
elytral declivity, and by the shorter elytral ves- titure.
Male.— Length 1.7 mm (paratypes 1.6-1.7 mm), 2.2 times a.s long a.s wide; color pale yellow.
Frons rather deeply concave from upper level of eyes to cpistomal margin; surface reticu- late, finely granulate; eyes rather narrowly sepa- rated above, separated by distance ecjual to 1..5 times width of an eye; vestiture of rather fine, short, moderately abundant setae. Antennal club 1.6 times as long as wide; sutures 1 and 2 distinct, .3 obscurelv indicated.
Pronotum proportions and outline as in hisi)idum; surface reticulate on anterior third, obscurely subreticulate behind, punctures uni- formly rather small, shallow, close. Vestiture very short, of fine and stout setae intennixed.
Elytra 1.4 times as long as wide; sides almost straight and parallel on basal two-thirds, rather narrowly rounded behind; striae weakly im- pressed, punctures very shallow, obscurely im- pressed, of moderate size, interstriae about one and one-half times as wide as striae, surface not regular, perhaps very minutely rugulose, punc- tures minute, many very minutely elevated par- ticularly along median row. Declivity rather steep, convex; striae and interstriae slightly nar- rower than on disc, striae more distinctly im- pressed, punctures less distinct. Vestiture con- sisting of short, stout, moderately abundant ground cover mostly in indefinite marginal rows on each interstriae, not subplumose, and inter- strial rows of erect, stout bristles, each bristle about one and one-third times as long as ground setae, very slightly shorter in length than width of an interstriae, at least six times as long as wide.
Female.— Similar to male except frons mod- erately convex from upper level of eyes to near epistoma; surface very finely subrugulose.
Type Locality.— Olanchito, Honduras.
Type Material.— Tlie male holotypc and four paratypes were collected at the type locality on 21 June (type), 15 and 19 May 1949, at light, by E. C. Becker. The female allotype and 40 paratypes were taken at La Ceiba, Honduras, on various dates from May to July 1949, at light, by E. C. Becker. Eighty-seven paratypes were taken at 5 km west of El Pino ( near southeast- ern shore of Lake Maracaibo), Merida, Vene- zuela, on 20 October 1969, 10 m elevation. No. 143, from strangler fig, by S. L. Wood. The gallerv svstems were basically as in Jus-pkhim.
The holotype, allotype, and paratypes are in mv collection.
Ptjcmirtlirum perdiitim, n. sp.
This species and lucidum Wood have the setae on the basal half of the elytra much more slender and the erect bristles longer than in other representatives of the genus. From lucidum this species is distinguished by the larger, more widely spaced pronotal punctures, by the more abrupt slope of the elytral declivity, and by the less strongly impressed male frons.
Male.— Length 1..3 mm (allotype 1.6 mm), 2.3 times as long as wide; color dark brown, elytra somewhat lighter.
Frons weakly convex except flattened in median area on lower half; surface reticulate, rather finely, shallowly punctured, a conspicu- ous median fovea at center; eyes separated above by 1.4 times width of an eye. Anteimal club 2.0 times as long as wide; sutures 1 and 2 indicated.
Pronotum proportions, outline and sculpture as in hispidum except punctures uniformly rather large, deep, not as close; vestiture of fine hair except intennixed with stouter setae on margins and on anterior fifth.
Elytra 1.5 times as long as wide; sides almost straight and parallel on basal tsvo-thirds, rather narrowly rounded behind; striae 1 feebly, others not impressed, punctures moderately large, rather deep; interstriae as wide as striae, punc- tures of two sizes, a median row of small punc- tures and less abundant, very minute punctures on margins of interstriae. Declivity rather steep, convex; striae distinctly impressed, narrower than on disc; interstriae weakly convex. Vesti- ture of fine strial hair and e(jually fine, short, recumbent, sparse, interstrial hair, and inter- strial rows of longer, erect, slender bristles, each bristle one and one-half times as long as ground vestiture, ecjual in length to width of an inter- striae, stout, more than six times as long as wide.
Female.— Similar to male except frons more stronglv convex to epistoma, interstrial bristles more slender.
Type Locality.— La Ceiba, Adantida, Hon- duras.
Type Material.— The male holotype and female allotype were taken at the type locality on 26 August 1949, at light, by E. C'. Becker.
The holotype and allotype are in my collec- tion.
Pycnartlirum lucidum. n. sp.
This species is allied to perditum Wood, but mav be distinguislied bv the concave male frons, by the closer, finer, pronotal punctures, by the less steep elytral declivit\', by the larger, deeper
Biological Series, Vol. 15, No. 3 New Neotropical Scolytiuae
13
strial punctures, and by the less abundant, finer pronotal and elytral vestiture.
Male.— Length 1.7 mm (paratypes 1.5-1.9 mm), 2.3 times as long as wide; color brown, elytra lighter.
Frons moderately concave from upper level of eyes to epistoma, a shining, transverse callus on median half of epistoma, a pair of calli in lateral areas of concavity just above level of antemial insertion; surface shining, obscurely reticulate; punctures rather fine, deep, rather close; eyes separated by distance equal to width of an eye. Antennal club 1.6 times as long as wide, sutures 1 and 2 distinct, 3 obscurely indi- cated.
Pronotum proportions and outline as in per- ditum; surface brightlv shining except obscurely reticulate on anterior third; minute points pres- ent, punctures rather small, deep, close. Vesti- ture mostly abraded, verv fine, a few coarse setae in anterolateral areas.
Elytra 1.4 times as long as wide; sides straight and parallel on basal two-thirds, rather narrowly rounded behind; striae weakly im- pressed, punctures coarse, close, moderately deep; interstriac slightlv wider than striae, al- most smooth, shining, punctures very fine, in three obscure ranks, median row very slightly larger. Declivitv rather steep, convex; striae narrowlv impressed, narrower than on disc; in- terstriae twice as wide as striae, rather weakly convex. Vestiture consisting of fine, short strial hair and interstrial short ground cover of very fine hair on disc becoming scalelike on declivity, and interstrial rows of erect bristles, each bristle almost twice as long as ground cover, equal in length to width of an interstriae. about six to eight times as long as wide.
Female.— Similar to male except frons rather stronglv convex from upper level of eyes to epistoma.
Type Locality.— Finca Gromaco on Rio Goto Brus, Puntarenas, Gosta Rica.
Type Material.— The male holotype and four paratypes were collected at the tvpe locali- ty on 14 July 1963, .500 m elevation, No. 60, from a broken limb, by S. L. Wood. The allo- type and 63 female paratypes were taken at Playon, San Jose, Gosta Rica, 20 m elevation. No. 117, in Cedro amarga, by S. L. Wood.
The holot}'pe, allotype, and paratypes are in my collection.
Piicnarthnim carinatum, n. sp.
This species is verv similar to fici Wood, but it may be distinguished by the sharply elevated.
transverse epistomal carina of the male, and by the longer, very stout interstrial bristles.
Male.— Length 1.8 mm, 2.1 times as long as wide; color very dark brown, elytra lighter.
Frons deeply concave from just below upper level of eyes to epistoma as in jici; epistoma armed by an acnately elevated, transverse carina on median fourth; eyes separated above by 2.2 times width of an eye. Other features of head and pronotum essentially as in jici.
Elytra 1.25 times as long as wide; outline as in jici; striae weakly impressed, punctures rather small, distinctly not deeply impressed, not at all confluent or rugulose; interstriae shining, almost twice as wide as striae, median row of punctures fine, uniseriate, punctures on margins of each interstriae minute. Declivity rather steep, con- vex; strial punctures deeper than on disc, very close, shining, not at all confluent or reticulate. Vestiture of fine, short, recumbent strial hair and rows of short, rather slender, subplumose scales on each margin of each interstriae; and inter- strial rows of erect very stout bristles, each bristle slightlv shorter than width of an inter- striae, each about four times as long as wide.
Type Locality.— Near Leonpampa, Depart- ment Huanuco, Peru.
Ty'PE Material.— The unique male holotype was collected at the type locality between 11 and 30 December 1937, 800 m elevation, in jungle. No. 3811, by F. Woytkowski.
The holotype is in my collection.
Pijcnarthrum subcarinahnn, n. sp.
This species is very closely related to carina- tum Wood except as noted below.
Male.— Length 1.9 mm (paratypes 1.6-1.9 mm), 2.1 times as long as wide; color very dark brown, vestiture pale.
Frons as in carinatum except epistomal ca- rina weaklv developed, shorter, not acutely pro- duced. Pronotum as in carinatum except vesti- ture finer. Elytra as in carinatum except erect interstrial setae slightly longer and much more slender, each bristle six or more times as long as wide.
Female.— Similar to male except frons con- vex, epistomal elevation not indicated.
Type Locality.— Eight km southwest of Bumbum, Barinas, Venezuela.
Type Material.— The holotype, allotype, and
10 paratypes were taken at the type locality on
11 February 1970, 150 m elevation, No. 327,
14
Brigham Young University Science Bulletin
from Charo Blanco (Brosmium sp. ), by S. L. Wood.
Pijcnarthnim hrosmii, n. sp.
This species is allied to lucidum Wood, but it may be distinguished by the much liner pro- notal and elytral discal punctures, by the more strongly impressed elytral declivity, and by the flattened male frons.
Male.— Length 2.0 mm (paratypes 1.9-2.1 mm), 2.3 times as long as wide; color reddish brown.
Frons flattened from just below upper level of eyes to epistoma; surface shining, obscurely reticulate, punctures fine, moderately close; ves- titure rather sparse, fine, erect, moderately long; eyes very large, coarsely faceted, separated above by a distance ecjual to width of an eye. Antennal club 1.8 times as long as wide, sutures
1 and 2 indicated.
Pronotum equally as long as wide; widest in front of middle, broadly rounded in front; sur- face shining, except subreticulate on anterior third, a few minute points, punctures fine, rather deep, moderately close; vestiture of fine, short bristles, a few longer setae in marginal areas.
Elytra 1.3 times as long as wide; sides almost straight and parallel on basal two-thirds, rather broadly rounded behind; striae 1 slightly, others feebly or not at all impressed, punctures rather small, moderately close; interstriae twice as wide as striae, almost smooth, median row of punc- tures uniseriate, fine, becoming finely then rather conspicuously granulate in progressing from anterior to posterior areas, supplementary punctures on margins of each interstriae minute, irregularly spaced. Declivity rather steep, broadly flattened; strial punctures conspicuously larger and deeper than on disc; interstriae slightly wider than striae, 1 moderately elevated,
2 distinctly impressed, all interstriae with upper half uniseriately, rather finely granulate. Vesti- ture consisting of ground cover of fine, short, moderately abundant strial and interstrial hair, strial setae absent on declivity, interstrial ground cover on declivity of subplumose scales; and interstrial rows of erect bristles, each bristle slightly more than twice as long as ground cover, about equal in length to width of an interstriae, each at least eight times as long as wide.
Female.— Similar to male except frons moderately convex from upper level of eyes to epistoma.
Type Locality.- Nine km south of Barrancas, Barinas, Venezuela.
Host.— Brosmium sp. (Charo amarillo).
Type Materl\^l.— The male holotype, female allotype, and six paratypes were collected at the type locality on 1 October 1969, 150 m ele- vation, No. 23, from Charo amarillo, by S. L. Wood; 12 paratypes bear identical data except 5 November 1969, No. 106; two paratypes are labeled 8 km SW Bumbum, Barinas, Venezuela, 11-11-70, No. 319; two paratypes 10 km SE Miri, Barinas, Venezuela, 8-II-70, No. 299; six para- types 27 km NE Montoya, Santander, Colombia, 2-VII-70, No, 591; and 24 paratypes 8 km S Colonia (near Buenaventura), Valle de Cauca, Colombia, 9-VII-70, No. 645, all from the same host and collector.
Gymnochihis alni, n. sp.
This species is rather similar to reitteri Eichhoff, but may be distinguislied by the concave male frons, by the elongate body, by the absence or near absence of sutures on the antennal club, and by the obsolescent strial punctures on the disc.
Male.— Length 2.2 mm (paratypes 2.0-2.2 mm), 2.2 times as long as wide; color light brown to bicolored.
Frons rather shallowlv, broadlv concave from well above eyes to epistoma; surface strongly reticulate, a transverse arcuate callus at and slightly above level of antennal insertion, punc- tures fine, shallow, moderatelv abundant; vesti- ture fine, very short, sparse. Antennal club 1.3 times as long as wide, sutures not clearly e\ident except basal portion slightly sclerotized to point where suture 1 normally located.
Pronotum 0.S6 times as long as wide; outline, sculpture and vestiture as in reitteri except pos- terior area finelv, densclv punctiued.
Elytra 1.3 times as long as wide; sides almost straight and parallel on more than basal half, narrowly rounded behind; striae 1 distinctly, others feebly or not at all impressed, punctiires small, scarceJN' distinguishable from those of interstriae; interstriae three to four times as wide as striae, obscurely subreticulate, rather finely, densely punctured. Declivity commencing at middle, moderately steep, convex; strial punc- tures small, distinct; interstriae 1 weakly ele- vated. Wstiture of short, recumbent, very abundant, fine hair; and sparse interstrial rows of erect bristles, those on even-numbered inter- striae onl\- slightlv longer than ground vestiture, on disc only, those on odd interstriae slender, equal in length to width of an interstriae on both disc and declivity.
Biological Series, Vol. 15, No. 3 New Neothoi'ical Scolytidae
15
Female.— Similar to male except frons weak- ly convex, callu.s present; .short interstrial bristles on even-numbered interstriae slightly longer, ex- tending to declivity.
Type Loc.\lity.— Ten km SE Teziutlan, Puebla, Mexico.
Type MATEni.\L.— The male holotvpe, female allotype, and 24 parat>'pes were taken at the type' locality, from AInus, on 2 July 1967, 1600 m elevation. No. 141, by S. L. Wood.
The holotvpe, allotv'pe, and paratypes are in my collection.
Scohjtodes ominateus, u. sp.
This unique species is distinguished from all other species in the genus by the large, very narrowly separated eyes on the vertex, by the strongly impressed lower half of the female frons, by the fine, greatly reduced vestiture, and by the sulcate elytral declivity. It evidently rep- resents a new species group in the genus. It would be placed in Ptjcnartlinim except for the antennal club.
Female.— Length 2.0 mm (paratypes 1.7-2.0 mm), 2.6 times as long as wide; color yellowish to reddish brown.
Frons very strongly concave on upper half, abruptly, rather strongly impressed on lower half; entire surface reticulate, with rather ob- scure, shallow, rather fine, moderately sparse punctures; vestiture fine, sparse, inconspicuous. Eyes greatly enlarged, approximate above, sep- arated by a distance equal to less than diameter of two facets. Antennal club with suture 1 partly septate, others obsolete.
Pronotum 1.2 times as long as wide; widest on basal fourth, sides almost straight and paral- lel on basal two-thirds, rather broadly rounded in front; moderately declivous on anterior fourth; surface reticulate, rather fine, shallow, moderately abundant. Vestiture minute, incon- spicuous, a few longer hairs on anterior margins.
Elytra 1.5 times as long as wide, 1.5 times as long as pronottim; sides almost straight and parallel on basal two-thirds, rather broadly rounded behind; striae not impressed, punctures shallow, moderately large, in somewhat indefi- nite rows; interstrial punctures slightly smaller than those of striae, not easily disHnguLshed. Declivity steep, convex and rather narrowly bisulcate; declivital punctures fine, confused, indistinct; sutural interspace moderately elevat- ed, a moderately deep, narrow sulcus on inter- striae 2; lateral areas convex. Vestiture consist-
ing of minute, fine, strial and interstrial hair, and a few long, erect hairlike setae on and near declivity.
Male,— Similar to female except frons not impressed, convex.
Type Locality.— Eight km south of Colonia (near Buenaventura), Valle de Cauca, Colombia.
Type Material.— The female holotype, male allotype, and 97 paratypes were taken at the type locality on 9 July 1970, 70 m elevation. No. 624, from CJu.sia twigs, by S. L. Wood.
Tlie holotype, allotype, and paratypes are in my collection.
Scolytodes punctifer, n. sp.
This pubescent species is allied to punctatus Eggers, but it may be distinguished by the much smaller strial punctures with the inter- strial punctures much smaller than those of the striae, by the smooth pronotum surface with smaller punctures, and by the more abundant elytral hair.
Male.— Length 1.5 mm, 2.0 times as long as wide; color very dark brown, elytra slightly lighter, abundant vestiture pale.
Frons convex, a slight transverse impression just above epistoma; surface obscurely reticulate, punctures rather small, deep, moderately close; vestiture fine, sparse, short. Sutures of antennal club almost obsolete.
Pronotum 0.95 times as long as wide; sides parallel, feebly arcuate on basal half, broadly rounded in front; surface obscurely reticulate behind, distinctly reticulate toward anterior mar- gin, punctures coarse, deep, close. Vestiture fine, abundant, moderately long.
Elytra 1.3 times as long as wide; sides almost straight and parallel on basal two-thirds; striae 1 feebly, others not impressed, punctures rather coarse, deep; interstriae smooth about one and one-half times as wide as striae, punctures rather large, uniscriate except slightly confused on 2. Declivity convex, rather steep; all punc- tures distinctly smaller, otherwise similar to disc. Vestiture of rather abundant, fine, erect, strial and interstrial hair, and slightly longer, uniseriate rows of slightly coarser, interstrial hair; each long hair about one and one-half times as long as distance between rows.
Type Locality. —Volcan Irazu, Cartago, Costa Rica.
Type Material.— The imique male holotype was taken at the type locality on 26 September
16
Brigham Young University Sc:ienc:e Bulletin
1963, 2300 m elevation, No. 207, from a twig of Oreopanax nubigenus, by S. L. Wood. The holotypo i.s in my collection.
Scolytodes Idrsuttis, n. sp.
This .species is allied to punctifer Wood, but it may be distinguished by the fine, confused elytral punctures, by the strongly reticulate, dull pronotal surface, and by the much finer pronotal punctures.
Malk.— Length 1.7 mm, 2.1 times as long as wide; color black.
Frons moderately convex, surface strongly reticulate, punctures fine, deep, moderately abundant; vestiture inconspicuous.
Fronotuni ().9.5 times as long as wide; outline as in punctifer; surface strongly reticulate, dull, punctures fine, rather shallow, moderately abun- dant. Vestiture fine, rather abundant, moderate- Iv long.
Elytra 1.2 times as long as wide; outline as in punctifer: strial and interstrial punctures equal in size, small, rather shallow, interstrial punc- tures confused, those of striae distinguished with difficultv. Declivitv convex, rather steep; punc- tures minute, contused. Vestiture of fine, long strial ;ind interstrial hair, and slightly longer uniseriate rows of interstrial hair; each longer hair slightly longer than distance between rows.
Type Locality.— Tapanti, Cartago, Costa Rica.
Typk Matkhial.— The unique male holotype was taken at the type locality on 2 July 1963, 1300 m elevation. No. 8, from ;m unidentified vine, by S. L. Wood.
The holotype is in mv collection.
Scohjtodes crasstis, n. sp.
The position of this small, stout species is probleiiKitical, but the rows of very fine, strial and interstrial hair suggest a possible relation- ship to liirsutus Wood.
Male.— Length 1.0 nun, 1.9 times as long as wide; color vellowish brown, apparently not fully colored.
Frons convex, surhice obscuielv reticulate, shining, punctures rather large, deep, not close; fine, moderately abundant toward epistoma.
Pronotum 0.94 times as long as wide; widest at base, sides indistinctlv arcuate, converging very slightly on basal two-thirds, rather narrow- ly rounded in front; surface smooth and shining, punctures small, rather shallow, moderately close, irregularlv spaced. Vestiture of short, very fine, moderately abundant recumbent hair.
Elytra 1.2 times as long as wide, 1.5 times as long as pronotum; sides almost straight and parallel on basal two-thirds, rather broadly rounded behind; striae feebly if at ail impressed, punctures large, moderately deep, close; inter- striae almost as wide as striae, smooth, shining, punctures very fine, uniseriate, close. Declivity convex, rather steep; strial punctures gradually reduced to about t^vo-thirds size on disc; inter- striae slightly wider than striae, very feebly con- vex. Vestiture consisting of unLseriate rows of very fine, recumbent, moderately long strial and interstrial hair; in addition, odd-numbered inter- striae on posterior half of elytra bear widely spaced, erect, spatulate bristles of moderate length, about six such bristles on each inter- striae.
Type LocALiTi.— Barro Colorado Island, Canal Zone, Panama.
Type Material.— The male holotype was taken at the type locality between November 1952 and March 1953, presumably at light, by J. Zetek.
The holotype is in my collection.
Scoh/todes pannticetis, n. sp.
This species and plicatus Wood are unique in having unusually broad, eostiform pronotal asperities. Tliis species is distinguished from ])Iicatm- by the smaller, shallow strial and inter- strial punctures, and by the very different sculp- ture of the female frons.
Female.— Length 1.7 mm (male 1.5 mm), 2.4 times as long as wide; color vellowish brown.
Frons flattened from epistoma to upper level of eyes, surface minutely, denselv pilose over almost entire flattened area; epistomal margin very slightly elevated, smooth, shining, a narrow median extension reaching level of antennal in- sertion; lateral and upper extreme margins bear- ing a row of long, yellow hair.
Pronotimi and elytra as in plicatus except pronotal punctures slightly larger, strial and interstrial punctures much larger and deeper.
Male.— Similar to female except frons eon- vex, reticulate, punctures moderately large, shal- low, rather sparse, subglabrous; pronotal asperi- ties slightlv larger; pronotal and elvtnd punc- tures distinctly finer.
Type Locality. —La Ceiba, Atlantida, Hon- duras.
Type Matekial.- The female holotype was taken at the type locality on 7 June 1949, at
Biological Series, Vol. 15, No. 3 New Neotropical Scolytidae
17
light, by E. C. Becker. The male allotype bears identical data except that it was taken on 17 June 1949.
The holotype and allotype are in my collec- tion.
ScoJijtodes micidti.s, n. sp.
This species is very closely related to tenuis Wood, but it may be distinguished by the closely set bristles on all interstriae, by the finer interstrial punctures, and by the glabrous cen- tral area on the female frons.
Female.— Length 1.4 mm (paratypes 1.2-1.4 mm), 2.9 times as long as wide; color almost black.
Frons flattened from epistoma to upper level of eves, median third on lower half smooth, shining, slightlv elevated, remaining area coarse- Iv, closely, deeply punctured; punctured area liearing a tuft of long, vellow hair.
Pronotum 1.2 times as long as wide; sides rather strongly constricted on basal half, other- wise as in tenuis except posterior area smooth, shining (feebly reticulate in a few specimens).
Elytra 1.8 times as long as wide; outline and sculpture as in tenuis. Vestiture consisting of fine, short strial hair and long, erect interstrial bristles; each interstrial bristle slightly longer than distance between rows and betsveen l)ristles within a row.
Male.— Similar to female except frons con- vex, deeplv, narrowlv, transverselv impressed above epistoma, surface coarsely, deeply punc- tured.
Type Locality.— Four miles north of Totola- pan, Oaxaca, Mexico.
Type Material.- The female holotype, male allotype, and 28 paratypes were collected at the t\^pe localitv on 20 June 1967, 1 100 m elevation. No. 68, in Ficus. bv S. L. Wood. One paratype was taken at Palin, Es(|uintla, Guatemala, 19 Mav 1964, 300 m elevation. No. 683, S. L. Wood.
The holotvpe. allotvpe, and paratypes are in my collection.
Microborus lectus, n. sp.
This species is allied to hoops Blandford, but it is distinguished by the more widely sepa- rated eyes and elongate frons, by the more coarsely punctured pronotum, by the more strongly reduced punctures on striae 1 on the lower half of the declivity particularly in the male, and bv the much more strongly elevated
male declivital interstriae 7 which continues to the apex.
Male.— Length 1.5 mm (paratypes 1.4-1.5 mm), 2.8 times as long as wide; rather dark reddish brown.
Frons moderatelv convex, a slight summit just above level of antennal insertion, a weak, transverse impression just above epistoma; sur- face reticulate at sides and above, shining, with sparse, moderately coarse punctures; eyes sepa- rated by twice width of an eye.
Pronotmn 1.4 times as long as wide; widest at base, sides feebly constricted, almost straight on basal three-fourths, rather broadly rounded in front; surface shallowly reticulate, punctures rather coarse, close, deep, oval. Glabrous.
Elytra 1.8 times as long as wide, 1.5 times as long as pronotum; outline about as in hoops except apex narrower; striae strongly impressed, punctmes large, deep, close; interstriae smooth, shining, strongly convex, as wide as striae, punc- tures fine, uniseriate, rather widely spaced. De- clivity steep, convex; striae strongly impressed except lower half of 1, punctures strongly re- duced, minute on 1; interstriae strongly nar- rowed on left side, moderately so on right, 2 feebly elevated and armed by about three to six small teeth (usuallv different numbers on right and left sides), 3 and 4 convex but unanned and continuing to 7, 7 strongly, acutely elevated and continuing to apex. Vestiture confined to declivity, of coarse, rather short hair; a few scales on interstriae 1 and 2.
Female.— Similar to male except pronotum more finely sculptured; elytral declivity sculp- tured about as on disc except interstrial punc- tures very finely granulate and punctures on striae 1 much smaller; declivital interstriae 7 only moderately elevated as in female hoops; scalclike setae on declivity absent.
Type Locality.— Carbonera Experimental Forest, about 50 km (airline) northwest of Merida, Venezuela.
Type Material.— The male holotype, female allotype, and 19 paratypes were collected at the type locality on 14 October 1969, 2500 m ele- vation. No. 57, from Chisici bark, by S. L. Wood. Most of the galleries were at the fracture point between the outer and inner bark of Chisia logs from which other bark beetles had pre- viously emerged. Evidently several successive generations were produced without emerging from the host between generations.
The holotvpe, allotype, and paratypes are in my collection.
18
)5ni(;iiAM Young Univi-hisity Sf:iiiN(;E Bulletin
Scolijlii.s hdiinensis, n. sp.
The frons resembles proximus Chapuis, the elytra are as in cristatus Wood. Stemum 2 re- sembles cristatus, but it is much more steeply elevated and more finely sculj^tured, and the median elevated process arises at the anterior margin of sternum 2, it is slightly thicker, of rather high unifonn height and descends abrupt- ly behind.
Male.— Length 2.6 mm (paratypes 2.0-2.9 mm), 1.7 times as long as wide; color almost black.
Frons broadly, weakly convex, a slight trans- verse impression on lower third, a moderately large, rounded, shining tubercle just above mid- dle, tubercle convex and ecjually precipitous on all sides; surface obscurely reticulate on upper half, fine, moderately deep punctures ar- ranged in obscure, shallow aciculate grooves, punctures on lower half replaced by close, rounded granules of ecjual size; epistomal mar- gin smooth; vestiture of moderately abundant, very fine, long hair, slightly longer laterally. Antennal club with suture 1 indicated.
Pronotum 0.97 times as long as wide; widest one-third length from base, sides weakly ar- cuate, converging slightly on basal two-thirds then moderatelv constricted laterally just before broadly rounded anterior margin; surface smooth shining, punctures on disc slightly oval in shape, rather close, moderately large for this genus, about two to three times larger in lateral areas. Glabrous.
Elytra outline as in cii.itdtus; striae and inter- striae equally, narrowly, stronglv impressed, punctmes moderatelv large, rather deep; inter- spaces between rows smooth and shining, about as wide as punctures. Glabrous except for a few scattered bristles on or near declivity.
Stemum 2 vertical, rather finely, deeply punctured; anned on slightlv more than anterior h;ilf by a median process that begins in cleft of anterior m;irgin of segment 2, process rising abruptly to m;ixiinum height and continuing at this level to its posterior extremity, rather ;ibruptly terminated behind; abdominal sterna 2-.5 with moderatelv ;ibuiulant erect, nither long bristles.
Female.— Similar to male except frontal tu- bercle not as high, granules on lower half of frons more poorlv developed to ol)solete.
Type L()c:ality.— Campamento Gachicamo. 40 km east of Canton, Barinas, Venezuela.
Type Matehial.— The male holotype, female allot)'pc, and 97 paratypes were taken at the
type locality on 8 March 1970, 70 m elevation, No. 356, from a large vinelike tree known local- ly as Hevecito, by S. L. Wood. The parental tunnels were biramose and transverse; two fe- males were associated with each male.
The holotype, allotype, and paratypes are in my collection.
Scohjtopsis orinocanus, n. sp.
This species apparently is allied to peruanus Eggers. From the key and description of the female (Eggers, 19.37, Rev. de Ent. 7:83) it is distinguished from Eggers' species by the broad- ly oval or round punctures on the female frons, and the broadlv raised median area of the epistoma (not carinate), by the unifonnly short, frontal vestiture and, evidently, by the coarser pronotal punctures and shorter elytral setae.
Female.— Length 3.0 mm (paratypes 2.9-3.3 mm), 2.0 times as long as wide; very dark brown to black, pronotum usually darker, scales rather dark.
Frons broadly convex, a distinct, transverse impression alcove epistoma, median area of im- pression broadly, distinctly elevated (not cari- nate); epistoma rather well developed, slightly elevated; surface smooth, shining, with dense, deep, moderately coarse, round or broadly oval punctures; vestiture of short, moderately abun- dant, coarse setae of almost unifonn length.
Pronotum 0.95 times as long as wide; widest one-third length from base, sides rather strongly arcuate, converging only slightly before an- terior third, rather broadly rounded in front; surface smooth, shining, with a few \'erv minute, impressed points, punctures rather fine (almost as large as in pimcticoUls Blandford), oval on disc, up to twice as large, subcircular and closer in latend areas, a row of larger punctmes at lateral and basal margins. Glabrous.
Elvtra 1.3 times as long as wide, 1.3 times as long as pronotum; slightlv naiTOwer than pronotum; sides shallowlv eniarginate on basal half, rather broadlv rounded Ix-hind; striae and interstri;ie on basal half equallv, rather deeply impressed, punctures moderatelv coarse, deep, mostiv of about equal size; on posterior half interstriae gradually, less strongly impressed until almost flat toward apex, interstrial punc- tmes also decr<>asing in depth and clarity pos- teriorlv, centnil third of each interstriae on pos- terior h;df etched, dull, spreading to entire inter- striiie, then to entire declivital surface near apex; eost;il margin near apex almost smooth. Vesti- ture of slender, suberect scales; each scale slight- ly shorter than distance between rows; very
Biological Series, Vol. 15, No. 3 New Neotropical Scolytid.\e
19
slightly more slender and e(jual in length to puncticoUis.
Male.— Similar to female except flat from vertex to epistoma ( no arch ) , without elevations except a feeble one at epistoma, central area smooth and shining, lateral areas with dense, long pubescence slightly shorter than in other species.
Type Loc.'Vlitv.— Campamento Rio Grande, 30 km east of Palmar, Bolivar, Venezuela.
Type Material.— The female holotype, male allotype, and 4S paratypes were taken at the type locality on 12 June 1970, 70 m elevation. No. 573, in Tenninaliu guianensis, by S. L. Wood. These monogamous beedes made bira- mous, longitudinal tunnels similar to those of Scohjtus.
The holotype, allotype, and paratypes are in my collection.
Carphodicticus, n. gen.
Blandford ( 1895, Ann. Mag. Nat. Hist., Sen 6, 15:317) described from Ceylon a peculiar genus, Craniodicticus, "of doubtful relationship," which he placed in the Scolytidae but suggested a possible relationship to ScJwdlarius Wood {=Chapui-si<i Duges). Except for the addition of one other species, minor Eggers ( 1936, Ann. Mag. Nat. Hist,, Ser. 17, 10:635), from India, this genus has scarcely been mentioned in the literature. It is very primitive and has several characters suggesting a relationship to the Platy- podidae, although it clearly belongs in Scoly- tidae. Except for the fact that it was taken from a liana ("jungle rope") nothing of its habits is known.
An equally peculiar insect belonging to the same generic group with Craniodicticus recently was collected in Venezuela. Both genera have the body elongate, the antennal funicle 5-seg- mented, the scape simple, the club with two, straight, transverse sutures, the pronotum lateral- ly excised as in primitive Platypodidae, the an- terior coxae narrowly separated, the prostemum extending well behind the anterior coxae, the mesostemum feebly inflated, the scutellum small, the tibiae short, rather broad, with four socketed teeth, and the basal margias of the elytra unarmed. These genera do not belong to any previously recognized tribe of Scolytidae; therefore, the new tribal name Cai-phodicticini is proposed to include them.
Diagnosis.- This genus Ls rather closely allied to Craniodicticus, but it is distinguished by the larger, somewhat asymmetrical antennal club,
by the very large, elongate, shallowly emargin- ate, coarsely faceted eyes, by the very broad oral region, ;uid by entirely different types of characters on the frons and elytral declivity.
Description.— Eyes large, broadly emargin- ate; antennal scape simple, funicle 5-segment- ed, club moderately flat, slightly asymmetrical, with two straight, transverse sutures clearly in- dicated and at least partly septate, a third su- ture obscurely indicated. Pronotum elongate, laterally excised as in primitive Platypodidae, anterior coxae moderately separated, prostemum extending well behind coxae. Scutellum small, not depressed. Elytral bases unanned; inter- striae 10 obsolete on posterior half. Anterior coxae with raised cusp on anterodistal margin; tibiae short, broad, anned by several teeth on outer margin; tarsi longer than tibiae, segments cylindrical, 1, 2, and 3 about equal in length.
Type Species.— Carphodicticus cristatus Wood, described below.
Carphodicticus cristatus, n. sp.
The only known member of the family that could possibly be confused with this species is Craniodicticus mucronatus Blandford, but the generic characters cited above distinguish it. The very elongate platypodid body form and armature of the elytral declivity are unique.
Male.— Length 2.2 mm (paratypes 1.9-2.4 mm), 3.4 times as long as wide; color reddish brown.
Frons broad, very strongly convex, protrud- ing abruptly above elevated, shining epistoma, very closely, deeply, rather finely punctured; glabrous except a few short bristles just above epistoma and a broad, dense epistomal brush of long coarse setae; oral area very broad. Eye elongate, broadly, shallowly emarginate, coarse- ly faceted; about four times as long as wide; more than half of eye below antennal socket. Antennal scape moderately short; funicle 5- segmented; club as described above.
Pronotum 1.4 times as long as wide; widest at basal and anterior angles, basal and anterior margins very broadly rounded, lateral margins on basal half broadly, deeply excised as in Platy- podidae; surface smooth, shining, a few minute points, punctures close, deep, moderately coarse. Glabrous, except a few minute, bifid hairs on basal margin.
Elytra 2.0 times as long as wide, 1.5 times as long as pronotum; sides almost straight and parallel on basal three-fourths, then abruptly rounded slightly to projecting scoop formed by
20
Brigham Young University Science Bulletin
declivital costae, broadly, shallowly emarginate behind; bases feebly emarginate at scutelluin; scutellum small, oval, attaining elytra! surface; striae 1 very feebly impressed, others not im- pressed, punctures coarse, deep, close; inter- striae half as wide as striae, smooth, shining, punctures minute, shallow, irregularly uniseriate. Declivity commencing three-fourths of elytral length from base, steep; striae 1 and 2 rather coarsely punctured; interstriae 1 distinctly ele- vated, convex, summit smooth, 2 impressed, nar- row, obsolete before apex, 3 very strongly, nar- rowly elevated from just below declivital base to just before apex, crest smooth; punctures on lateral interstriae near declivity rather coarse, not granulate.
Protibiae two-thirds as long as femur, broad, terminal mucro curved toward tarsal insertion, four socketed teeth on lateral margin; tarsi about as long as tibia, segments 1, 2, and 3 cylindrical, about equal in length.
Vestiture confined to declivity outside of concave area, consisting of coarse bristles in interstrial rows.
Female.— Similar to male except frons less strongly protuberant immediately above epi- stoma.
Type Locality. —Eight km west of Bumbum, Barinas, Venezuela.
Type Material.- The male holotype, female allotype, and 110 para types were collected at the type locality on 11 February 1970, 150 m elevation. No. 326, by S. L. Wood.
The large, cut host tree was not recognized by local professional foresters; it may have been a rare Lauraccae. The tree had been cut several months and had been largely abandoned by a species of Pliloeotrihus. This species entered through Phloeotribus entrance tunnels, followed along an old egg gallery to a convenient point where their tunnels commenced along the cam- bium, engraving both phloem and xylem tissues. The beetles were monogamous and worked in pairs. The egg tunnels branched occasionally, with a different pair of beetles working on each branch, all using the same original entrance hole. Egg niches were fonned along the cam- bium on each side of the gallery where eggs were deposited individually. The larvae con- structed short, irregular mines in the phloem next to the cambium. The wood was exceed- ingly dry and hot, but the infestation was thriving.
The holotype, allotype, and paratypes are in mv collection.
Pseudothtjsanoes quemeus. n. sp.
This species i.s rather closely related to quer- cintK Wood, but it is distinguished by the more coarsely punctured, rugulose pronotal disc, by the coarser strial punctures, and by the much wider elytral scales.
Male.— Length 1.3 mm (paratypes 1.2-1.5 mm), 2.5 times as long as wide; color black.
Frons convex, a slight transverse impression just above epistoma; surface obscurely, rather coarsely punctured, rugulose; vestiture of mod- erately abundant, rather short, coarse setae. Antennal scape short, broad, slightly longer than pedicel; club rather small, widest through seg- ment 2, sutures 1 and 2 slightly procurved, 1 marked by setae only at sides.
Pronotum 0.87 times as long as wide; widest on basal third, sides arcuately converging to- ward broadly rounded anterior margin; anterior margin armed by six low teeth; summit at mid- dle, moderately high; anterior slope asperate; posterior area coarsely, indistinctly punctured, rugulose. Vestiture of moderately abundant, stout setae.
Elytra 1.6 times as long as wide; sides straight and parallel on basal two-thirds, rather narrowlv rounded behind; striae not impressed, punctures moderately coarse, rather deep; inter- striae as wide as striae, somewhat irregular, punctures rather fine, uniseriate. Declivity steep, convex; striae irregularly, weakly im- pressed, interstrial punctures with coarse, low granules; most of surface obscurely subgranu- lose. Vestiture of semirecumbent strial hair, and rows of equally long, erect interstrial scales; each scale about twice as long as wide.
Fenl\le.— Similar to male except antennal scape as wide as long, bearing a tuft of very long white hair; teeth on anterior margin of pronotum smaller, some obsolete; interstrial scales slightly longer and more slender, each about three to four times as long as wide.
Type Localit\.— Seventeen km ( 10 miles ) east of Pachuca, Hidalgo, Mexico.
Type Material.— The male holotype, female allotype, and 15 paratypes were taken at the type locality on 10 June 1967, at an elevation near 2500 m. No. 5, in Qucrcus. by S. L. Wood. Ten paratypes were taken S km (5 miles) west of Tulancingo, Hidalgo, Mexico, 11 June 1967, 2400 m elevation. No. 10, in Qucrcus, by S. L. Wood. Nine paratypes were collected 15 km (9 miles) east of Huatasco, Veracruz, 7-VII-67, .300 m elevation. No. 173, in Qucrcus, by S. L. Wood.
Biological Series, Vol. 15, No. 3 New NEOTiioriCAL Scoi.ytidae
21
The holotype, allotype, and paratypes are in mv collection.
Pseudothijsanoes furvesren.s, n. sp.
This species is in the phoradendri group very closely related to furvtts Wood. From furvus it may be distinguished by the absence of gran- ules on the male discal interstriae and by the shorter elytral scales.
M.\LE.— Length 1.3 mm (paratypes 1.1-1.4 mm), 2.3 (female 2.8) times as long as wide; color black.
Frons con\ex, transversely impressed above epistoma; surface finely rugulose, punctures not evident; vestiture of sparse, coarse, short setae. Antennal scape elongate; moderately large, oval, widest at middle, suture 1 straight, 2 obscure.
Pronotum as in furvus.
Elytral proportions and outline as in furvus; striae not impressed, punctures small, shallow; interstriae slightlv wider than striae, smooth, punctures very fine, not at all granulate or ele- vated. Declivitv' convex, steep; striae as on disc; interstriae each with a row of fine granules. Vestiture consisting only of rows of interstrial scales, each scale about three times as long as wide, slightly shorter than distance between rows, spaced within a row by about one and one-half times length of a scale.
Female.— Similar to male except more slen- der, 2.8 times as long as wide; frons less strongly convex; anterior margin of pronotum unanned; strial punctures much smaller, obscure; occa- sional strial hair present.
Type Locality.— Eighteen km (11 miles) north of Huajuapan, Oaxaca, Mexico.
Type Material.— The male holotype, female allot\'pe, and 14 paratypes were taken at the type localitv on 15 June 1967, No. 42, from Phoradendron branches, by S. L. Wood.
The holotvpe, allotype, and paratypes are in my collection.
Pseudothijsanoes funereus, n. sp.
This species Ls very closely related to phora- dendri Blackman, but it is distinguished by the more slender body form, by the more rugulose pronotal disc, by the much smaller strial punc- tures on both disc and declivity, and by the near absence of interstrial granules.
Male.— Length 1.4 mm (parat}'pes 1.3-1.5 mm), 2.4 times as long as wide; body color black.
Head and prothorax as in phorandendri ex- cept pronotal disc slightly more rugulose.
Elytra 1.5 times as long as wide; sides straight and parallel on basal two-thirds, nar- rowly rounded behind; striae not impressed, punctures small, shallow, separated by two or more diameters of a puncture; interstriae smooth, at least twice as wide as striae, punc- tures uniseriate, very small, feebly if at all granu- late. Declivity beginning well behind middle, convex; interstrial granules small, regular. Ves- titure as in phoradendri.
Type Locality.— Volcan Colima, Jalisco, Mexico.
Type Material.— The male holotype and six male paratypes were taken at the type locality on 23 June'l965, 2.500 m elevation. No. 108, from Phoradendron longifolium, by S. L. Wood.
The holotype and paratypes are in my col- lection.
Pseudothijsanoes graniticus, n. sp.
This species is allied to verdicus Wood and viscicolens Wood, but it is distinguished by the shallowly impressed female frons with setae on the vertex short, and by the more coarsely granu- late male declivital interstriae with scales on lower half of interstriae 1 smaller.
Female.— Length 1.9 mm (male 1.8 mm), about 2.7 times as long as wide, color very dark brown.
Frons shallowly concave from epistoma to upper level of eyes, flattened to vertex; surface of concave area smooth, shining, impunctate, finely punctured above, somewhat strigose at sides; epistomal brush conspicuous at sides; flat- tened area above eyes bearing rather abundant, moderately short, coarse hair. Antennal scape elongate, ornamented by abundant, long hair; club about 2.5 times as long as wide; antenna as long as pronotum.
Pronotum 0.96 times as long as wide; widest at base, sides almost straight, converging feebly on basal half, broadly rounded in front; anterior margin unarmed; summit rather high, at middle; anterior area asperate; posterior area rugose- reticulate, with fine, obscure punctures, fine granules behind summit. Vestiture sparse, of fine and coarse, short hair.
Elytra about 1.6 (slightly spread) times as long as wide, 1.8 times a,s long as pronotum; striae not impressed except 1 weakly, punctures rather small, deep; interstriae slightly less than twice as wide as striae, almost smooth, uni- seriately, finely granulate. Declivity convex,
22
Bricham Young University Science Bulletin
steep; interstrial granules slighdy larger, con- fused on 2 and 3. Vestiture of very fine strial hair and rows of erect interstrial scales; each scale on disc three to four times as long as wide, four to five times as long as wide on declivity, each scale slightly longer than distance between scales within a row or between rows.
Male.— Similar to female except stouter, about 2.5 times as long as wide (estimated); frons not visible; pronotum distinctly constrict- ed on anterior half, anterior margin anned by ten teeth; strial punctures coarser, interstriae one and one-half times as wide as striae, much more coarsely granulate; elytra! scales slightly longer, proportions as on female; scales on low- er half of interstriae 1 conspicuously smaller; granules on declivital interstriae 2 and 3 uni- seriate.
Type Locality.— Forty-three km (26 miles) southeast of Nochixtlan, Oaxaca, Mexico.
Type Matfrial.— The female holotype and male allot\'pe were collected at the type locality on 17 June 1967, No. 55, from Phoradendwn, by S. L. Wood.
The holotj'pe and allotype are in my collec- tion.
Pseiidotlnjsanoes verticillus, n. sp.
This species is very closely allied to viscico- lens Wood, but it may be distinguished by the more shallowly concave female frons, with a more dense tuft of hair on the vertex, and by the much shorter elytral scales.
Female.— Length L6 mm, 2.8 times as long wide; color almost black, pronotal summit red- dish brown.
Frons as in viscicolens except less deeply con- cave, tuft of hair on vertex slightly shorter, more dense. Antenna and pronotum as in viscicolens.
Elytra L7 times as long as wide, L8 times as long as pronotiim; striae not impressed, punc- tures rather small, moderately deep; interstriae about twice as wide as striae, punctures uniseri- ate, finely granulate. Declivity as in groniticu.s Wood, with indistinct granules and punctures on interstriae 2 and 3 confused. Vestiture of fine, short strial hair, and rows of erect, short, inter- strial scales; each scale one to one and one- half times as long as wide, half as long as dis- tance between rows, equal in length to distance between scales in a row.
Type Locality.— Twenty-seven km ( 16 miles ) north of Ixmiquilpan, Hidalgo, Mexico.
Type Material.- The unique female holo- type was taken at the type locality on 10 July 1967, 1900 m elevation, probably from Phomden- dron, by S. L. Wood.
The holotype is in my collection.
Pseudotlujsanoes cuspidi.s, n. sp.
This species is allied to suhulahis Wood and spicatus Wood, but it may be distinguished by the smaller size, by the flattened and grooved female frons, by the coarser punctures on the elytral declivity, and by the slender, rather long elytral setae.
Female.— Length 1.3 mm (paratypes 1.2-1.4 mm), 2.7 times as long as wide; color very dark brown.
Frons transversely flattened, longitudinally concave from epistoma to upper level of eyes, median line narrowly sulcate from level of an- tennal insertion to upper level of eyes; surface smooth and shining on epistomal area, then finelv substrigose becoming more coarsely re- ticulate-granulate above eyes; vestiture sparse, fine, inconspicuous. Antennal scape elongate; club moderately large, sutures very obscure.
Pronotum 1.0 times as long as wide; sides feebly arcuate, subparallel on basal half, dis- tinctly constricted on anterior half, broadly rounded in front; summit at middle, moderately high; sparsely asperate on anterior slope; finely rugose-reticulate behind, a few minute granules. Vestiture of sparse hair.
Elytra 1.8 times as long as wide, 1.8 times as long as pronotum; sides almost straight and parallel on more than basal two-thirds, rather narrowly rounded behind; striae not impressed, punctures small, shallow; interstriae not smooth, less than twice as wide as striae, punctures fine, obscured by surface sculpture. Declivity convex, steep; interstriae uniseriately, finely granulate. Vestiture of rows of very fine strial hair and rows of slightly longer, coarser, interstrial setae, not at all scalelike; interstrial setae about as long as distance between rows.
Last visible sternum narrowly produced to fonn a mucronate process.
Type Locality.- Ten km (6 miles) north- east of Teziutlan, Puebla, Mexico.
Type Material.— The female holotv'pe and 13 female paratypes were taken at the type lo- cality on 2 July 1967, 1600 m elevation. No. 144, by S'. L. Wood.
The holotv'pe and paratypes are in my col- lection.
Biological Series, Vol. 15, No. 3 Nevv Neotbopical Scolytidae
23
Pseudothysanoes tenellus, n. sp.
Presumably this species is allied to Jwi)kin.si Blackmail, but it is distinguished by the very elongate interstrial scales on the male declivity, by the flattened frons in both sexes with a feeble central impression.
Male.— Length 1.6 mm (paratypes 1.3-1.6 mm), 2.4 times as long as wide; color almost black.
Frons almost flat from transverse impression just above epistoma to upper level of eyes, con- vex above, a small median impression at upper level of eyes; smooth and shining near median line, becoming rugose-reticulate in lateral areas, rugose-punctate above eyes; vestiture of sparse, coarse setae. Scape elongate, slender; club long- er than scape, broadlv oval, two sutures clearly marked, raflier strongly procurved.
Pronotum 1.0 times as long as wide; widest at base, sides strongly arcuate, very strongly con- stricted on anterior half, rather narrowly round- ed in front; anterior margin armed by six coarse teeth; summit behind middle; asperate on an- terior slope; reticulate-granulate in posterior area, with sparse, fine, obscure punctures, a few granules behind summit. Vestiture of sparse, short, stout setae.
Elytra 1.4 times as long as wide, 1.5 times iis long as pronotum; sides almost straight and parallel on basal two-thirds, obtusely angulate behind; striae not impressed, punctures mod- erately large, deep; interstriae slightly wider than striae, almost smoodi, uniseriately, coarsely granulate on posterior third, punctures feebly to not at all granulate to base. Declivity broadly convex, rather steep; striae weakly impressed on upper two-thirds, punctures smaller; uniseri- ate interstrial granules rather coarse at base, de- creasing in size on upper two-thirds, almost ob- solete below. Vestiture consisting of rows of interstrial scales, short on disc, each about three times as long as wide, increasing in length to- ward declivity, very long and delicate on de- clivity, some about four times as long as distance between rows.
Female.— Similar to male except frons more extensively flattened, more finely sculptured; pronotum similarly, almost equally armed; in- terstrial granules slightly smaller; declivital scales little longer than those on disc, each about three times as long as wide, those on interstriae 1 and 2 smaller.
Type Locality.— Thirty-three km (21 miles) west of Morelia, Michoacan, Mexico.
Type Materl\l.— The male holotype, female allotype, and 17 paratypes were taken at the type locality on 16 June 1965, at 2300 in eleva- tion. No. 64 from Phoradendron twigs, by S. L. Wood.
The holotype, allotype, and paratypes are in my collection.
Pseudothysanoes yuccavorus, n. sp.
This species is allied to hopkinsi Blackman, but it may be distinguished by the flattened frons with the absence of a central fovea, by the smaller, more slender antennal club, and by the shorter, stouter interstrial scales.
Male.— Length 1.2 mm (paratypes 1.1-1.3 mm), 2.2 (female 2.5) times as long as wide; color black.
Frons weakly convex, abnost flat to upper level of eyes, a weak transverse impression just above epistoma; epistoma shining, remaining area rather coarsely reticulate-granulate, punc- tures fine, obscure; vestiture of sparse, short, coarse setae. Antennal scape elongate, slender; club small, 1.6 (female 1.8) times as long as wide, sutures very obscure, weakly procurved.
Pronotum 0.92 times as long as wide; sub- triangular, widest at base, sides weakly arcuate, converging toward narrowly rounded anterior margin; anterior margin armed by four coarse teeth; summit high, narrow; anterior area coarse- ly asperate; posterior area rugulose, a few fine granules behind summit; vestiture of sparse, coarse setae of moderate length.
Elytra 1.3 times as long as wide, 1.6 times as long as pronotum; sides almost straight and parallel on slightly more than basal hvo-thirds, rather broadly rounded behind; striae not im- pressed, punctures small, moderately deep; in- terstriae twice as wide as striae, almost smooth, uniseriately, rather coarsely granulate. Declivity rather steep, convex; as on disc. Vestiture con- sisting of minute, fine strial hair, and erect, inter- strial scales; each scale three times as long as wide, separated betsveen rows and within a row by distances slightly greater than length of a scale.
Female.— Similar to male except body form more slender; antennal club longer; anterior margin of pronotum more broadly rounded, un- armed; interstrial granules almost obsolete on disc; interstrial scales slightly longer, very slen- der, each scale eight or more times as long as wide.
Type LocALirv.— Fifteen km (9 miles) west of Durango, Durango, Mexico.
24
Brigham Young University Science Bulletin
Type Material.— The male holotypc, female allotype, and 45 paratypcs were collected at the type locality on 4 June 1965, at 1300 m elevation, No. 1, from dying Yucca leaves on a large, healthy plant, by S. L. Wood.
The holotype, allotype, and paratypes are in my collection.
Thysanoes inornatiis, n. sp.
Thi.s species is distinguished from the closely allied mexicanus Wood by the greatly reduced size of strial punctures on the declivity, by the shorter, more widely spaced interstrial scales, by the finer interstrial granules on the declivity, and by the abnost glabrous female frons.
Female.— Length 1.6 mm (paratypes 1.4-1.7 mm), 2.7 times as long as wide; color very dark brown.
Frons moderately, rather broadly concave from epistoma to upper level of eyes; surface above concavity reticulate-granulate, becom- ing reticulate on upper part of concavity, smooth and shining over lower areas; a fovea at center; almost glabrous except near lateral mar- gins of epistoma. Antennal scape bearing nu- merous long setae.
Pronotum 1.04 times as long as wide; widest one-third length from base, sides arcuate, not converging on basal half, rather strongly con- stricted in front of middle, moderately rounded in front; anterior margin anned by six teeth; summit at middle; posterior area reticulate to obscurely reticulate-granulate, a few minute, ob- scure punctures. Vestiture sparse, consisting of a few short, stout setae.
Elytra 1.6 times as long as wide; sides almost straight and parallel on basal three-fourths, rather narrowly rounded behind; striae not im- pressed, punctures small, shallow; interstriae al- most smootli, with obscure lines, almost three times as wide as striae, puncture sparse, uniser- iate, very finely granulate. Declivity steep, con- vex; strial punctures reduced, minute to obsolete; interstrial granules regularlv spaced, fine. Ves- titure of rows of very minute, fine, strial hair, and rows of erect interstrial scales; each scale three to four times as long as wide, spaced with- in a row by one and one-half to three times length of a scale, between rows by about one and one-half times length of a scale.
Male.— Similar to female except stouter, 2..3 times as long as wide; frontal impression smaller, about two-thirds as large, reticulation extending almost to epistomal area; scape sparsely pubes- cent; teeth on anterior margin of pronotum larg-
er; interstrial scales wider, each two to three times as long as wide.
Type LocALiTi.— Volcan Colima, Jalisco, Mexico.
Type Material.— The female holotype, male allotype, and five paratypes were taken at the type locality on 23 June 1965, 2500 m elevation. No. 103, from an unidentified shrub, by S. L. Wood. Eight paratypes bear identical data ex- cept they were No. 106, taken from small branches of a Rhus near aromatlca. Ten para- types were collected at Aguascalientes, Aguas- calientes, Mexico, 12 June 1965, 2100 m eleva- tion. No. 43, from Mimosa branches, by S. L. Wood.
The holotype, allotype, and paratypes are in my collection.
MicraciseUa adnata, n. sp.
This species is distinguished from the allied knulli Blackman bv the presence of six teeth on the anterior margin of the pronotum, bv the finely tuberculate, almost unelevated declivital interstriae 3, bv the wider interstrial scales, and by the median impression on the female frons.
Female.— Length 1.9 mm (paratypes 1.9-2.0 mm), 2.8 times as long as wide; color very dark reddish brown.
Frons convex, with a narrow, subtriangular, median impression from near upper level of eyes to level of antennal insertion; surface re- ticulate-granulate, apparently with a few larger granules; vestiture of moderately abundant, coarse, moderately short setae. Eye slightly more than twice as long as wide; antenna as in knulli.
Pronotum 1.1 times as long as wide; as in knuUi except anterior margin armed by six teeth, and posterior area rugulose-reticulate, punctures fine, shallow, obscurely subtuberculate. Vesti- ture abraded.
Elytra 1.8 times as long as wide, 1.6 times as long as pronotum; outline as in knuUi: striae not impressed, punctures fine, shallow; inter- striae twice as wide as striae, subnigulosely marked bv irregular lines, punctures uniseriatc, very fine, shallow, obscure. Declivit)' steep, con- vex, apex protruding and minutely divaricate, mucro absent; surface on lower Uvo-thirds nigu- lose-reticulate; striae obscure but \isible; inter- striae 1 obscurely, very minutelv, uniseriately granulate, 2 with a few granules on upper fourth, 3 with larger granules to junction with 9, 7 and 9 with granules similar to 3. Vestiture abraded on disc; on declivitv consisting of rows of min- ute strial hair and rows of suberect, interstrial
Biological Series, Vol. 15, No. 3 New Neotropical ScoLYTn)AE
25
scales; each scale with a shght arcli or curl to- ward posterior direction, two to four times as long as wide, each almost as long as distance be- tween rows, slightly closer within a row, scales absent on lower half of interstriae 2.
Male.— Similar to female except (head con- cealed) strial punctures slightly deeper; declivi- tal interstriae 1 and 3 feebly elevated, 9 distinct- ly elevated; interstrial scales averaging slightly more slender.
Type Locality.— Fourteen km (9 miles) east of Huatusco, Veracruz, Mexico.
Type Material.— The female holotype, male allot)'pe, and one damaged female paratype were taken at the type locality on 7 July 1967, 270 m elevation. No. 173, from Quercus twigs, by S. L. Wood.
The holotype, allotype, and paratype are in my collection.
Micracisclla serjaniae, n. sp.
This species is allied to striata Wood, but it may be distinguished by the much smaller strial punctures, by the less nearly granulate discal interstriae, by the distinctly, uniseriately granu- late declivital interstriae, by the absence of a preapical emargination on the costal margin of the elytra and by a distinct sutural emargination at the elytral apex.
Female.— Length L3 mm (paratypes 1.2-L3 mm), 3.0 times as long as wide; color very dark reddish brown, pronotmn lighter.
Frons, antenna, and pronotum as in striata, except segment 1 of antennal club distinctly shorter.
Elytra 1.8 times as long as wide, 1.7 Hmes as long as pronotum; outline as in striata except apex wider and suturally emarginate; striae not impressed, punctures moderately small; inter- striae shining, somewhat irregular, one and one- half times as wide as striae, punctures fine, uni- seriate, not at all granulate. Declivity steep, con- vex, produced apically; strial punctures deeper than on disc; interstrial punctures granulate. Vestiture as in striata except interstrial scales very slightly larger; each scale about two to three times as long as wide.
Male.— Similar to female except tuft of hair on scape much smaller; discal interstriae evi- dently less irregular.
Type Locality.— Zamorano, Morazan, Hon- duras.
Type Material.— The female holotype, male allotype, and 83 paratypes were taken at the
type locality on 18 April 1964, at 700 m eleva- tion. No. 547, from Scrjania racemosa, by S. L. Wood.
The holotype, allotype, and paratypes are in my collection.
Micracis amplinis, n. sp.
This is the largest known species in the genus; it is also distinguished from all other representatives of the genus by the moderately impressed discal striae and by the very broad mterstrial scales on the declivity.
Male. -Length 3.2 mm (paratype 3.3 mm), 3.0 times as long as wide; color reddish brown.
Frons convex, with a moderate, transverse impression just above epistoma; surface finely reticulate-granulate, with rather numerous large, low, rounded granules widely distributed on convex area; epistomal area smooth, shining; glabrous except for a few setae lateral to epis- toma. Anteimal scape only slightly flattened; su- ture 1 reaching middle of club.
Pronotum L03 times as long as wide; widest on basal third, sides weakly arcuate, shghtly constricted on anterior third, rather broadly rounded in front; anterior margin armed by ten coarse serrations; summit in front of middle; anterior area coarsely serrate; posterior area reticulate-subgranulate, dull, a few, small, shin- ing granules to base. A few stout setae in lateral areas.
Elytra 1.8 times as long as wide, 1.8 times as long as pronotum; sides almost straight and parallel on basal three-fourths, strongly, acutely acuminate and mucronate behind; striae im- pressed except on basal fourth, punctures mod- erately small, deep; interstriae shining, almost smooth, three times as wide as striae, punctures very minute, almost obsolete, each with two or three large granules at base of declivity. De- clivity rather steep, convex, apex produced; entire surface minutely granulose; strial punc- tures small, deep; interstriae 1 and 3 weakly elevated, punctures on all interstriae as large as those of striae, confused, deep. Vestiture con- fined to declivity (possibly abraded on disc), of large interstrial scales; each scale about as wide as long, a few as wide as an interstriae.
Type Locality.— Five km (3 miles) west of El Salto, Durango, Mexico.
Type Material.— The male holotype and one damaged male paratype were taken at the type locality on 7 June 1965, 2500 m elevation. No. 41, from a Quercus branch, by S. L .Wood.
26
Bricham Young University Science Bulletin
The holotype and paratype are in my col- lection.
Micracis incertus, n. sp.
This species is allied to lepidus Wood, but it is distinguished by the presence of a few shallow punctures on the pronotal disc, by the larger interstrial tubercles on the declivity, and by the distribution.
Female.— Length 2.4 mm (paratypes 2.0-2.4 mm), 3.0 (male 2.8) times as long as wide; color vei-v dark brown.
Frons convex, a moderate, transverse im- pression just above epistoma; surface finely reticulate-granulate, dull, upper half with rather abundant, shallow punctures of moderate size; a few stout setae in lateral and epistomal areas. Antennal scape moderately triangidar, orna- mented by rather abundant, long hair; suture 1 not (juitc reaching middle of club.
Pronotuin 1.1 times as long as wide; as in amplinus except moderately large, shallow, rather abundant punctures intermixed with shining granules on disc. Vestiture of sparse scales and minute hair.
Elytra 1.8 times as long as wide, 1.7 times as long as pronotum; outline as in amplinis; striae not impressed, punctures moderately large; interstriae shining, subrugose, twice as wide as striae, j^^nctures moderately large, uniseriate, their anterior margins finely granu- late. Declivity steep, convex, apex produced; surface dull; strial punctures small, deep, reduced toward apex; interstrial punctures on 1 and 2 fine, irregular and reduced on lower half, on .3 and lateral areas moderately large, rounded. Vestiture of rows of interstrial scales, largely abraded on disc, each scale about five times as long as wide, as long as distance be- tween rows.
Male.— Similar to female except frons more strongly convex, punctures replaced by granules; discal interstrial granules smaller except larger near declivity, dechvital granules larger, ex- tending to apices of 1 and 2; interstrial scales on upper fourth of declivity longer, some six or eight times as long as wide.
Type LocALrrY.— Thirty-seven km ( 22 miles ) west of Durango, Durango, Mexico.
Type Matehial.— The female holotype was taken at the type locality on 4 June 1965, at 2000 m elevation. No. 4, from Qiierctis brandies, by S. L. Wood. The male allotype and 5 para- types (in poor condition) were taken 64 km
(10 miles) west of Durango on the same day, from the same host and collector. No. 18, at 2500 m elevation. One paratype was taken 5 km (3 miles) west of El Salto, Durango, 7 June 1965, 2500 in elevation. No. 41, from Qttercns by S. L. Wood.
The holotype, allotype, and paratypes are in mv collection.
Micracis torus, n. sp.
This species is distinguished from the allied evanescens Wood by the smaller size, by the stout, hairlike elytral setae, and by the absence of a sharply defined lower margin of the female frontal elevation.
Female.— Length 2.1 mm (paratypes 1.8-2.2 mm), 3.1 (male 2.7) times as long as wide; color dark reddish brown.
Frons planoconvex to vertex, median half on lower half elevated and brightly shining, its margins rounded not sharply defined, re- maining area minutely reticulate-granulate; ves- titure fine, short, unifonnly distributed on upper area, slightly longer toward vertex. Antenna about as in evaiiescem except hair on scape sUghtly shorter.
Pronotum 1.2 times as long as wide; as in evancscetis except reticulation in posterior areas finer.
Elytra 1.8 times as long as wide, 1.4 times as long as pronotum; outline as in evanescens; striae not impressed, punctures small, shallow; interstriae about three times as wide as striae, almost smootli, punctures very fine, uniseriate. Declivity steep, convex, apex produced; strial punctures reduced, obscure; interstrial punc- tures small, indistinct. Vestiture of rows of fine, short, strial hair, and rows of longer, coarse, pointed, interstrial hair; slightly longer on de- clivity, longest interstrial setae equal in length to distance between rows, all setae somewhat confused on declivity.
Male.— Similar to female except body stout- er; frons convex, a transverse impression on lower third, elevation absent, surface with sev- eral rounded granules.
Type Locality.— Vol can Colima, Jalisco, Mexico.
Type Material.- The female holotype, male allotype, and 17 paratypes were taken at the type locality on 23 June 1965, 2500 m elevation, No. 102, from branches of a leguminose tree, by S. L. Wood.
Biological Sehies, Vol. 15, No. 3 Ne\v Neothopical Scoi.ytidae
27
The holotype, allotype, and paratypes are setae sometimes abraded and appear as short in mv collection. scales.
Micracis exilis, n. sp.
This species is more closely allied to cosfari- censis Wood than to others in the genus, but it is distinguished by the much smaller size, by the more abundant, longer vestiture on the female vertex, by the smaller, smooth, shining area on the female frons, and by the much finer sculp- ture on the pronotum and elytra.
Female.— Length 1.9 mm (paratypes: fe- males 1.7-1.9 mm; males 1.3-1.7 mm), 3.3 (male 2.7) times as long as wide; color moderately dark yellowish brown, vestiture pale.
Frons rather shallov/ly concave to upper level of eves, an acute median carina on upper half; frons smooth and shining on median half below level of antennal insertion, obscurely reticulate and minutely punctured over remain- ing area; vestiture of short, coarse, subplumose setae over central reticulate area becoming much longer on upper lateral margins and very long on vertex, tips of longest setae on vertex could reach below middle of frons. Antennal scape rather broadly triangular, ornamented by long setae; club moderately large, 1.3 times as long as wide; suture 1 not reaching middle of club.
Pronotum 1.3 times as long as wide; sides straight and parallel on basal two-thirds, rather broadly rounded in front; anterior margin un- anned; summit well in front of middle, asperi- ties fine; posterior area reticulate, punctures sparse, fine, obscurely, minutely granulate. Ves- titure of a few coarse setae.
Elytra 1.9 times as long as wide, 1.45 times as long as pronotiun; sides straight and parallel on basal three-fourths, rather abruptly rounded then narrowly acuminate at apex; striae not impressed, punctures small, shallow, rather ob- scure; interstriae slightly more than twice as wide as striae, minutely irregular, punctures small, obscure. Declivity very steep, convex, acuminately produced at apex; surface toward apex very minutely subrugulose; interstriae each with a row of fine granules. Vestiture of short strial hair, and rows of moderately long, coarse, interstrial hair.
Male.— Similar to female except slightly smaller, stouter; frons convex above, a moderate transverse impression on lower third, surface rugose-reticulate, vestiture sparse, short, incon- spicuous; scape small, sparsely pubescent; an- terior margin armed by six small teeth; declivital
Type Locality.— Nine km south of Barran- cas, Barinas, Venezuela.
Type Material.— The female holotype, male allotype, and 11 paratypes were taken at the type locality on 2 December 1969, 150 m eleva- tion. No. 163 from Espinito de Sabana (type). No. 155 from Acahjpha sp.. No. 186 from a tree twig, by S. L. Wood. The gallery systems were typical of the genus.
The holotype, allotype, and paratypes are in my collection.
Micracis vitulus, n. sp.
This species is allied to lignicohts Wood, but it is distinguished by the large size, by the smaller, less pubescent female scape, by the larger antennal club, by the larger, much deeper strial punctures, by the hairlike elytral vestiture, and by the very different elytral declivity in both sexes. This species superficially resembles some species of Hylocurus.
Female.— Length 3.6 mm (paratypes 3.2-3.6 mm), 3.6 times as long as wide; color very dark reddish brown, parts of pronotum often lighter.
Frons as in lignicohis. Antennal scape broad- ly club-shaped, at least twice as long as wide, moderately ornamented by rather long hair; club 1.5 times as long as wide, suture 1 extend- ing beyond middle.
Pronotum 1.2 times as long as wide; outline about as in lignicolus; anterior margin unarmed; posterior area strongly reticulate, with rather small, isolated, low, rounded granules. Sparse vestiture limited to margins.
Elytra 2.2 times as long as wide, 1.9 times as long as pronotiun; sides straight and parallel on basal three-fourths, strongly produced and narrowly mucronate behind; striae weakly to moderately impressed toward declivity, punc- tures rather large, deep; interstriae smooth, shining, convex near decUvity, as wide as striae, punctures small, uniseriate, granulate at margin of declivity. Declivity moderately abrupt, broad- Iv convex, moderately steep, resembling some male Hylocurus; striae more strongly impressed; interstriae all irregularly sculptured, a few granules at base, 1 moderately elevated, gran- ules rather coarse, continumg almost to apex, 2 somewhat convex, moderately impressed, devoid of granules, 3 slightly elevated, rather finely, irregularly subserrate, 4-8 with several granules below base, 9 moderately elevated, joining cos-
28
Bricham Young University Science Bulletin
tal margin. Vestiture c-ontiiicd to declivity, of coarse, moderately abundaiit. rather long hair in interstrial rows. Basal half of front tibia slightly narrower than in li^nicohis; posterior face devoid of tubercules.
M.VLE.— Similar to female except frons as in male li<^nicolus; anterior margin of pronotum with about four indefinite, small serrations; a circumdeclivital ring of rather well-developed, blunt tubercles (largest in this genus); decUvity more nearly flat, interstriae 1 higher, 2 lower, tubercles reduced except for a few toward base, 9 more acutely elevated.
Type Locality.— Carbonera Experimental Forest, about 50 km (airline) northwest of Meri- da, Venezuela.
Type Material.— The female holotype, male allotype, and 13 paratypes were taken on 10 November 1969, 2500 m elevation. No. 128, from an unidentified tree seedling, by S. L. Wood. The galleries were typical of the genus.
The holotype, allotype, and paratypes are in my collection.
Micracls sentus, n. sp.
Except for the antenna and protibia this species would be placed in the genus Htjlocunts because of the coarse clytral sculpture. It is remotely allied to lionicohis Wood, but it is distinguished by the coarse spines on the male interstriae 1 and 3 and at the base of the de- clivity on 1-9.
Male.— Length 2.2 mm, 2.7 times as long as wide; color rather dark reddish brown.
Frons convex, a narrow, transverse impres- sion just above epistoma; surface rather coarsely granulate-punctate, a rather large median tuber- cle just above level of antennal insertion; ves- titure of rather coarse, short bristles over entire surface. Antennal scape moderately long, rather slender; club moderately large, ovate, 1.4 times as long as wide, sutures narrowly, strongly procurved, 1 reaching middle.
Fronotnm about as long as wide; sides weak- ly arcuate and subparallel on basal half, broadly rounded in front; anterior margin armed by 10 coarse teeth; summit slightly in front of middle; posterior area deeply reticulate, with moderately coarse, isolated granules bchiinl summit to base. Vestiture of rather sparse, short scales.
Elytra 1.6 times as long as wide, 1.6 times as long as elytra; sides straight and parallel on basal three-fourths, subacutely angulate behind; striae not impressed, punctures rather small.
deep; interstriae shining, almost smooth, almost twice as wide as striae, punctures moderately coarse, uiiiseriate, widely spaced, replaced at declivital base by rather large, rounded gran- ules. Declivity steep, convex, produced at apex; surface dull, densely mgulose-reticulate; striae almost obsolete; even-numbered interstriae each with two rounded nodules at base of declivity, tubercles on odd-numbered interstriae slightly larger, becoming pointed and rather widely spaced on declivity, those on 1, 3, and 9 extend- ing below middle, only one or two supplemental tubercles at base of 5 and 7; interstriae 2 weakly impressed; costal margin ascending rather strongly at apex. Vestiture of erect, interstrial scales mostly at base of dechvity and at tuber- cles on declivity, sparse on disc; each scale on disc two to four times as long as wide, longer and more slender on declivity.
Type Locality.— El Laurel Experimental Farm, 12 km southwest of Caracas, Venezuela.
Type Material.— The male holotype was taken at the type locality on 1 May 1970, 1300 m elevation. No. 460, from a branch of an un- identified tree, by S. L. Wood.
The holotype is in my collection.
Hijlocurus torosus, n. sp.
This species is very closely related to rudis LeConte but it is distinguished by characters of the frontal excavation and its protuberances as noted below.
Female.— Length 2.0 mm, 2.7 times as long as wide; color very dark brown.
Frons deeplv excavated on median two-thirds from just above epistomal margin to well above eyes, broadly oval in outline, its margins abrupt on all sides; surface etched as in nidis; lateral walls of concavity with a pair of large, protrud- ing, subcircular areas, almost as high as wide, pile longer than in rudis.
Antenna, pronotum, and elytra as in rudis.
Type Locality.— Florence, South Carolina.
Type Material.— The female holotv'pe was taken at the type locality on 12 May 1961, in flight, by V. M. Kirk.
The holotype is in my collection.
Hijlocurus verrucosus, n. sp.
Thus species is allied to noduhis Wood, but it is distinguished by the smaller size, by the dull, reticulate elytral declivity with the de- clivital strial punctures largely obsolete, and by
Biological Series, Vol. 15, No. 3 New Neothopicai. Scolytidae
29
the di.stinctly elevated declivital interstriae 9 with a large tubercle at the apex of the eleva- tion.
M.\LE.— Length 1.5 mm (paratypes 1.4-1.8 mm), 2.5 times as long as wide; color black.
Frons convex, a transverse carina on median half just below upper level of eyes; surface reticulate-granulate, a few fine, obscure punc- tures below, sparse, low, elongate, shining gran- ules abo\e; \estitiire inconspicuous. Antennal club as in tiochihis except middle third of suture 1 totally obsolete.
Pronotum as in noduhis except teeth on an- terior margin much smaller; posterior area more finely sculptured, with fewer granules. Half of setae on disc scalelike.
Elvtra 1.5 times as long as wide; outline as in noduhis except for a large tubercle on inter- striae 9; disc as in noduhis except interstrial nodules slightly narrower, perhaps slightly higher. Declivity slightly steeper than in nodu- his; surface finely reticulate-granulate; strial punctures obsolete except at base; interstriae 2,
4, 6, and 8 with tubercles only at base, on 1, 3,
5, and 7 tubercles extend about one- third de- clivital length from base, 3 weakly elevated and with a moderately large tooth at middle, 9 slightly more strongly elevated, its elevation ending at middle of declivity in a coarse, point- ed tooth. Vestiture of minute, fine, strial hair, and interstrial rows of scales; scales on basal half of disc short, becoming much longer to- ward and on upper half of declivity, scales much wider than in noduJus.
Fem.\le.— Similar to male except frontal car- ina shorter; interstrial nodules poorly developed; interstrial scales more slender.
Type Locality.— Nine km south of Barran- cas, Barinas, Venezuela.
Type Material.— The male holotype, female allotv-pe, and 3-3 paratypes were taken at the type localitv on 5 November 1969, 150 m eleva- tion. No. ill, from cut Inga branches, by S. L. Wood. The galleries were typical of the genus.
The holotype, allotype, and paratypes are in my collection.
Htjlocurus dilutus, n. sp.
This species is allied to simplex Blandford and ruber Wood, but it is distinguished by the smaller average size, by the smaller interstrial tubercles in both sexes, and by the small, con- fused, strial punctures on the declivity.
Male.— Length 2.1 mm (paratypes 2.1-2.5
mm), 2.6 times as long as wide; color rather dark reddish brown.
Frons convex, transversely impressed on lower third; surface obscurely reticulate-granu- late, convex area with rather numerous, very large, low, shining granules over more than central half; vestiture fine, sparse, inconspicu- ous except more abundant along epistoma. An- tennal club broadly oval; sutures 1 and 2 strong- ly procurved, 1 almost reaching middle.
Pronotum LI times as long as wide; as in simplex except sides weakly arcuate, anterior margin anned by eight coarse serrations, pos- terior area more strongly reticulate with isolated granules slightly smaller.
Elytra 1.5 times as long as wide, 1.4 times as long as pronotum; outline as in simplex; striae not impressed, punctures moderately large, rather deeply, not sharply impressed; in- terstriae ahnost smooth, with a few obscure, transverse lines, evidently narrower than striae, punctures not clearly defined; each interstriae near declivity with about two low, poorly form- ed granules. Declivity steep, convex, produced at apex; strial punctures small, deep, confused on lower two-thirds; interstriae 1 and 3 weakly elevated, 1 with very small granules on upper two-thirds, 3 with small granules on upper third, 9 rather strongly elevated on basal half, its summit subserrate, a rather broad impression between 9 and costal margin. Subglabrous; very sparse, short, fine hair on some specimens.
Female.— Similar to male except frons flat- tened on at least median half, transverse im- pression absent, surface reticulate, dull, with rather abundant, small, deep punctures and fine, short, abundant hair; serrations on anterior mar- gin of pronotum smaller, irregular; strial punc- tures on disc smaller, not as deep, shaiply de- fined; tubercles on declivity evidently slightly smaller.
Type Locality.— Volcan, Colima, Jalisco, Mexico.
Type Material.— The male holotype, female allotype, and 12 paratypes were collected at the type locality on 23 June 1965, at 2500 m eleva- tion. No. 102, by S. L. Wood, from a branch of a leguminose tree.
The holotype, allotype, and paratypes are in my collection.
Ilylocurus disparilis, n. sp.
This species is allied to inaequali.s Wood, but it is distinguished by the exaggerated de- velopment of the male circumdeclivital spines.
30
BRIGHAM YoUNO UNlVEHSITi' SCIENCE BULLETIN
Male.— Length 2.2 mm, 2.5 times a.s long a.s wide; color very dark brown.
Frons, antenna, and pronotimi a.s in inaeqtial- is Wood except circuindeclivital spines longer, particularly on alternate odd-numbered inter- striae, spines 1 and 9 conspicuously longer than others; elytra! apex much more slender than in allied species. Dcclivital face with punctures on lower two-thirds of striae 1 and 2 very small, in rows, all other punctures coarse, deep, con- fused, becoming much larger toward base; inter- striae 3 not elevated, anned by two blunt tu- bercles.
Type Locality.— Zamorano, Morazan, Hon- duras.
Type Materl\l.— The unique male holotype was taken at the type locality on 18 April 1964, 700 m elevation. No. 558, from a branch of Ficus glabrata, by S. L. Wood.
The holotype is in my collection.
Hijlociirus dissidens, n. sp.
This species is rather closely allied to ab- errans Wood, but it is distinguished by the slightly smaller spines in the male circumde- clivital ring, by the more strongly convex male declivital face, by the more abundant pubes- cence on the female frons, and by the more strongly procurved suture 1 on the antennal club.
Male.— Length 1.9 mm (paratypes L9-2.2 mm), 2.7 times as long as wide; color dark reddish brown.
Frons as in aberrans. Antennal club with sutures moderately procurved. Pronotiun as in aberrans except granules on discal area slightly larger, punctures not indicated.
Elytra L6 times as long as wide; widest at base of dechvity, sides straight on more than basal three-fourths, abruptly, serrately rounded, then obtusely produced at apex; striae not im- pressed except near declivital margin, punc- tures moderately coarse, deep; interstriae almost as wide as striae, smooth, punctures small, sparse; each interstriae ending at margin of de- clivity in a coarse nodule of equal length, 3-9 acutely pointed on their lateral margins and projecting slightly. Declivity abnipt, very steep, broadly convex, produced behind; strial punc- tures in rows, those near base of declivity larger; interstriae minutely granulate, 1 slightly elevated to apex and bearing about five small granules on middle third, 3 rather weakly ele- vated on upper two-thirds, with four closely set, acutely pointed teeth on middle third;
interstriae 9 acutely elevated, joining costal mar- gin, but with a very small notch at its apex as it joins costal margin. Glabrous except for a few minute, hairlike setae at declivital margin.
Female.— Similar to male except frons shal- lowly concave on central half, upper two-thirds of concavity with dense, erect, coarse, subplu- iiiose setae, surface minutely reticulate-granu- late, devoid of larger granules; anterior margin of pronotum more finely seiTate; strial punctures slightly smaller; circumdcclivital ring of nodules absent, declivity more strongly convex; declivital face similar, more finely sculptured, all inter- striae with a few minute granules near base, 9 rather weakly elevated, ending remotely from costal margin; vestiture rather widely distributed, sparse, coarse, moderately long.
Type LocALiTi.— Laguna Santa Maria, Na- yarit, Mexico.
Type Material.— The male holotype, female allotype, and 19 paratypes were taken at the type locality on 6 June 1965, 1000 m elevation, No. 197, from a large, square-stemmed vine, by S. L. Wood.
The holotype, allotype, and paratypes are in my collection.
Hylocurus viUifrotis, n. sp.
This species is remotely allied to dissidens Wood, but it is distinguished by the very differ- ent female frons, by the very different sculpture of the male declivity, and by other characters described below.
Female.— Length 2.6 mm (paratypes 2.0-2.6 mm), 3.2 (male 2.9) times as long as wide; color very dark reddish brown.
Frons flattened from epistoma to vertex, area above eyes larger and wider than area below; surface entirely concealed by vestiture; vestiture below level of eyes of abundant, coarse, erect, moderately long, subplmnose setae, increasing in length and apparently in thickness to upper margin, those on upper margin appearing as coarse, broad, ribbonlike setae each subdivided near its apex into about six hairlike fOaments, tips of these large setae extend more than half distance to epistomal margin. Antennal scape short, twice as long as pedicel, not expanded, ornamented by a small tuft of hair; club sub- circular; sutures broadly procur\ed, 1 reaching middle of club.
Pronotum 1.3 times as long as wide; sides almost straight and parallel on basal half, rather broadly rounded in front; anterior margin un-
Biological Series, Vol. 15, No. 3 Nevv Neotoopical ScoLvrmAE
31
armed; summit well in front of middle; posterior area shininsr, finely subrugose-reticulate, with fine, isolated granules behind summit almost to base. Vestiture confined to margins.
Elytra 1.9 times as long as wide; sides straight and parallel on basal three-fourths, acutely angulate behind; striae not impressed, punctures moderately coarse, rather deep; in- terstriae slightly wider than striae, almost smooth, shining, punctures almost entirely ob- solete except replaced near declivity by uni- seriate, moderately coarse granules. Declivity steep, convex, produced at apex; striae distinctly impressed, punctures not as deep as on disc; inters triae distinctly convex except near apex, each with a row of low, rounded tubercles on upper half, extending well below middle on 3 and 7, 9 moderatelv elevated and continuing \vithout interruption to costal margin. Vestiture confined to declivity, consisting of short, stout and long, slender scales of moderate abundance.
Male.— Similar to female except averaging shorter and stouter; frons convex and very coarsely deeply punctured on upper half, flatten- ed and very finely sculptured below, vestiture limited to lower area, fine, inconspicuous; an- terior margin of pronotum irregularly armed by about six small teeth; interstriae rather strongly convex at base of declivity, granules larger at base, largely obsolete on declivital face; interstriae 9 more strongly, acutely ele- vated; declivital vestiture evidently more abun- dant, scales distinctly wider.
Type Locality.— Rancho Grande, Pittier National Park, Aragua, Venezuela.
Type Material.— The female holotype, male allotype, and 21 paratypes were taken at the type localit)' on 9 April 1970, 1100 m elevation, No. 432, from a species of Guttiferae by S. L. Wood. Five paratypes bear identical data except No. 409, from a NIeleaceae; 37 paratypes are from El Laurel, 12 km southwest of Caracas, Venezuela, 1 May 1970, 1800 m elevation. No. 511, from tree branches by S. L. Wood; 13 para- types are from Colonia Tovar, Aragua, Venezu- ela, 4 May 1970, 1700 m elevation, No. 499, by S. L. Wood.
The holotype, allotype, and paratypes are in my collection.
Hylocurus singularis, n. sp.
This species is allied to viUijrons Wood, but it is distinguished by the subcarinate, transverse elevation of the male frons, by the concave,
finely pubescent female frons, and by the differ- ent elytral declivity.
Female.— Length 2.4 mm (paratypes 2.0-2.4 mm), 3.0 times as long as wide; color dark red- dish brown.
Frons rather deeply concave on circular area on about central half, center of impressed area at upper level of eyes; surface almost smooth, shining, finely punctured, vestiture very fine hairlike, rather abundant, of moderate uniform length. Antennal scape short, ornamented by a few setae; club wider than long, sutures broadly procurved.
Pronotum 1.2 times as long as wide; as in villifrons except posterior area very slightly more finely sculptured.
Elytra 1.9 times as long as wide; sides straight and parallel on basal three-fourths, acutely acuminate behind; striae feebly im- pressed, punctures large, moderately deep; in- terstriae ahnost smooth, weakly convex, as wide as striae, punctures almost obsolete except sub- granulate toward declivity. Declivity steep, con- vex; surface densely punctate-reticulate, striae obsolete or nearly so on lower three-fourths; interstriae 1, 2, and 3 weakly elevated, vWth rather large, rounded granules to lower fourth, 5 and 7 with similar granules to middle, 4, 6, and 8 with granules on upper fourth, 9 acutely, moderately elevated, not serrate, its crest fusing with costal margin. Vestiture limited to declivity where tubercles occur, consisting of rather short, stout, interstrial bristles.
MALE.-Similar to female except slightly stouter; frons broadly convex, with a thick, cal- luslike, transverse carina on median half im- mediately below upper level of eyes, vestiture sparse, limited to lower half; anterior margin of pronotum armed by eight rather coarse teeth; tubercles at basal margin of declivity somewhat larger, those on interstriae 2 and to a lesser ex- tent on 4 fused to form a short, acutely elevated, longitudinal crest; declivital tubercles slightly larger, usually pointed, interstriae 9 slighdy higher; declivital bristles flattened, much wider, slightly longer.
Type Locality.— Eight km southwest of Bumbum, Barinas, Venezuela.
Type Material.- The female holotype, male allotype, and 27 paratypes were taken at the type locality on 11 February 1970, 150 m ele- vation, No. 313, from an unidentified tree seed- ling (somewhat similar to Guava) by S. L. Wood.
32
Brigham Young University Science Bulletin
The holotype, allotype, and paratypes are in my collection.
Hylocurus flagellatus, n. sp.
This species is closely allied to dimorphiis (Schedl), but it is distinguished by the more elaborately decorated female scape, by the deeply, longitudijially sulcate female frons which bears a remarkable tuft of hair on the vertex, and by the flatter male declivity, with a dense covering of scales and shorter tubercles on the circumdeclivital ring.
Female.— Length 2.3 mm (paratypes; fe- males 2.3 mm, males 1.8-2.0 mm), 3.0 times as long as wide; color very dark brown.
Frons with median half deeply, broadly sul- cate from vertex to epistoma to accommodate a long, stiff pencil of apparently reddish fused setae arising on a small, circular area of vertex; surface reticulate, becommg subrugulose lateral- ly; other setae minute, recumbent, yellowish. Antennal scape broadly triangular, dorsal margin bearing a fringe of long, yellow hair, ventral margin near insertion of pedicel bearing a small penicillate tuft of reddish, fused hair consider- ably longer than length of pronotum, and from ventral margin near base a similar tuft of curi- ously curved hair equal in length to combined lengths of funicle and club; scape very similar to dimorplius but all tufts of hair longer; club 1.3 times as long as wide, sutures rather nar- rowly procurved, 1 not reaching middle.
Pronotum 1.4 times as long as wide; sides almost straight and parallel on basal two-thirds; anterior margin shallowly notched for reception of tuft of setae on vertex, unarmed; summit well in front of middle; asperities on anterior area low; posterior area reticulate, punctures minute, obscure. Vestiture of rather long, coarse, sparse hair.
Elytra 2.0 times as long as wide; sides straight and parallel on basal three-fourths, rather broadly rounded then subobtusely acu- minate beliind; striae not impressed, punctures small, shallow, obscure toward declivity; inter- striae as wide as striae, minutely somewhat irregular, punctures fine, obscure. DecHvity steep, convex; surface minutely granular; striae obsolete; interstriae 1 and 3 feebly elevated, small granules in rows on 1-3, scattered in later- al areas, 9 not separately elevated. Vestiture of fine, sparse hair on disc, of short strial and interstrial scales on declivity, scales rather abundant, each about two to three times as long as wide.
Male.— Similar to female except shorter, stouter; frons convex, without special ornamen- tation; antennal scape short, simple; anterior margin of pronotum rather narrowly rounded, anned by four teeth; elytra truncate behind, declivity very steep, almost flat; margin of declivity with circumdeclivital ring of interstrial tubercles, each wider than long; declivital striae evidently obsolete, interstriae with fine, evident- ly confused punctures, vestiture of abundant scales in obscure rows, each scale slightly less than twice as long as wide.
Type Locality.— Rancho Grande, Pittier National Park, Aragua, Venezuela.
Type Materl\l.— The female holotype, male allotype, and five paratypes were taken at the type locality on 9 April 1970, 1100 m elevation, No. 431, from Nectandra branches, by S. L. Wood. The gallery systems were typical of the genus.
The holotype, allotype, and paratypes are in my collection.
Stegomerus mirandus, n. sp.
This species is distinguished from the four previously described species by the small, con- fused, elytral punctures on both disc and de- clivity and by the hairlike vestiture on the ely- tral disc with no scales intermixed.
Female.— Length 1.4 mm (paratypes 1.2-1.5 mm), 2.5 times as long as wide; color very dark brown to black.
Frons rather narrow, convex, a small, trans- verse impression immediately above epistomal margin; surface reticulate, with moderately coarse punctures above, finer below; vestiture of sparse, inconspicuous, fine, long hair. An- tennal club large, subcircular, with three moder- ately procurved sutures obscurely marked by rows of fine, short hair.
Pronotum 1.0 times as long as wide; widest on basal third, sides moderately arcuate on basal half, weakly constricted in front of middle and behind indistinct anterolateral angles, rather narrowly rounded in front; anterior margin arm- ed by six small teeth ( variable ) ; anterior slope armed by low asperities; summit at middle; posterior area subreticulate, with small, isolated granules of irregular size to base. Vestiture of sparse, short hair.
Elytra 1.5 times as long as wide. 1.6 times as long as pronotum; sides almost straight and parallel on slightlv less than basal two-thirds, rather narrowly rounded behind; disc shining, all punctures small, moderately deep, confused.
Biological Series, Vol. 15, No. 3 New Neotropical Scolytidae
33
Declivity convex, rather steep; punctures slightly smaller than on disc, confused. Vestiture on disc of fine, short, hairlike setae; on declivity of abundant, rather short scales, each scale about four to six times as long as wide.
Male.— Similar to female except frons flatten- ed on lower half; reticulation on pronotal disc less evident.
Type Locality.— El Laurel Experimental Farm, 12 km southwest of Caracas, Miranda, Venezuela.
Type Material.— The female holotype, male allotvpe, and 57 paratypes were taken at the type localitA' on 1 May 1970, 1300 m elevation, No. 468, from an unidentified vine, by S. L. Wood.
The holotA'pe, allotvpe, and paratypes are in my collection.
Phacnjhts pruni, n. sp.
This species differs from bosqui Schedl and robtistus Schedl in having either two (male) or no (female) teeth on the anterior margin of the pronotum, in having the frons more strongly convex, and in having clearly defined striae and strial punctures.
Female.— Length 1.7 mm paratypes 1.4-1.8 mm), 2.4 (male 2.2) times as long as wide; color black, with whitish scales.
Frons broadly, evenly convex; surface reticu- late-granulate, a few small punctures almost obsolete; vestiture sparse, inconspicuous. Seg- ment 2 of antennal funicle as long as scape, larger than in other species.
Pronotum 0.85 times as long as wide; widest near base, sides arcuately converging to nar- rowly rounded anterior margin; summit be- hind middle; asperities on anterior slope narrow, rather small, on median third only; anterior mar- gin unarmed; entire surface finely reticulate- granulate, punctures fine, obscure. Lateral and posterior areas with moderately abundant, short scales and recumbent hair intermixed.
Elytra 1.5 times as long as wide, 1.9 times as long as pronotum; sides almost straight and parallel on basal two-thirds, rather narrowly rounded behind; striae not impressed, punctures small, deep; interstriae three times as wide as striae, smooth, shining, punctures abundant, fine confused. Declivity rather steep, rather broadly convex; striae weakly impressed, punc- tures smaller than on disc; interstriae feebly con- vex. Vestiture of abundant, confused, short scales of uniform length; each scale shghtly longer than wide.
Male.— Similar to female except anterior submargin of pronotum anned by two closely set teeth; pronotal asperities evidently very slightly larger; scales on elytra and pronotum evidently more abundant and shghtly longer.
Type Locality.— Carbonera Experimental Forest, about 50 km (airline) northwest of Merida, Venezuela.
Type Material.— The female holotype, male allotype, and 250 paratypes were collected at the type locality on 16 September 1969, 2.500 m elevation. No. 21, from Prunus sphaerocarpa, by S. L. Wood.
The holotype, allotype, and paratypes are in my collection.
Periocryphahts, n. gen.
This genus belongs to the Cryphalini. While a general description of characters might sug- gest a relationship to Ptilopodius Hopkins or other genera from the Indo-Malayan region, its true affinities probably lie more nearly to, but still remote from Phacnjlus Schedl and perhaps other South American Cryphalini. The emargin- ate eye, the 3-segmented antennal funicle, and the complete absence of sutures on the antennal club serve to distinguish it.
Description.— Length about 1.0 mm, about 2.3 times as long as wide; vestiture of scales and hair.
Frons simple, broadly convex; eye oval, emarginate, finely faceted; antennal scape rather short; funicle 3-segmented, segments 2 and 3 both narrow, their combined length shorter than pedicel; club flattened, subcircular in out- line, pubescent but totally devoid of indications of sutures, slightly shorter than combined length of scape and funicle. Pronotum wider than long, coarsely asperate on strongly declivous anterior slope, anterior margin armed by coarse teeth; basal and posterior third of lateral margin with a fine, raised line. Elytra rather strongly ascend- ing on costal margin toward apex; finely sculp- tured. Vestiture of scales and hair. Anterior tibiae with few teeth on outer margin on less than distal third. Presumably the male is of re- duced size and flightless.
Type-species.— Periocri/p/ia/us pxiUus Wood, described below.
Periocnjphalus jmllus, n. sp.
This species superficially might be confused with several other Cryphalini, but the emargin-
34
Brigham Young University Science Bulletin
ate eye, the 3-.segmented antennal funicle, the unscgmcntcd antennal club, and the siiiall, stout body serve to distinguisli it.
Female.— Length 1.0 inin (paratypes 1.0-1.1 mm), 2.3 times as long as wide; color almost black.
Frons rather strongly convex, weakly im- pressed above epistoma except on median line; surface minutely, somewhat obscurely rugose- reticulate, punctures rather coarse, moderately close, not clearly defined; vestiture of fine, short, sparse, inconspicuous hair. Eye less than one- third divided by a narrow emargination. Anten- nal scape rather short, slender; as described above.
Pronotum 0.91 times as long as wide; widest just behind middle, sides moderately arcuate on basal two-thirds, broadly rounded anteriorly but with median fourth slightly produced and armed by four closely set teeth, median pair much larger; summit slightly in front of middle; an- terior slope steep, rather coarsely asperate; posterior area subshining or shining, irregularly, very finely reticulate, part of lateral areas smooth, shining, punctures minute, indefinite, a few fine granules behind summit. Vestiture of short, suberect, fine and stout setae of mod- erate abundance.
Elytra 1.3 times as long as wide, 1.4 times as long as pronotum; sides almost straight and parallel on basal tvvo-thirds, rather narrowly rounded behind; striae not impressed, punctures minute, almost obsolete; interstriae minutely ir- regular, punctures uniseriate, minute, not always clearly defined, slightly larger than those of striae. Declivity confined to posterior third, con- vex, moderately steep; punctures evidently slightly larger than on disc, largely obscured by vestiture; costal margin distinctly flanged (or elevated) to accommodate rather strongly ascending abdomen. Vestiture of fine, very short strial hair and interstrial rows of erect scales; each scale about four times as long as wide, close; both scales and hair more abundant on declivity.
Protibiae with three small teeth on outer, apical m;xrgin.
Type LocALrri-.-Forty km southeast of So- copo, Barinas, Venezuela.
Type Mateiual.— The female holotype and 8 female paratypes were taken at the type locality on 2.5 January 1970. 150 m elevation, No. 274, from the vine knowai locally as Bejuco Blanco, by S. L. Wood; four female paratypes bear identical data except the collection No.
267, one additional female paratype is from No. 273, and one female paratype No. 250, from the twig of an unidentified tree. The pith tunnels in tiny stems were very similar to those made by certain Xi/losandrus species.
The holotype and paratypes are in my col- lection.
Ht/pothenemus naneUus, n. sp.
Apparently this species is more closely allied to puhescens Hopkins than to other representa- tives of the genus, but it is easily distinguished by the stouter body, by the more narrowly rounded apices of pronotum and elytra, and by the much larger antennal club.
Female.— Length 1.0 mm (allotype 0.75 mm, female paratype 1.0 mm), 2.2 times as long as wide; color rather dark yellowish brown.
Frons rather weakly convex, a small median pit or impression at upper level of eyes; surface finely rugose-reticulate; sparsely clothed with fine hairlike setae of moderate length. Antennal club large, 1.1 times as long as wide; eye 1.3 times as long as club.
Pronotum 0.9 times as long as wide; widest on basal third, sides moderately arcuate, rather narrowly rounded in front; anterior margin anned by six teeth, teeth .spaced by distances not greater than their basal width; simimit at middle, slightly impressed behind; posterior surface rather coarsely reticulate, with small seti- ferous granules of moderate abundance on disc and lateral areas. Vestiture consisting of small erect scales in posterior areas, small bristles in asperate area.
Elytra 1.3 times as long as wide, 1.5 times as long as pronotum; sides straight and parallel on basal half, then gradually arcuately converg- ing toward narrowly rounded apex; striae weak- ly impressed, punctures small, shallow; inter- striae about one and one-half times as wide as striae, feebly convex, subreticulate, punctures granulate, granules in uniseriate rows, rather coarse, moderately close. Declivity beginning at middle, convex; essentially as on disc. Vestiture consisting of uniseriate rows of erect interstrial scales and shorter, inconspicuous strial hair; each scale about twice as long as wide, spaced between rows by slightly more than length of a scale, and within rows by slightly less than length of a scale.
Male.— Similar to female except smaller and eye slightly reduced.
Type Locality.— Turrialba, Cartago Prov., Costa Rica.
Biological Series, Vol. 15, No. 3 New Neotropical Scolytidae
35
Type Material.— The female holotype, male allotype, and four female paratypes were collect- ed at the tvpe locality on 5 July 1963, at an elevation of about 800 m, by S. L. Wood, from dead twigs of a large, unidentified shade tree in a coffee plantation.
The holotype, allotype, and paratypes are in mv collection.
Ht/potheneintts a.scitiis, n. sp.
This species probably is more closely related to nanclhis Wood, described above, than to other representatives of the genus, but it is readilv distinguished by the larger size, by the narrower elvtral scales and the more abundant elytral ground vestiture, by the smaller, oval antennal club, and by other characters.
Female.— Length 1.3 mm (paratypes: males 0.9 mm, females 1.2-1.3 mm), 2.2 times as long as wide; color dark reddish brown, elytra almost black.
Frons convex above, slightly, transversely impressed above epistoma; surface coarsely re- ticulate above upper level of eyes, punctate and slightly granulate below; a weak median carina on lower third; vestiture sparse, rather short, hairlike, inconspicuous. Antennal club rather small, oval, 1.5 times as long as wide; eye 1.5 times as long as club.
Pronotum 0.9 times as long as wide; as in imncUus except teeth on anterior margin spaced, by distances at least twice basal width of a tooth, and posterior area densely, not clearly punctured, with scattered granules; vestiture of slender scales and bristles.
Elytra 1.2 times as long as wide, 1.5 times as long as pronotum; sides almost straight and parallel on basal two-thirds, rather narrowly rounded behind; striae moderately impressed, punctures rather coarse, deep; interstriae almost as wide as striae, somewhat convex, almost smooth, punctures uniseriate, rather close, min- utely subvulcanate. Declivity confined to pos- terior third, convex, rather steep. Vestiture con- sisting of uniseriate rows of interstrial scales; each scale about three times as long as wide; and shorter, rather abundant, fine, strial and in- terstrial hair.
Male.— Similar to female except smaller, eye partly reduced.
Type Locality-.— Puerto Viejo, Heredia Prov., Costa Rica.
Type Material.— The female holotype, male allotype, and 10 paratypes were collected at the
type locality on 12 March 1964, at an elevation of about 70 in by S. L. Wood, from just below the outer surface of living bark on the bole of a large, standing healthy tree at least 1 m in diameter.
The holotype, allotype, and paratypes are in my collection.
Hijpothcnemus teretls, n. sp.
This species has scalelike interstrial setae only on the basal half of the elytra; on the posterior half these setae are shorter, hairlike, and mav be semirecumbent. In this respect it resembles dipterocarpi Hopkins to which it is not closely related. Except for the frons, which is somewhat similar to jxirallchis Hopkins, and the elytra setae, this species resembles eruditus Westwood to which it is compared below.
Female.— Length 1.1 mm (paratypes 1.0-1.2 mm), 2.2 times as long as wide; color almost black.
Frons moderately, transversely impressed on lower two-thirds, convex above, with a low, distinct elevation at upper level of eyes occupy- ing median sixth; surface rugose-reticulate, punctures moderately coarse, shallow; vestiture inconspicuous.
Pronotum 0.94 times as long as wide; as in eruditus.
Elytra 1.5 times as long as wide; outline as in eruditus; striae not impressed, punctures fine, very shallow, seen with difficulty; interstriae subrugulose, at least twice as wide as striae, punctures fine, obscure, uniseriate. Declivity rather steep, convex; striae weakly impressed, otherwise as on disc. Vestiture of rows of very minute strial hair, and rows of interstrial setae; interstrial setae on disc erect, scalelike, each about four to six times as long as wide, these setae on posterior half of elytra represented by shorter, fine, slender, hairlike setae resembling strial setae.
Type Locality.— Finca La Lola, Limon, Costa Rica.
Type Material.— The female holotype and two paratypes were taken at the type locality on 10 January 1963, from Theobronw cacao, by J. L. Saunders. Other paratypes include: one from Beverley, Limon, Costa Rica, 26 August 1963, 7 m. No. 154, vine; one from Playon, San Jose, Costa Rica, 22 February 1964, 50 m, No. 449, Canavalia villosa; one from 5 km W EI Pino, Zulia, Venezuela, 20 October 1969, 10 m, No. 139, hif^a; five from 20 km SW El Vigia,
36
Bricham Young UNrvERSixv Science Bulletin
22 October 1969, 100 ni. No. 81, Cecropki peti- ole, and one 10 December 1969, No. 184, same locality; one from 9 km S Barrancas, Rarinas, Venezuela, 5 November 1969, 150 m. No. 101, Serjania; two, same locality, 1 October 1969, No. 33; two from 30 km E Palmar, Bolivar, Vene- zuela, 12 June 1970, 200 m. No. 575, Sterculia pruriens; and two from 40 km E Canton, Bari- nas, Venezuela, 8 March 1970, 70 m. No. 370, tree seedling; all were females and were taken by S. L. Wood.
The holotvpe and paratvpes are in my col- lection.
Cryptocarenus coroiuitus, n. sp.
This species is distinguished from diadema- tus Eggers by the larger size, by the very dif- ferent frontal sculpture, and by several other characters mentioned below.
Female.— Length 3.0 mm (paratypes 2.8-3.1 mm), 2.5 times as long as wide; color reddish brown.
Frons transversely impressed from upper level of eyes to epistomal margin, this area trans- versely flat, longitudinally concave; impressed area rather coarsely, closely punctured, more nearly granulate at upper limits of impression and on convex area above to vertex; upper mar- gin of impression with a small, median eleva- tion; epistoma distinct, not strongly elevated; vestiture inconspicuous except for epistomal brush.
Pronotum about as in diadematus except anterior slope steeper, asperities larger; anterior margin armed by eight to ten coarse teeth. Glabrous except at margins.
Elytra as in diadematus except declivity slightly steeper, more strongly convex, declivital interstriae 2 less strongly impressed, more nearly convex; vestiture more abundant, apices of setae wider.
Type LocALiTi'.— Twenty km southwest of El Vigia, Merida, Venezuela.
Type Material.— The female holotype was taken at the type locality on 21 November 1969, 50 m elevation. No. 150, from a Bignoniaceae vine, by S. L. Wood. Three female paratypes were takcMi in Venezuela, one from each of the following localities: 8 km SW Bumbum, Barinas, 11-11-70,' No. 325, Nectandra twig; 40 km SE Socopo, Barinas, 2.5-1-70, No. 2.55, from a cut tree seedling; Campamento Rio Grande, 30 km E Palmar, Bolivar, 20() m elevation, 12-VI-70, No. 566, Suipo; all by S. L. Wood. Pith tunnels in small stems were made by this species.
The liolotypc and paratypes are in my col- lection.
Cryptocarenus lepidus, n. sp.
This species is rather closely allied to lae- vigatus Blandford, but may be distinguished by the longitudinally carinate tubercle (often a complete carina from epistoma to vertex), by the shorter elytral vestiture, and by the lighter color.
Female.— Length 1.7 mm (female paratypes 1.6-1.9 mm), 2.6 times as long as wide; color rather dark reddish brown (some specimens very dark).
Frons rather weakly, transversely impressed from just above epistomal margin to just below upper level of eyes, convex above, a fine, low carina extending from epistoma to vertex but higher at upper level of eyes (not evident on lower half in several specimens); epistomal mar- gin shallowly, broadly emarginate; surface finely granulate-punctate, with several low, irregular, longitudinal rugae on upper two-thirds; vesti- ture consisting of rather sparse, fine hairlike setae.
Pronotum 1.0 times as long as wide; widest just behind middle, sides on ba.sal two-thirds rather weakly arcuate, constricted just behind anterior margin, rather broadly rounded in front; anterior margin armed by 11 closely placed teeth (seven to ten in most specimens); summit at middle; posterior area obscurely marked by fine points, lines, and a few punc- tures, subreticulate laterally. Anterior and later- al areas bearing a few short, stout hairs.
Elytra 1.5 times as long as wide, 1.6 times as long as pronotum; sides almost straight and parallel on basal two-thirds, rather narrowly rounded behind; striae not impressed except 1 feebly impressed toward declivity, punctures small, distinct, rather shallow; interstriae about twice as wide as striae, snbshining. surface obscurely marked by numerous minute points. Declivity rather steep, convex; striae 1 narrowly rather deeply impressed, 2 rather weakly, ir- regularly impressed, punctures smaller, less regular than on disc; interstriae 1 weakly ele- vated, surface more irregular; scattered punc- tures on all interstriae, a few of them very feebly granulate. Vestiture confined to sides and de- clivity, consisting of sparse, slender, spatulate bristles and a few minute, strial, hairlike setae.
Male.— Smaller than female. Only one male, with head and pronotimi lost, was available for study.
Biological Series, \^ol. 15, No. 3 New Neotropicai. Sccjlvtidae
37
Type Locality.— Beverley, Limon Prov., Costa Rica.
Host.— A woody vine, probably Serjanui sp. (type), Canavalia viUosa. and at least two tree .species (paratypes).
Type Material.— The female holotype and seven paratv'pes were collected at the type localitv on 26 August 1963, at an elevation of about 10 m, from a cut woody vine 1-3 cm in diameter. Other paratypes were taken in Costa Rica as follows: 6 from Pandora, Limon Prov., 23 August 1963, .50 m elevation, from a broken branch; 2 from Playon, 22 February 1964, and 2 from Volcan, 11 December 1964, Puntarenas Prov.; 7 from Santa Ana. .30 August 196.3, 1.300 m elevation and 1 from San Isidro del General, 13 December 1963, 1000 m elevation, San Jose Prov. One paratope was taken at Palin, Es- quintla Prov., Guatemala, 19 May 1964, 1000 m ele\'ation from a broken branch of a large tree. All were collected by S. L. Wood.
The holotype and paratypes are in my col- lection.
Dendrocranulus tardulus, n. sp.
This species is readily distinguished from tardus Schedl by the smaller size and by the distinctly impressed declivital interstriae 2.
Male.— Length 1..5 mm (paratypes 1.4-1.5 mm), 3.0 times as long as wide; color light brown.
Frons convex above eyes, rather shallowly, transversely impressed below; surface shining, closely, subgranulately punctured; vestiture very fine, long, rather sparse, hairlike.
Pronottun 1.2 times as long as wide; widest just beliind middle, sides moderately arcuate, rather broadly rounded in front; summit indefi- nite, well in front of middle; surface almost smooth, \\ith a few minute points, punctures on disc rather coarse, close, separated by distances about equal to their diameters, becoming some- what finely asperate in lateral areas. Vestiture consisting of rather sparse, erect hair.
Elytra 1.6 times as long as wide, 1.3 times as long as pronotum; sides almost straight and feebly dilated on basal two-thirds, then slightly narrowed to just before apex, and abruptly rounded to very broadly rounded, almost straight posterior margin; striae not impressed, punctures rather large, deep; interstriae slightly wider than striae, smooth and shining, punctures almost as large and deep as those of striae, separated by distances about equal to twice diameter of a
puncture. Declivity steep, almost flat; postero- lateral margin subacutely elevated from apex to interstriae 7, interstriae 2 almost flat, impressed, 1 and area lateral to 2 moderately elevated; strial and interstrial punctures somewhat smaller than on disc, those of interstriae very finely granulate. Vestiture consisting of erect, blunt, rather long interstrial bristles.
Female.— Similar to male except frons evenly convex and more closely pubescent; declivital interstriae 2 feebly if at all impressed, weakly convex, 1 and lateral areas less strongly ele- vated; vestiture finer.
Type Locality.— Rio Damitas in the Dota Mountains, San Jose Prov., Costa Rica.
Type Material.— The male holotype, female allotype, and 10 paratypes were collected at the type locality on 18 February 1964, at an eleva- tion of about 250 m, by S. L .Wood; the holo- type and two paratypes were taken from an unidentified vine, the others from Chayote.
The holotype, allotype, and paratypes are in my collection.
Dendrocranulus divermis, n. sp.
Tliis species appears to be somewhat vari- able, but it may be distinguished from costari- censis Schedl and other allied species by the more broadly impressed declivity, the impres- sion reaching interstriae 4.
MALE.-Length 2.7 mm (paratypes 2.0-2.9 mm), 2.5 times as long as wide; color very dark brown.
Frons convex above eyes, rather strongly transversely impressed from upper level of eyes to epistoma; epistomal margin broadly emargin- ate, upper tooth on mandibles projecting anteri- orly and partly filling this emargination; surface reticulate almost to epistomal margin, rather closely, deeply punctured, many punctures granulate; vestiture consisting of rather long, sparse hair.
Pronotum 1.1 times as long as wide; widest just behind middle, sides strongly arcuate from base to rather narrowly rounded anterior mar- gin; posterior surface almost smooth and shining, punctures moderately large, deep, close, each puncture with a very minute lateral granule (almost imperceptible at center), granules in- creasing in size laterally. Vestiture brisdelike, almost entirely limited to lateral and asperate areas.
Elytra 1.3 times as long as wide, 1.2 times as long as pronotum; sides arcuately narrowed
38
Hhigiiam Young Univehsitv Science Bulletin
from level of declivital base to level of sutural apex, posterior margin straight or feebly, broad- ly emarginate over about two-thirds of elytral width; striae 1 feebly, others not impressed, punctures small, close, rather deep; interstriae about twice as wide as striae, surface obscurely etched by minute points and lines, punctures slighth' smaller than those of striae and about half as numerous. Declivity rather abrupt, steep, broadlv, shallowly concave; posterolateral mar- gin obtusely rounded, strongly elevated from apex to interstriae 4, conspicuously higher than 1; interstriae 1 weakly elevated, 2 wider and im- pressed, all interstriae with a median row of fine granules; striae 1, 2, and 3 with punctures rather strongly impressed. Vestiture consisting of interstrial rows of erect bristles.
Female.— Similar to male except frons less strongly impressed (flat or weakly concave in some paratypes); in some specimens postero- lateral margin of declivity less strongly devel- oped.
Type Localiti-.— Puerto Viejo, Limon Prov., Costa Rica.
Type Material.— The male holotype, female allotype, and 14 paratypes were collected at the type locality on 12 March 1964, at an elevation of about 70 m, by S. L. Wood, from an uniden- tified vine. Other paratypes were taken in Costa Rica as follows: 4 at Pandora, Limon Prov., 23 August 1963; 3 at Turrialba, Cartago Prov., 9 March 1964; 16 at Rio Damitas in the Dota Mountains, San Jose Prov., 18 February 1964; and 5 at Playon, Puntarenas Prov., 22 February 1964; all were taken from unidentified vines by S. L .Wood. Three specimens not included in the type series are from La Lima, Honduras, 5 May 1964, in Cmjaponia microdonta.
The holotype, allotype, and paratypes are in my collection.
Dendrocranulus ptimihis, n. sp.
This species is very clo.sely related to schedU Wood, but may be distinguished by the reticu- late pronotum, by the smaller elytral punctures, and by the small nodules associated with discal punctures on the pronotum.
Male.— Length 1.3 mm (allotype 1.45 mm), 2.7 times as long as wide; color yellowish brown.
Frons convex above, transversely impressed one-third below upper level of eyes; surface reticulate, punctures coarse, deep; vestiture hairlike, rather sparse. Antennal club with sutures 1 and 2 weaklv procurved. 1 near mid- dle of club.
Pronotum evidently 1.3 times as long as wide (partly obscured by glue); widest just behind middle, sides moderately arcuate from base to rather narrowly rounded anterior margin; as- perities small, isolated, poorly developed; pos- terior surface reticulate, punctures coarse, mod- erately close, each with a minute granule on lateral side; vestiture hairlike, inconspicuous.
Elytra 1.9 times as long as wide, 1.4 times as long as pronotum; sides almost straight and parallel on basal two-thirds then gradually narrowed to broadly rounded posterior margin; striae not impressed, punctures coarse, deep; interstriae equal to or slightly narrower than striae, punctures about two-thirds as large as those of striae and slighdy less abundant. De- clivity steep, flattened between third striae; strial punctures almost as large and as deep as on disc; interstriae 1 moderately elevated, 2 impressed (at least medially), gradually as- cending laterally to interstriae 3; interstrial punctures less abundant than on disc, not at all granulate. Vestiture consisting of rows of erect interstrial brisdes (abraded on most of de- clivity ) .
Female.— Similar to male except frontal im- pression feebly developed; elytral declivity weakly, more narrowly impressed.
Type LocALrri.— Rio Damitas in the Dota Momitains, San Jose Prov., Costa Rica.
Type Matebial.— The male holotype and female allotj^pe were collected at the type lo- caUty on 18 February 1964, at an elevation of about 250 in, by S. L. Wood, from the stems of an unidentified vine.
The holotvpe and allotype are in my col- lection.
Dendrocniiudiis limtis. n. sp.
This species is distinguished from the closely allied turd us Schedl by the rather strongly con- vex elNtral declivity, by the less strongly ele- vated posterolateral margin of the declivity, and by the more widely spaced strial punctures.
Male.— Length 1.8 mm (paratypes 1.7-2.0 mm), 2.7 times as long as wide; color light red- dish brown.
Frons weakly convex; surface reticulate and rather coarsely, not deeply punctured, reticula- tion and punetiires reduced toward center on lower half; vestiture rather sparse, consisting of fine, long hair.
Pronotum 1.2 times as long as wide; sides of middle third almost straight and parallel,
Biological Series, Vol. 15, No. 3 New Neotropical Scolytidae
39
posterior angles broadly rounded, anterior angles arcuately converging toward broadly rounded anterior margin; dorsal profile arcuate from base, a little more strongly declivous on anterior third; surface shining, with few points, punc- tures rather coarse, deep, separated by distances equal to width of a punctiire, lateral margin of each puncture very finely asperate, tubercles becoming larger laterally. Vestiture hairlike, inconspicuous except laterally and in asperate area.
Elvtra 1.6 times as long as wde, 1.3 times as long as pronotuni; sides straight and parallel on slightly more than basal two-thirds, broadly rounded behind; striae not impressed, punctures large, deep; interstriae slightly wider than striae, almost smooth, shining, punctures two- thirds as large as those of striae, spaced one to two times diameter of a puncture. DecHvity steep, distinctly convex; posterolateral margin acutely elevated from apex to interstriae 7; striae weakly impressed; interstriae weakly convex, punctures reduced and feebly granulate. Wstiture consisting of erect, interstrial bristles.
Fem.\le.— Similar to male except frons bear- ing a rather dense fringe of long yellow hair to well above eyes, except central area glabrous.
Type LocALiTi'.— Rio Damitas in the Dota Mountains, San Jose Prov., Costa Rica.
Type Material.— Tlie male holotype, female allotype, and 19 paratypes were collected at the tv'pe locality on 18 February 1964, at an ele- vation of 250 m, by S. L. Wood, from stems of an unidentified vine; 22 paratypes were taken at Playon, Puntarenas Prov., 22 February 1964, at 50 m elevation, by S. L. Wood, from a wild cucurbit vine.
Xyleborus ebenus, n. sp.
This species is very closely related to san- guinicollis Blandford; it is readily distinguished from that species by the black pronotum, by the shining elytral declivity, with strial punctures small and clearly impressed. The declivital dif- ferences are conspicuous at a magnification of 80 diameters.
Female.— Length 3.8 mm (paratypes: fe- male 3.7-3.9 mm, male 3.1 mm), 2.0 times as long as wide; mature color uniformly black.
Frons as in sanguinicoUK except punctures reduced in number and size. Posterior area of pronotum as in sanguinicoUis but more uniformly reticulate, punctures less clearly impressed. Gen- eral sculpture of elytral declivity as in sanguini-
coUis; both species vary within limits but aver- age dentition about the same; strial and inter- strial punctures minute but clearly impressed, general surface between punctures appearing smooth on lower half, small points visible on upper half of declivity.
Male.— Slightly smaller than female; pro- notal asperities reduced, posteromedian part of pronotum gradually elevated to acutely pre- cipitous posterior margin; elytral declivity more gradual, upper denticles absent, lateral margin rather weakly elevated above major spine.
Type Locality.- GuapUes, Limon Prov., Costa Rica.
Type Material.— The female holotype and 15 paratypes were collected at the type locality on 22 August 1966, at an elevation of about 100 m, from a recently cut sapling. Other paratypes include: 2 from Turrialba, Cartago, Costa Rica, 9 March 1964, 700 m elevation; 3 from Cerro Campana, Panama, 26 July 1966, 1000 m ele- vation; 1 from Volcan Chiriqui, Panama, 11 January 1964, 1800 m elevation; 1 from Limon Bay, Canal Zone, Panama, 30 December 1963, sea level; all collected by myself; and 1 from Zant, Limon, Costa Rica, Flora E, Salas No. AB, 30 May 1956, at 57A4B, in Cacao.
The holotype, allotype, and paratypes are in my collection.
Mimips mimicus Schedl
Fig. 1
This species was described (Schedl, 1961, Pan Pacific Ent. 37:227) from specimens taken
Fig. 1. Mimips mimictis: posterolateral aspect of male elytral declivity.
40
BiircHAM Young UNrvERsixY Science Bulletin
at light in Costa Rica. Since then it has been taken from the followini^ locaUties. PANAMA: Barro Colorado Island, Canal Zone, 27-XII-63, 70 m elevation, No. .340, tree limb; Ft. Clayton, Canal Zone, 22-XII-63, .30 rn elevation. No.' 303, tree limb. VENEZUELA: 9 km S Barrancas, Barinas, ,5-XI-69, .50 in elevation. No. 11.5, Spon- dias mombin; 40 km E Canton, Barinas, 8-III-70, 70 m elevation, No. .394, Spondias mombin. All were taken by me; apparently they all came from the same host species. Its length is 1.6-2.0 mm.
Mhnips analogtis, n. sp.
Fig. 2
This species is almost identical to mimicus Schedl, but it is distinguished by the male frons and by other characters indicated in the fol- lowing description.
Fic. 2. Mimips analogus: posterolateral .aspect of male elvtral declivity.
Male.— Length 2.0 mm (paratypes 1.9-2.4 mm), 2.4 times as long as wide; color yellowish brown.
P'rons rather strongly convex above, lower third strongly, abruptly, transversely impressed; surface rather coarsely punctate-granulate; ves- titure sparse, fine, long. Antenna as in mimicus.
Pronotum 1.03 times as long as wide; as in mimicus except anterior margin unarmed and discal area smooth, shining (partly reticulate in mimicus.
Elytra 1.4 times as long as wide; as in mimicus except ventrolateral margin of declivity more evenly elevated, basal area of second tooth
usually much more strongly developed, and setae on declivita! interstriae 1 slender, longer, these setae stout and blunt in mimicus; punc- tures on elytral disc very slightly smaller.
Female.— Similar to female of mimicus ex- cept frons much more convex, with a slight, transverse impression above epistoma, its sur- face smoother, more sharply punctured, its ves- titure finer, not quite as long; anterior margin of pronotum unarmed; declivities differ as in males.
Type Locality. -Forty km E Canton, Ba- rinas, Venezuela.
Type Materlvl.— The male holotype, female allotype, and 30 paratypes were taken at the type locality on 8 March 1970, 70 m elevation, No. 394, from Spondias mombin, by S. L. Wood. They were taken from branches .3-8 cm in dia- meter; mimicus was in branches 2-3 cm in dia- meter in the same limb.
The holotype, allotype, and paratypes are in my collection.
Mimips fortis, n. sp.
Fig. 3
Evidently this species is allied to analogus Wood, but it is easily distinguished by the larger size and by the presence of only two pairs of teeth on the elytral dechvity.
Fic. .3. Mimips fortii: posterolateral aspect of male elytral declivity.
Biological Series, Vol. 15, No. 3 New NEomopiCAi. Scoi.ytidae
41
Male.— Length 2.5 mm (paratype.s 2.3-2.4 mm), 2.6 times as long as wide; color very dark brown.
Frons as in analogus except transverse im- pression much weaker; vestiture sparse, fine. Pronotiuii as in atuilogus except anterior margin indistinctly serrate, discal punctures slightly larger, not as close.
Elvtra 1.5 times as long as wide; disc as in flJUi/ogi/i- but punctures very slightly smaller, deeper; strial and interstrial punctures equal in size. Declivity very steep, tnmcate, broadly concave; lower two-thirds margined by an ele- vated, subacute, continuous, slightly undulating ridge, upper third armed by two coarse teeth (directed caudad), one in line with interstriae 2, second in line with interstriae 4; face of de- clivity shining, with moderately abundant, large, shallow punctures. Vestiture of rather long, fine, interstrial hair, a few supplemental setae near declivity.
Type Locality.— Finca Taboga, about 15 km southwest of Caiias, Guanacaste, Costa Rica.
Type Material.— The male holotype and tAvo male paratypes were taken at the type lo- calit)' on 8 February 1967, from a fallen tree. One male paratype was taken at Fort Clayton, Canal Zone, Panama, on 22 December 1963, pre- sumably from Spondias mombin, by S. L. Wood.
The holotype and paratypes are in my col- lection.
Mwiips hidens, n. sp.
Fig. 4
This species is somewhat intermediate be- tween chiriqucmis (Blandford) and /ort!,s Wood, but it is distinguished by the larger size, and by having only one pair of widely separated declivital teeth.
MALE.-Length 2.8 mm (paratypes 2.6-3.0 mm), 2.6 times as long as wide; color reddish browai.
Frons broadly convex, a rather large, low median elevation just below upper level of eyes; surface minutely subaciculate, with isolated, uniformly distributed, moderately large gran- ules; vestiture fine, sparse.
Pronotum as in fortis except discal surface subreticulate, punctures fine.
Elytra 1.4 times as long as wide; about as in jortis except strial and interstrial punctures rather small, deep, moderately confused from striae 3 to suture. Declivity steep, truncate, broadlv excavated; margin acutely, continuously,
strongly elevated on lower two-thirds, one coarse, blunt tooth in line with interstriae 4, upper margin between tooth and suture rounded; in declivital face suture moderately elevated, surface shining, punctures rather coarse, im- pressed, confused. Vestiture of sparse, moder- ately long, fine, interstrial hair, with supplemen- tary setae in declivital margin.
Female.— Similar to male except frons flat- tened, feebly impressed above, a feeble median elevation below, surface uniformly covered by small, isolated, setiferous granules, vestiture fine, long, abundant; declivital teeth displaced mesally, in line with striae 2, smaller, located slightly below upper margin of declivity.
Type Locality.— Eight km south of Colonia (near Buenaventura), Valle de Cauca, Co- lombia.
Type Material.— The male holotype, female allotype, and 102 paratypes were taken at the type locality on 9 July 1970, 30 m elevation. No. 628, in Icica altisima, by S. L. Wood.
The holotype, allotype, and paratypes are in my collection.
Mimips uncinatus, n. sp.
Fig. 5
This species evidently is closely allied to bidem Wood and chiriquensis (Blandford), but
Fic. 4. Mimips hidenx: posterolateral a.spect of male elvtral dedivitv.
42
Brigham Young University Science Bulletin
Fig. 5. Mimips uncinatus: posterolateral aspect of male
elytral declivity- it differs in the details of the frontal and elytral declivital sculpture.
Male.— Length 1.4 mm (paratypes 1.4-1.7 mm), 2.7 times as long as wide; color reddish brown.
Frons broadly convex; surface minutely aci- culate, with small, moderately sparse granules to upper level of eyes, a small, central fovea usually present; vestiture fine, long, moderately abundant. Pronotum as in hidens.
Elytra 1.7 times as long as wide; strial and interstrial punctures small, shallow, in rows, of equal size, surface almost smooth, shining. Declivity steep, truncate, broadly concave; low- er t^vo-thi^ds of margin subacutely elevated, ridge ending above in large tooth in line with interstriae 2, tooth directed mesad, slender at its apex and hooked toward suture; declivital margin rounded between teeth; concavity about as in fortis. Vestiture of short strial and long interstrial, fine hair, longer, coarser and with supplemental setae at base of declivity.
Female.— Similar to male except median third of vertex transversely etched as on a strid- ulating organ.
Type Locality.- Colonia Tovar, Aragua, Venezuela.
Type Matkkial.— The male holotype, female allotype, and 15 paratypes were taken at the type locality on 4 May 1970, 1700 m elevation,
No. 483, by S. L. Wood, from branches of an unidentified tree. Additional paratypes bear identical data except three arc No. 498, two are No. 495, 42 are No. 500, and one is No. .509.
The holotype, allotype, and paratypes are in my collection.
Mimips ocularis, n. sp.
Fig. 6
This small, conservatively sculptured species is not closely related to any described species. It is distinguished by the rather narrowly sep- arated eyes, and by other characters.
Female.— Length 2.0 mm (paratypes 1.7-2.0 mm), 2.8 times as long as wide; color reddish brown.
Frons convex above, flattened toward epis- toma; lower third subaciculate, coarse isolated granules in upper and lateral areas, shining, smooth between granules in upper areas; ves- titure very fine, long, rather sparse hair. Eyes very large, separated above by a distance almost equal to width of an eye.
Pronotum L14 times as long as wide; out- line as in mimicus; anterior margin finely ser- rate; posterior area rather coarsely, sharply punctured, with obscure indications of reticula- tions at margins of punctures, shining. Vestiture of fine hair at margins.
Elytra 1.7 times as long as wide, 1.4 times as long as pronotum; sides straight and parallel on basal three-fourths, broadly rounded behind; strial and interstrial punctures efjual in size, small, confused. DecHvity steep, narrowly, shal- lowly concave; margin obtusely rounded on low- er two-thirds, summit anned on upper two- thirds by three verv small, rounded granules, a
bic. 6. Miviips oculuris: posterolateral aspect of male elvtral dcilivitv.
Biological Series, Vol. 15, No. 3 New Neotropical Scolytidae
43
moderately large, pointed tooth in line with interstriae 2, mesad and very slightly above upper granule of lateral margin; concave area .shallow, transversely oval, with numerous small, confused punctures. \'estiture of moderately abundant, short, hairlike ground cover, and rather sparse, longer setae, long setae rather stout on declivity.
Male.— Similar to female except frons more sparsely pubescent.
Type LocALiTi'.— Eight km south of Colonia (near Buenaventura), Valle de Cauca, Co- lombia.
Type Material.— The female holotype, male allotype, and 19 paratypes were taken at the type locality' on 9 July 1970, 30 m elevation, No. 628, from Idea altisinm, by S. L. Wood. These specimens were in the same small tree with bidens.
The holotype, allotype, and paratypes are in mv collection.
stm
iJilosoma granulatum Blandford
The genus Stijphlosoirm Blandford ( 1904, Biol. Centr. Amer., Coleopt. 4[6]:232) was named from a unique male specimen from Vol- can de Chiriqui, Panama. This genus and spe- cies has remained almost unknown since its description except for references to the type. A long series was taken at Finca Taboga, about 15 km southeast of Cartas, Guanacaste Prov., Costa Rica, on 2 February 1967, by R. W. Mat- thews, from the bark of Spondias jrwmbin. Since the remarkable female has not been de- scribed the following will aid in the recognition of this species.
Female.— Length 1.3-1.5 mm, 2.4 times as long as wide; color very dark brown.
Frons flat from eye to eye from epistomal margin to vertex; epistomal margin very deeply emarginate, emargination almost attaining upper level of eyes, rather broadly U-shaped, wider than deep, its arms diverging slightly; surface almost smooth, with dense, uniformly distrib- uted, rather coarse, deep punctures over entire flattened area; dorsomesal angle of cutting sur- face of mandibles greatly extended and curved cephalad into a median, projecting process; this process projecting from fundus of epistomal emargination a distance equal to almost half length of scape; vestiture rather abundant, very short, inconspicuous. Antenna as described and figured by Blandford; funicle clearly 5-segment- ed; suture 1 moderately procurved, 2 very
strongly procurved, segments 1 and 2 smooth and glabrous except for rows of setae at sutures.
Styphlosonui subulatum, n. sp.
This species is very closely related to granu- latum Blandford, but it is distinguished by the shallowly impressed female frons with finer punctures, by the deeper, narrower female epis- tomal emargination, by the finer interstrial piuic- tures, and by the slightly narrower declivital impression.
Female.— Length 1.6 mm (paratypes 1.4-1.6 mm), 2,4 times as long as wide; color very dark brown.
Frons as in granulatum except feebly plano- concave, upper margin more abrupt; surface reticulate, punctures smaller, obscured in cen- tral area; sides of epistomal emargination more nearly parallel; mandibular processes very slightly longer.
Pronotum 1.1 times as long as wide; as in granulatum but very slightly more finely sculp- tured.
Elytra 1.4 times as long as wide; sides almost straight and parallel on basal two-thirds, very broadly rounded behind; striae 1 weakly im- pressed, strial punctures moderately coarse, rather deep, close; interstriae as wide as striae, smooth, punctures fine, close, in rows except slightly confused toward base (larger and much more strongly confused in granulatum). De- clivity very steep, appearing somewhat flattened from dorsal aspect; strial punctures smaller than on disc; interstrial punctures about as on disc; interstriae 2 narrowed, strongly, narrowly im- pressed on central two-thirds of decUvity. Vesti- ture on striae and alternate interstrial setae of short, suberect, rather coarse hair; alternate in- terstrial setae fomiing rows of slightly longer scales, each scale about three times as long as wide (alternate arrangement of hair and scales on interstriae of granulatum not at all regular ) .
Male.— Similar to female except frons with a strongly developed, transverse carina at upper level of eyes, convex above, strongly, transverse- ly impressed below carina, surface coarsely, closely punctured, upper area also reticulate.
Type Locality.— Nine km south of Barancas, Barinas, Venezuela.
Type Material.— The female holotype, male allotype, and 51 paratypes were taken at the type locality on 2 December 1969, 150 m ele- vation, No. 165, from Astronium graviolens, by
44
Brigham Young University Scienc:e Bulletin
S. L. Wcx)d. The flattened nuptial chamber is apparently standing on its edge in thick bark, with one axis parallel to the grain of wood, the other perpendicular to the cambium surface, but not reaching the cambium. From its inner angles one to four egg galleries branch, usually two each from the upper and lower angles; these angle diagonally from the nuptial cham- ber 2-4 mm to the cambium. The egg tunnels then continue along the cambium on a straight or slightly curved course. Eggs are deposited in niches and larval development starts in the cambium. Only newly hatched larvae were present.
The holotype, allotype, and paratypes are in my collection.
Dcndroterus eximius, n. sp.
Among described species this one is most nearly allied to texamis Wood, although the re- lationship is not close. It is distinguished by the much more finely punctured pronotimi and elytra, and by the impressed female frons.
Female.— Length 2.3 mm (paratypes 1.7-2.1 mm), 2.8 times as long as wide; color reddish brown.
Frons broadlv, transversely impressed on lower half, epistoma slightly elevated; surface reticulate above, becoming smooth below, upper area with small, moderately abundant, shallow punctures; vestiture moderately abundant, of rather long, fine hair. Antennal funicle with four strongly compressed segments; club longer than wide.
Pronotiim 1.2 times as long as wide; widest on basal half, sides weakly arcuate, narrowly rounded in front; anterior margin armed by about six teeth, median pair much larger; sum- mit at middle; asperities rather coarse, not con- tinued behind summit; posterior area reticulate, punctures very small, shallow, rather sparse. Vestiture rather sparse, hairlike.
Elytra 1.7 times as long as wide, 1.4 times as long as pronotum; sides almost straight and parallel on basal two-thirds, narrowlv rounded behind; striae not impressed, punctures rather small, shallow; interstriae wider than striae, rather obscurely reticulate, dull, punctures fine, very sparse except toward declivity. Declivity rather steep, convex; strial punctiu'es minute, weakly impressed; uiterstrial punctures replaced by fine, uniseriatc rows of granules. Vestiture largely confinetl to declivity, consisting of mi- nute strial hair and rows of much longer, erect interstrial hair; each hair stout, about as long
as distance between rows, spaced more closely withhi a row; all declivital setae diverge very slightly away from suture.
Male.— Similar to female except frons con- vex, weakly impressed immediately above epis- toma, surface very finely, obscurely punctured, vestiture sparse, inconspicuous.
Type Localit\-.— Lago Amatitlan, Esfjuintla, Guatemala.
Type Material.— The female holotype, male ;dlotype, and 67 paratvpes were collected at the type locality on 10 June 1964, 700 m elevation. No. 707, Bursera branch, by S. L. Wood.
The holotype, allotype, and paratypes are in my collection.
Dendroterus parilis, n. sp.
This species is very closely related to eximius Wood, but it is distinguished by the more strongly impressed female frons, by the deeper pronotal punctures, by the greatly reduced strial punctures, by the much more numerous, deeper, impressed points on the elytra, and by the much less abundant elytral hair.
Female.— Length 2.0 mm (paratype 1.9 mm), 2.8 times as long as wide; color very dark reddish brown.
Frons as in eximius except very slightly more strongly impressed. Pronotum as in eximius ex- cept discal reticulation stronger, punctures deep- er, more sharply impressed.
Elytra as in eximius except surface shining, strial punctures minute, shallow, some of them little larger than ;ibundant, sharply impressed, minute points; surface of disc with many shal- lowly impressed, irregular lines; interstrial punc tures on declivitv less distinctly granulate. Ves- titure about half as abundant as in exi7nius.
Type Locality'.— Santa Ana, San Jose, Costa Rica.
Type Material.— The female holotype and one female paratvpc were taken at the type locality on 8 November 196.3, 1.300 m elevation, No. 2.53, from a Bursera simarubra branch, by S. L. Wood.
The holotype and paratype are in my col- lection.
Dendroterus sodalis, n. sp.
This species is allied to eximius Wood, but it is distinguished bv the convex, subglabrous female frons, bv the subshining pronotum and elytra with sharply impressed, moderately large
Biological Series, Vol. 15, No. 3 New Neotropical Scolytidae
45
puncture.s, and bv the weakly impressed declivi- tal interstriae 2.
Female.— Length 1.7 mm (paratypes 1.4-1.8 mm), 2.6 times as long as wide; color dark red- dish brown.
Frons broadly convex; surface subshining, small, finely granulate punctures rather close, miifomily distributed; vestiture very short, hair- like, moderately abundant. Funicle 4-segmented.
Pronotum 1.2 times as long as wide; as in exi7nius except teeth on anterior margin much smaller; posterior area subshining, surface sub- reticulate with many impressed pomts, punc- tures moderately large, deep, sparse. Glabrous.
Elytra 1.6 times as long as wide, 1.4 times as long as pronotum; sides straight and parallel on basal two-thirds, rather broadly rounded be- hind; striae not impressed, punctures moderate- ly large, deep; interstriae abnost twice as wide as striae, almost smooth, shining, impunctate except on 1 and on others toward declivity. Declivity steep, broadly convex; strial punctures slightly smaller, not as deep; interstriae 1 abruptly, slightly elevated, 2 impressed, 3 weakly elevated, each with a row of fine rounded gran- ules except absent on lower half of 2. Vestiture of moderately long, coarse, rather widely spaced hair on and near declivity.
Male.— Similar to female except frons more narrowly convex, vestiture less distinct.
Type Locality.— Volcan de Agua, Esquintla, Guatemala.
Type Material.— The female holotype, male allotype, and 43 paratypes were taken at the type locality on 19 May 1964, 1000 m elevation, No. 595, from Biirsera, by S. L. Wood.
The holotype, allotype, and paratypes are in my collection.
Dendroteriis resohitus, n. sp.
This species is distinguished from the closely allied salJaei Blandford by the very different frons in both sexes, by the smooth, shining pro- notal disc with the punctures much more sharply defined, and by the finer elytral punc- tures.
Female.— Length 1.9 mm (paratypes 1.5-1.9 mm), 2.7 times as long as wide; color yellowish brown to almost black.
Frons moderately convex on upper two- thirds, lower third broadly, rather strongly, transversely impressed, epistomal margin shghtly elevated, a distinct, median tubercle on episto- mal margin; surface shining, punctures moder-
ately large, rather deep, sparse in central area, rather dense on lateral and upper margins, ex- tending well above upper level of eyes; vesti- ture of fine, long hair, sparse at center, rather abundant on margins. Antennal funicle 4-seg- mented; club 1.3 times as long as wide, sutures straight, 1 partly septate.
Pronotum 1.2 times as long as wide; as in sallaei except asperities distinctly larger; pos- terior area shining, deeply, coarsely, closely punctured. Hairlike vestiture short on disc, longer at sides, moderately abundant.
Elytra 1.5 times as long as wide, 1.3 times as long as pronotum; sides almost straight and parallel on basal two-thirds, broadly rounded behind; striae not impressed, punctures mod- erately small, deep; interstriae shining, about as wide as striae, punctures rather small, uni- seriate. Declivity steep, flattened, striae 1 and 2 moderately impressed, punctures as on disc; interstriae 1 convex, weakly elevated, 2 im- pressed but weakly convex, 3 weakly elevated; all interstriae with a row of fine, rounded tuber- cles, those on 2 smaller. Vestiture of rows of stout, moderately long, interstrial hair, much shorter on declivity.
Male.— Similar to female except frons con- vex to epistoma, punctures rather coarse, uni- formly distributed, vestiture sparse, evenly dis- tributed, epistomal tubercle absent.
Type Locality.— Playa del Coco, Guana- caste, Costa Rica.
Type Material.- The female holotype, male allotype, and 28 paratypes were taken at the type locality on 18 October 196.3, 15 m eleva- tion, from the bole of Bursera simambra, by S. L. Wood.
The holotype, allotype, and paratypes are in my collection.
Dendroteriis defectus, n. sp.
This species is not closely related to other de- scribed species of this genus. It is distinguished by the rather large size, by the unanned anterior margin of the pronotum, by the distinctive frons, by the small strial punctures, and by the sculp- ture of the interstriae.
Female.— Length 2.5 mm (paratypes 2.3-2.9 mm), 2.4 times as long as wide; color yellowish brown.
Frons broadly impressed from epistomal margin to well above eyes, flat on transverse axis, shallowly concave on longitudinal axis; sur- face very densely, rather coarsely punctured and
46
BiuciiAM Young University Science Bulletin
evidently very finely gnuiulate except smooth, impunctate and shining on median area along epistoma; vestiture of rather abundant, fine, moderately long hair of unifonn length. An- tennal funicle 4-segniented; club wider than long.
Pronotum 1.1 times as long as wide; widest at base, sides weakly arcuate, converging slight- ly on basal half, feebly constricted in front of middle, rather narrowly rounded in front, with a feeble, pseudoemargination on median line; anterior margin weakly elevated, not at all ser- rate; summit indefinite, behind middle; asperi- ties small, decreasing in size gradually from center, posterior area punctate-subasperate to base, punctures rather small, moderately deep, some reticulation present. Glabrous.
Elytra 1.3 times as long as wide, 1..3 times as long as pronotum; sides almost straight and parallel on basal two-thirds, broadly rounded behind; striae not impressed, punctures small, moderately deep; interstriae shining, twice as wide as striae, surface with many rather deep, irregular, subtransverse lines and moderately large, impressed points, punctures small, slight- ly irregular. Declivity rather steep, rather broadly flattened; striae slightly impressed, punctures deeper than on disc; flattened be- tween interstriae 4. Almost glabrous, a very few minute, hairlike setae on declivity.
Male.— Similar to female except frons strong- ly convex above eyes, a very strong, narrow, transverse impression on median three-fourths, area between impression and epistomal margin moderately inflated, glabrous, coarsely punc- tured; posterior area of pronotum distinctly less subcrenulate.
Type LocALiTi-.— Near Rincon de Osa, Pun- tarenas, Costa Rica.
Type Material.— The female holotype, male allotype, and 47 paratvpes were taken at the type locality on 11 August 1966, 50 m elevation, from Bursera simaruhra, by S. L. Wood. Fifty para- types were taken at Ft. Clayton, Canal Zone, Panama, 27 July 1966, 70 m elevation, from Bursera simaruhra, by S. L. Wood.
The holotype, allotype, and paratypes are in my collection.
Deruhoterus cognatus, n. sp.
This species is very closely related to nicxi- canus Blandford, but it is distinguished by the more slender form, by the more broadly, evenly roimded, more coarsely punctured male frons,
by the more convex female frons with shorter, less abundant vestiture, by the more coarsely punctured pronotum and elytra, and by the more distinctly impressed elytral declivity on which there are numerous minute, impressed points. Future collecting could necessitate the reduction of this form to a subspecies.
FEMALE.-Length 2.0 mm (paratypes 1.8-2.2 mm), 2.8 times as long as wide; color black.
Frons broadly, moderately concave to epis- toma; surface rather coarsely, evenly granulate- punctate; vestiture finer, shorter, about half as abundant as in mexicanus.
Pronotum 1.23 times as long as wide; about as in nwxicanus except punctures slightly larger, deeper.
Elytra 1.67 times as long as wide; outline as in mexicanus but much more broadly rounded behind; strial punctures coarser and deeper than in mexicanus, interstriae narrower than striae. Declivity slightly steeper than in mexi- canus, very slightly more strongly impressed, sculpture as in mexicanus except minute im- pressed points larger and much more numerous. Vestiture averaging longer and coarser than in mexicanus.
MALE.-Similar to female except frons as in male itiexicanus but more broadly, evenly con- vex, more coarsely punctured, with lateral epi- stomal calli only moderately to poorly developed.
Type Locality.— Five km ( 3 miles ) west of El Salto, Durango, Mexico.
Type Material.— The female holotype, male allotype, and eight paratypes were taken at the type locality on" 7 June 1965, 2500 m elevation, No. 1(W, from Bursera, by S. L. Wood.
The holotype, allotype, and paratypes are in my collection.
Phelloterus, n. gen.
This genus has the distinctive gallery pattern and habits, and the ]>road oral region of Styphlo- sonui Blandford and some of the elytral char- acters of Dendroterus Blandford, but it has the antennal club more nearly like Pittjophthorus Eichhoff, and an acutely elevated lateral and basal margin on the pronotum. It is distinguish- able irom Pityophthurus only by a combination of characters.
Descriition.— Length 1..3-2.2 mm, body rather stout; color reddish brown, vestiture of hair and or scales.
Frons convex in both sexes, epistomal area very broad, a premandibular lobe usually pres-
Biological Series, Vol. 15, No. 3 Nisrvv Neotropical Scolytii>ae
47
ent at least in female; mandibles enlarged, elongae in female. Eye finely faceted, emargi- nate. Antennal scape elongate; funicle .5-segment- ed; club subcircular, sutures 1 and 2 straight, septate at margins onlv, marked by setae at mar- gins onlv, apparently suture 3 procui-ved, mark- ed by setae just below apical margin. Pronotum about as wade as long, anterior half coarsely asperate; summit indefinite, transition from as- perate to smooth areas gradual. Elytra striate, discal striae 1-3 usually slighdy confused; inter- strial punctures very sparse; declivity steep, con\ex, usually narrowly bisulcate, punctures on striae 1 and 2 large and usually deep; inter- striae 3 in male coarsely dentate and with several large teeth or granules in lateral areas, females less strongly sculptured. Tibiae slender, as in Pittjoplithorus.
Type-Species.— Pliellotenis terstis Wood, de- scribed below.
Biology.— The polygamous species of this genus infest trees with thick phloem tissues. Their nuptial chamber is tabular, with the longi- tudinal axis parallel to the grain of wood and the transverse axis perpendicular to the cambi- um surface. Compared to other phloem-inhabit- ing genera the nuptial chamber stands on its edge. From one to five egg galleries branch from the inner margin of the nuptial chamber well before they reach the cambium region. Eggs and larvae occurred only in the cambium region. From one to four females were found with each male.
Phellofertis terstis, n. sp.
This species has minute tubercles on declivi- tal interstriae 1, and dechvital interstriae 2 flat and very strongly impressed; the sparse elytra! setae are almost hairlike.
Male.— Length 2,0 mm (paratypes 1.8-2.1 mm), 2.2 times as long as wide; color reddish brown.
Frons very strongly convex, rather strongly, transversely impressed just above epistoma, epi- stomal margin slightly produced in median area; surface shining, very coarsely, deeply punctured to well above upper level of eyes. Mandibles normal. Eye about twice as long as wide, emar- ginate; rather finely faceted. Antenna as de- scribed for genus.
Pronotum 1.03 times as long as wide; widest one-third length from base, sides moderately arcuate on less than basal two-thirds, narrowly rounded in front; anterior margin serrate; sum- mit indefinite, at or behind middle; posterior
area shining, with many minute, impressed points, punctures moderately large, deep, rather close. Glabrous except at margins.
Elytra 1.3 times as long as wide, 1.3 times as long as pronotum; sides almost straight and subparallel on slightly more than basal half, very broadly rounded behind, posterior oudine inter- rupted by coarse denticles; striae 1 feebly im- pressed, 1-3 moderately confused, punctures rather coarse, deep; interstriae smooth, shining, with rather numerous, minute, impressed points, punctures absent. Declivity very steep, narrow- ly sulcate; strial punctures very shallow; inter- striae 1 abruptly, moderately elevated, with about six widely spaced, fine granules, 2 as wide as 1 or 3, strongly impressed, flat, im- punctate, 3 abrupdy, rather strongly elevated and armed by five or six widely spaced, coarse, pointed teeth, 4-9 each with one to three denti- cles of various sizes, some as large as those on 3. Vestiture confined to declivity; of sparse, short, stout, interstrial bristles.
Female.— Similar to male except frons less coarsely punctured, premandibular process on epistomal margin small, rectangular, as long as wide, mandibles slightly elongate; declivital in- terstriae 2 less strongly impressed, granules on all interstriae much smaller.
Type LocALixY.-Colonia Tovar, Aragua, Venezuela.
Type Material.— The male holotype, female allotype, and .32 paratypes were taken at the type locality on 4 May 1970, 1700 m elevation. No. 507, probably Eshweilera log, by S. L. Wood.
The holotype, allotype, and paratypes are in my collection.
Phelloterus aiiaxeiis, n. sp.
This species is closely related to fersus Wood, but it is distinguished by the more gradual ely- tral declivity, in the male interstriae 1 is im- pressed, less strongly convex and devoid of granules, and by the elongate female mandibles.
Male.— Length 1.8 mm (paratypes 1.6-1.9 mm ), 2.4 times as long as wide; color yellow- ish brown.
Frons as in tersus except transverse impres- sion very weak. Antennal club very slightly wider than long. Pronotum as in terstis except ten teeth on anterior margin more clearly de- fined.
Elytra 1.4 times as long as wide; posterior margin rather narrowly rounded, declivital teeth
48
Biur.HAM YouNO University Sciknce Bi'li-Etin
not intfinipting outline; disc as in tersus except shorter. Declivity gradual, sulcate; interstriae 1 impressed and convex, unanned, 2 ascending toward 3 and bearing a row of coarse punctures, 3 elevated and amied by three coarse, widely separated teeth; lateral denticles positioned as in tersus but smaller. Vestiture much stouter.
Female.— Similar to male except frons more finely punctured, premandibular epistomal lobe larger, twice as wide as long, mandibles greatly enlarged and elongate; elytral declivity very shallowlv sulcate, interstriae 2 convex, denticles as in male but slightly smaller.
Type LoCALrrv.— Eight km south of Colonia (near Buenaventura), Valle de Cauca, Colombia.
Type Material.— The male holotype, female allotype, and 49 paratypes were taken at the type localitv on 9 July 1970, 30 m elevation, No. 609, from Saco^lotliia procera, by S. L. Wood. Seventy-four paratypes bear identical data except they were No. 614, taken from Couma macrocarpa-, three paratypes bear iden- tical data except No. 612, taken from Licania sp.; eleven paratvpes bear identical data except No. 611, taken from Leajthia sp. Forty-eight paratypes were taken 27 km northeast of Mon- tova (Campamento Capote) on 2 July 1970, 150 m elevation, No. 589, Eshweilera (5), No. .586 in Sapotaceae ( 28 ) , and No. 587 in an unidenti- fied tree limb (15).
The holotype, allotype, and paratypes are in my collection.
Phellote nts atrocis, n. sp.
This species is closely related to anaxeus Wood, but it is distinguished by the smaller size, by the conspicuous transverse impression above the epistoma in both sexes, and by de- tails of sculpture on the elytral declivity.
Male.— Length 1.3 mm. (paratypes 1.3-1.6 mm), 2.4 times as long as wide; color yellowish browii.
Frons and antenna as in tersus. Pronotum and elytral disc as in atuixeus. Declivity mod- erately steep, rather broadly, shallowly sulcate; as in anaxeus except interstriae 2 more deeply impressed. Declivital vestiture with rather abundant ground cover of short, stout, almost scalelike interstrial setae and very fine, short, recumbent, strial hair; a few longer interstrial bristles also arise from the tubercles and latend areas.
Female.— Similar to female of anaxeus ex- cept mandibles slightly smaller; impression on
elytral declivity not as deep; declivital tubercles absent, a few minute granules present on all interstriae.
Type Locality.— Campamento Capote, 27 km northeast of Montoya, Santander, Colombia.
Type Matebl\l — The male holotype, female allotype, and 122 paratypes were taken at the type localitv on 2 July 1970, 150 m elevation. No. 589, Eshweilera, sp., S. L. Wood. Eleven paratypes bear identical data except No. 586, in Sapotaceae; 29 paratypes bear identical data ex- cept No. 587, in a limb of an unidentified tree. Twenty-five paratypes are from 8 km south of Colonia (near Buenaventura), 9 July 1970, 30 m elevation. No. 609, Sacogjothia sp., S. L. Wood. All series were associated in the same branches with anaxeus.
The holotvpe, allotype, and paratypes are in my collection.
Pitijophthorus arceiithobii, n. sp.
In Blackman's ( 1928 ) classification of the Pitijophthorus this species would be placed close to caclator Blackman, but the relationship to that or to named Mexican species is remote at best. From caelator it is distinguished by the very different frons in both sexes, by the much coarser pronotal and elytral punctures in both sexes, and by the shallower sulcus on the elytral declivity which is almost totally devoid of gran- ules on the lateral convexities.
Female.— Length 1.8 mm (paratypes 1.8-2.0 mm), 2.6 times as long as wide; color very dark brown, elytra usually shghdy lighter.
Frons planoconvex, with a slight transverse impression on lower third, epistomal margin weakly elevated; surface subshining, rather coarsely, closely punctured, on a hemispherical area; vestitiire of fine, rather long hair of equal length almost unifonnly distributed over punc- tured area. Antennal club with suture 1 straight, segments 2 and 3 about equal in width.
Pronotum 1.0 times as long as wide; widest near base, sides almost parallel on slightly less than basal half, weakly constricted on anterior half, rather broadly rounded in front; anterior margin anned bv about ten, small, irregularly formed teeth; posterior area reticulate, punc- tures coarse, deep. Glabrous except at margins.
Ellvtra 1.6 times as long as wide, 1.7 times as long as pronotum; sides almost straight and parallel on basal two-thirds, then shghtly con- verging, broadly rounded behind; striae 1 nar- rowly impressed on posterior half of disc; striae
Biological Series, Vol. 15, No. 3 New Neothopical Scolytidae
49
moderately confused, punctures large, deep, only moderately close, equal in size and appearance with sliffhtlv less abundant, confused, inter- strial punctures; surface shining, not completely smooth. Declivit)' rather steep, broadly, shal- lowly sulcate; punctures on striae 1 and 2 mi- nute but distinct; intcrstriac 2 wide, smooth, moderately impressed, 1 sharply, slightly ele- vated, bearing about si.\ minute granules, 3 gradually, broadly raised, punctures very feebly granulate. \'estiture of sparse, rather short, coarse hair, mostly on sides.
M.\LE.— Similar to female except frons more strongly convex above, lower half with a rather thick, low, median carina which gradually in- creases in height to summit just above epistoma, surface subreticulate, less regularly punctured, frontal vestiture sparse, short; granules on elytral declivity scarcely evident.
Type Locality.— Ninety-six km (60 miles) west of Durango, Durango, Mexico.
Type Material.— The female holotype, male allotype, and .38 paratypes were taken at the t}pe locality' on 5 June 1965, 2500 m elevation. No. 27, from a dying mistletoe, Arceiithobium glohosum. growing in a 5-needle pine, by S. L. W'ood. This is the first representative of the Pitvophthorini reported from the Loranthaceae.
The holotype, allotype, and paratypes are in my collection.
Pittjoborus hondurensis, n. sp.
Tliis species is distinguished from the closely allied vehitinus Wood by the less strongly im- pressed female frons, with the yellowish pubes- cence finer, shorter and less abundant, by the much smaller pilose areas on the female pro- notum, by the more strongly impressed, more abundant, confused punctures on the elytral de- clivity, and by the more abundant vestiture on the elytral declivity.
Female.— Length 2.0 mm (paratypes L8-2.0 mm), 2.6 times as long as wide; color reddish browm.
Frons as in vehitinus except very slightly less strongly impressed on a narrower area, with fine, sparse punctures; lateral vestiture finer, less abundant, much shorter. Pronotum as in vehitin- us except pilose areas distinctly smaller; pos- terior area with moderately abundant, small, shallow punctures clearly evident.
Elytra as in velutinus except punctures on disc deeper, slightly larger, with a few more
interstrial punctures particularly on posterior lialf; declivital punctures smaller than on disc but much larger and deeper than in vehitinus, interstriae 1 distinctly elevated, striae and inter- strial punctures in somewhat indistinct rows; sparse vestiture on declivity more abundant than in veltttinus.
Male.— Similar to female except frons not at all concave, vestiture greatly reduced; pilose areas on pronotum absent; pronotal punctures less distinct; punctures on discal striae slightly larger; declivital interstriae 1 and 3 with minute granules.
Type Locality.— Yuscaran, Paraiso, Hon- duras.
Type Material.— The female holotype, male allotype, and 11 paratypes were taken at the type' locality on 2.3 April 1964, 800 m elevation, from small shaded-out branches of Pinus cari- haea (No. 518, type) and P. oocarpa (No. 519), by S. L. Wood. Galleries were typical of the genus.
The holotype, allotype, and paratypes are in my collection.
Pitijobonis frontalis, n. sp.
This species is distinguished from the close- ly allied vehitinus Wood by the more deeply concave female frons, with the frontal vestiture finer and pale yellow in color, and by the much larger, deeper strial punctures on the declivity.
Female.— Length 2.0 mm (allotype 2.0 mm), about 2.6 times as long as wide (elytra spread); color rather dark reddish brown.
Frons as in vehitinus except much more deeply concave on a narrower circular area; vestiture in lateral areas much finer, pale yellow in color. Pronotum as in vehitinus except reticu- lation stronger, punctures more clearly evident.
Elytra as in vehitinus except reticulation stronger, declivity more nearly shining; strial punctures on dechvity larger, deeply impressed.
Male.— As in female except frons not con- cave, feebly impressed, surface reticulate, vesti- ture on lateral and upper margins similar but slightly less abundant; pilose areas on pronotum absent.
Type Locality.— Thirteen km southeast of El Cameron, Oaxaca, Mexico.
Type Material.— The female holotype and male allotv'pe were taken at the type locality on 21 June 1967, No. 75, from a shaded-out branch of Pinus, by S. L. Wood.
50
Bhigham Young UwivEnsiTY Science Bulletin
The holotypc and allotype are in my col- lection.
Pseudopittjophthorus .nngularis, n. sp.
This species is allied to tenuis Wood, but it is distinguished by the coarser pronotal punc- tures, by the longer elytral setae, by the weakly impressed male frons with slightly longer and more abundant setae, and by the subconcave fe- male frons which lacks a median carina.
Male.— Length 1.6 mm (paratypes 1.3-1.6 mm), 3.0 times as long as wide; color very dark brown (most of type series not fully colored).
Frons flattened, epistoma weakly elevated, surface indistinctlv punctured; vertex bearing a sparse brush of about two dozen very long setae with tips reaching to epistoma, a few additional setae in epistomal area. Antenna as in tenuis.
Pronotiun much as in tenuis, with anterior margin more broadly rounded; punctures on disc much larger, impressed points moderately abundant.
Elytra l.(S times as long as wide; sides al- most straight and parallel on basal two-thirds, rather broadly rounded behind; striae not im- pressed, punctures small, distinct, in rows; in- terstriae almost smoodi, a few minute, impressed points, at least three times as wide as striae, a few punctures toward declivity. Declivity steep, rather broadly convex; punctures of striae 1 mi- nute, in a row, others obsolete; sutural inter- striae feebly elevated, 1, 3 and some of lateral interstriae with sparse, minute, setiferous punc- tures. Vestiture of very minute strial hair, and rows of widely spaced, moderately long, fine in- terstrial hair on sides and declivity, almost obso- lete on disc.
Female.— Similar to male except frons con- vex above eyes, transversely impressed above epistoma, impression extending dorsad slightly at center, surface very coarsely punctured, vesti- ture sparse, inconspicuous.
Type Locality.— One km west of Las Vigas, Veracruz, Mexico.
Type Material.— The male holotype, female allotype, and five paratypes were taken at the type locality on 5 July 1967, No. 159, from a Quercus branch, by S. L. Wood.
The holotype, allotype, and paratypes are in m\' collection.
Pseudopiti/ophthorus virilis, n. sp.
This species has the frons ornamented by setae in both sexes; it is remotely similar to
•^ninulifer Wood but it is distinguished by the smaller size, by the distinctly punctured elytral declivity, and by the presence of granules only on interstriae 1 and 3.
Male.— Length 1.5 mm (paratypes 1.3-1.6 mm), 2.7 times as long as wide; color black.
Frons transversely impressed, flattened from above eyes to epistoma; surface somewhat coarsely punctured above and at sides, evidently impunctate below in median area; vertex orna- mented by a rather dense brush, its longest setae exceed epistomal margin, a few additional setae at sides and on epistomal area. Antenna about as in granulifer.
Pronotum as in granulifer except teeth on anterior margin of pronotum larger, punctures on disc very slightly larger, impressed points much smaller.
Elytra 1.6 times as long as wide; outline as in granidifer; punctures on disc confused, not in rows except near declivity, small; surface sub- shining, with a few irregular lines and impressed points. Declivity steep, convex; interstriae 1 and 3 each bearing a row of fine, rounded granules, three indistinct rows of minute, seti- ferous punctures between rows of granules, lateral areas similarly punctured. Vestiture al- most hairlike at base, becoming scalelike to- ward declivity, those setae apparently derived from interstriae distinctly longer than those de- rived from striae; longest scales on declivity rather short, about six times as long as wide.
Female.— Similar to male in all respects, in- cluding frontal sculpture and vestiture.
Type Locality.— One km west of Las Vigas, Veracruz, Mexico.
Type Materl\l.— The male holotype, female allotype, and 19 paratypes were taken at the type locality on 5 July 1967, No. 159, from Quercus branches, by S. L. Wood.
The holotype, allotype, and paratypes are in my collection.
Pseudopittjophthorus declivis, n. sp.
This species is unique in the genus; it is dis- tinguished by the subvertical, impressed elytral declivity on which the striae are clearly indi- cated by rows of moderately coarse punctures.
Female.— Length 1.8 mm, 2.6 times as long as wide; color very dark brown.
Frons broadly convex above eyes, almost flat below, with epistomal area weakly elevated; surface very coarsely, deeply punctured above
Biological Series, Vol. 15, No. 3 Nenv Neotropical Scolyiidak
51
and at sides, almost impunctate in median area below; vestiture rather sparse except on epi- stoma, of fine, long hair. Antennal club moder- ately large, \videst through segment 3.
Pronotum 1.1 times as long as wide; widest near base, sides feebly arcuate and converging very slightly on basal two-thirds, rather broadly rounded in front; anterior margin armed by a continuous, elevated, serrate costa; summit at middle; anterior area rather finely asperate; posterior area rather coarsely, deeply, closely punctured, general surface subreticulate. Sub- glabrous.
Elytra 1.6 times as long as wide, 1.4 times as long as pronotum; sides almost straight and parallel on basal two-thirds, rather broadly bi- sinuate behind; striae not impressed, punctures rather fine (coarse for this genus), deep; inter- striae three times as wide as striae, with very fine, irregular lines, numerous impressed points, impunctate. Declivity subvertical, flattened, con- fined to less than posterior fifth; striae 1-3 clear- ly indicated, punctures shallow; surface almost fiat except for strongly elevated interstriae 1; very fine punctures on all interstriae except 2. Vestiture confined to posterior third, of very minute strial hair, and rows of erect, moderately long, interstrial bristles of uniform length.
Type Locality.— Laguna Santa Maria, Na- yarit, Mexico.
Type Material.— The unique female holo- type was taken at the type locality on 6 July 1965, 1000 m elevation. No. 203, from a Quercus branch, by S. L. Wood.
The holotype is in my collection.
Pseudopitijophthonis colombkinus, n. sp.
This unique species is the first representative of the genus known from South America; it is distinguished from other species by the very slender form, by the shallowly bisulcate elytral declivity on which the strial punctures are in rows, and by the rather coarse granules on in- terstriae 3.
Male.— Length 1.4 mm (paratypes 1.3-1.5 mm), 3.0 times as long as wide; color brovvni.
Frons flattened from eye to eye, from epis- toma to vertex; surface smooth, shining, with fine punctures (evidently only near sides; ob- scured), central area glabrous; lateral and upper margins with a dense row of long yellow setae, tips of those on vertex reaching epistomal mar- gin. Antennal club rather small; sutures 1 and 2 almost straight.
Pronotum 1.2 times as long as wide; widest on basal third; sides feebly arcuate, converging slightly on basal two-thirds, rather broadly rounded in front; anterior margin armed by more than a dozen small teeth; summit in front of middle; asperities usually in poorly defined subconcentric rows; posterior area almost smooth, punctures moderately coarse, rather sparse. Vestiture of short, sparse hair, except bristlelike in asperate area.
Elytra 1.9 times as long as wide, 1.6 times as long as pronotum; sides almost straight and parallel on basal three-fourths, broadly rounded Ijehind; striae not impressed, punctures rather coarse, moderately deep; interstriae almost smootli, as wide as striae, punctures almost as large as those of striae but almost entirely con- fined to interstriae 1 and to posterior fourth of disc. Declivity steep, broadly, shallowly bi- sulcate; strial punctures fine, shallow; sutural interstriae slightly elevated, 2 slightly, rather broadly impressed, 3 gradually, distinctly ele- vated and bearing a row of three to five mod- erately coarse granules; lateral areas with oc- casional smaller granules. Vestiture of moder- ately abundant, rather long, hairlike setae, some interstrial setae rather coarse.
Female.— Similar to male except frons con- vex, lower third with a distinct transverse im- pression, surface rather coarsely punctured, ves- titure sparse, inconspicuous.
Type Locality.— Piedras Blancas, 10 km east of Medellin, Antiotjua, Colombia.
Type Material.— The male holotype, female allotype, and 50 paratypes were collected at the type locality on 15 July 1970, 2500 m ele- vation. No. 667, from small Quercus humboldtii branches, by S. L. Wood. The galleries and habits were as in other species of the genus.
The holotype, allotype, and paratypes are in my collection.
Gnathophthorus clematis, n. sp.
Fig. 7
The only previously knowTi species in this genus, sparsepilosus (Schedl), is not closely related to any of the species described here. That species has the frons more unformly sculptured and pubescent than those presented below, but it shares the very slender form and the distinctive antenna and elytral declivity of these four species. Two of the species for which hosts are known breed in vines of the genus Cletrwti'i and have similar habits.
52
Brigham Young University Science Bulletin
Fig. 7. Gntithophthorus clematis: dorsal aspect of out- line of female (above); ;ind ;uiterolateral iispect of female head (Ix-low).
Female.— Length 1.9 mm (paratypes 1.4-2.0 mm), 3.8 times as long as wide; color light yellowish brown, pronotum and elytral declivity slightly darker.
Frons flattened from just above epistoma to vertex, transversely divided at upper level of eyes by a feeble summit; lower area coarsely, shallowly, closely punctured, with sparse, uni- formly distributed, inconspicuous, hairlike pubes- cence; upper area appearing somewhat spongy, very minutely, densely punctured, with micro- scopic pilelike pubescence and with a marginal fringe of short hair of unifonn length, little if any longer than setae on lower frons. Eye large, emarginate, coarsely faceted. Antennal funicle 5-segmented, last four segments crowded, in- distinct; club very large, with three moderately procurved, nonseptate sutures.
Pronotum 1.4 times as long as wide; sides on basal half straight and parallel, constricted slightly in front of middle then weakly arcuate to broadly rounded imterior margin; anterior margin amied by eight widely spaced, coarse teeth; summit on anterior third, indefinite; an- terior third rather finely asperate; posterior area smooth, with fine, deep, sparse punctures. Gla- brous except for scattered hair near lateral margins.
Elytra 2.4 times as long as wide, 1.8 times as long as pronotum; sides straight and parallel on more dian basal three-fourths, broadly round- ed behind; declivity confined to posterior one- sixth; striae not impressed, punctures minute, scarcely indicated on siu^ace; interstriae almost smooth, impunctate. Declivity steep, rather strongly excavated; striae obsolete; interstriae 1 moderately elevated, with a row of moderately coarse punctures; area of interstriae 2 strongly impressed, widened at middle; lateral areas strongly elevated on upper two-thirds, summit broadly rounded, with rather coarse, confused pimctures, two very feeble granules on upper margin of elevated area. Disc glabrous; moder- ately abundant, coarse hair on declivity.
Male.— Similar to female except frons convex above, lower half abruptly impressed, subreticu- late, punctures rather coarse, vestitiu-e sparse, inconspicuous; granules on elytral declivity larger, pointed.
Type LocALiTi.— Volcan Colima, Jalisco, Mexico.
Type Material —The female holotype, male allotype, and 106 paratypes were taken at the type locaUty on 23 June 1965, 2.500 m elevation. No. 100, from Clematis, by S. L. Wood. Other specimens, not included in the type series are from Tapanti and Volcan Irazu, Costa Rica; and Volcan Chiriqui, Panama. The monogamous beetles construct transverse, biramose galleries
Biological Series, Vol. 15, No. 3 New Neotropical Scolytidae
53
in the cambium region; egg niches are small and packed with frass; larval mines are straight, in grooves between the longitudinal fibers, up to 40 mm long, of uniform diameter; fungal growth was noted in the parental tunnels but not in the larval mines.
The holotype, allotype, and paratypes are in my collection.
Gnathophthorus cracens, n. sp.
This species is allied to clematis Wood, but it has the frontal pile much longer and covering a large area, and the declivital sulcus is much narrower and anned by several denticles.
Female.— Length 1.2 mm, about 3.8 times as long as wide; color yellowish brown.
Frons as in clematis except lower area flat, very finely, sparsely punctured, ending well be- low upper level of eyes; upper area slightly more extensive, pile long, dense, setae on mar- ginal fringe on upper margin only, much longer, tips reaching middle of pilose area. Pronotum as in clematis except anterior margin armed by 10 teeth; punctures on posterior area minute.
Elytra as in clematis except declivity nar- rowly, deeply sulcate; sutural interstriae feebly elevated; lateral areas abruptly elevated from position of striae 2, summit armed by five or six small, laterally compressed teeth.
Type Loc.\LiTi'.— La Ceiba, Honduras.
Type Matebial.— The female holotype was taken at the type locality on 15 May 1949, at light, by E. C. Becker.
The holotype is in my collection.
Gnathophthorus ralhis, n. sp.
This species is allied to clematis Wood, but it is distinguished by the different sculpture of the frons, by the minutely etched pronotal disc, by the larger strial punctures, and by the dark color.
Female.— Length 1.8 mm (paratypes 1.5-1.9 mm), 3.7 times as long as wide; color very dark bro^vn, almost black.
Frons flattened from just above epistoma to vertex; area below upper level of eyes rather finely, shallowly punctured, with inconspicuous, moderately long, hairlike pubescence; upper area twice as large as lower area, pilose as in clematis, marginal fringe long, tips of upper setae extend beyond middle of pilose area.
Pronotum as in clematis except posterior
area reticulately, longitudinally etched, punc- tures slightly smaller.
Elytra as in clenmtis except strial punctures slightly larger, surface with minute, impressed points, and shallow, irregular, transverse lines; declivity devoid of granules.
Male.— Similar to female except frons as in male clematis with line between convex and impressed areas more nearly carinate.
Type LocALiTi.— Merida, Venezuela.
Type Materl\l.— The female holotype, male allotype, and 65 paratypes were taken at the type locality on 29 December 1970, 1700 m ele- vation, No. 210, 212, from recently cut stems of Clematis, by S. L. Wood. The habits were as in clematis.
The holotype, allotype, and paratypes are in my collection.
Gnathophthorus pertusus, n. sp.
This distinctive species differs from all others in the genus by the small, median eleva- tions on the male frons at the epistoma and at the upper level of the eyes, and by the pe- culiar cavity on the female frons. The small size and several other features also serve to dis- tinguish it.
Female.— Length 1.4 mm (paratypes 1.1-1.4 mm), 3.7 times as long as wide; color light yellowish brown.
Frons shallowly concave from just below upper level of eyes to vertex, a somewhat indefi- nite summit just below impressed area; median third of summit area occupied by a large cavity from upper level of eyes one-third distance to epistoma, inner diameter of cavity apparently larger than its opening on surface, its margins slightly elevated, its inner walls precipitous; surface of concave area subreticulate, impunc- tate, its upper margin with a fringe of short hair; surface of lower area with a few minute punctures and sparse, short pubescence. Funic- ular segments, except pedicel, very crowded, evidently partly fused to one another; club with suture 1 weakly procurved, 2 almost straight.
Pronotimi 1.4 times as long as wide; widest at base, sides straight and converging very slightly on basal three-fourths, rather narrowly rounded in front; anterior margin armed by about 12 small teeth; summit indefinite, one- third pronotum length from anterior margin; anterior third finely asperate; posterior area almost smooth, punctures not evident.
54
Bricham Young University Science Bulletin
Elytra 2.2 times as long as wide, 1.7 times a.s long as pronotum; sides straight and parallel on slightly more than basal two-thirds, then gradually converging, very broadly rounded be- hind; striae not impressed, punctures minute, almost totally obsolete, visible in subsurface structure; interstriae almost smooth, impunctate. Declivity moderately steep, rather deeply sul- cate; striae as on disc; sutural interstriae weak- ly elevated, 2 narrowly, deeply impressed, .3 abruptly, strongly elevated well above level of 1, its summit anned by about five granules of moderate size. Vestiture largely confined to de- clivity, of sparse, coarse, moderately long hair.
Mali£.— Similar to female except frons convex on upper half, broadly, transversely impressed below, a short, median elevation on epistoma and at upper limit of impressed area, cavity and setae absent; longer setae on elytral declivi- ty flattened, scalelike, small setae in ground cover much more conspicuous.
Type Locality.— Twenty km southwest of El Vigia, Merida, Venezuela.
Type Material.— The female holotype, male allotype, and 33 paratypes were taken at the type locality on 21 November 1969, 50 m ele- vation. No. 150, from a recently cut section of a large Bignoniaceae vine (liana), by S. L. Wood. The woody structure of the liana was layered and each pair of the monogamous beetles constructed transverse, biramous egg galleries at each of a half dozen or more suc- cessive layers; the connecting tunnel was more or less straight and was located where each pair of egg galleries branched. Larval mines were rather short and longitudinal. Fungal growth was not evident.
The holotype, allotype, and paratypes are in my collection.
Sphenoceros azteciis, n. sp.
This is the second known species in this genus and the first from North America. It is very closely related to Immx Schedl, but it is
distinguished by the very minutely punctured central area of the female frons, by the indis- tinctly more slender, angulate, anterior projec- tion of the female pronotum, by the more slen- der elytral bristles, and by the more finely punc- tured elytra with fewer interstrial punctures.
Female.— Length 2.0 mm (paratypes 1.8-2.2 mm), about 2.1 times as long as wide; color very dark reddish brown.
Frons exactly as in Umax except surface of large central area minutely, densely punc- tured. Antenna as in Umax except suture 2 more distinctly indicated. Pronotum as in Umax except anterior projecting angle very slightly more slender, and punctures on disc distinctly smaller.
Elytra as in Umax except punctures of striae and interstriae very slightly smaller, those of interstriae less numerous. Interstrial scales slightly longer and more slender.
Male.— As in male of Umax except lateral punctures on pronotal disc smaller; elytra differ as in females.
Type Locality.— Six km (4 miles) west of Tepic, Nayarit, Mexico.
Type Material.— The female holotype, male allotype, and eight paratypes were taken at the type locality on 13 July 1965, 1000 m elevation. No. 240, from cut branches of a large unidenti- fied tree, by S. L. Wood. One paratype is from El Salto, San Luis Potosi, Mexico, 19 June 1953, Ficus, S. L. Wood. A series from 8 km east of San Bias, Nayarit, Mexico, 12 July 1965, No. 235, is in very poor condition and is not part of the type series.
This genus was placed by Schedl in the Corthylini; however, the above species is mo- nogamous, the adults construct transverse, bira- mose tunnels in the cambium region and deposit their eggs in niches, and the larval tunnels are longitudinal and in the phloem. This genus is very closely related to and eventually may be united with certain species of Neodrtjocoetes.
The holotype, allotype, and paratypes are in my collection.
Brigham Young University Science Bulletin
ECOLOGY OF SCELOPORUS OCCIDENT AUS
LONGIPES BAIRD AND UJA
STANSBURIANA STANSBURIANA
BAIRD AND GIRARD
ON RAINIER MESA, NEVADA TEST SITE
NYE COUNTY, NEVADA
by
Wilmer W. Tanner
and James M. Hopkin
BIOLOGICAL SERIES — VOLUME XV, NUMBER 4 JANUARY 1972
BRIGHAM YOUNG UNIVERSITY SCIENCE BULLETIN BIOLOGICAL SERIES
Editor: Stanley L. Welsh, Department of Botany,
Brigham Young University, Provo, Utah
Members of the Editorial Board:
Vernon J. Tipton, Zoology Ferron L. Anderson, Zoology Joseph R. Murdock, Botany Wilmer W. Tanner, Zoology
Ex officio Members:
A. Lester Allen, Dean, College of Biological and Agricultural Sciences Ernest L. Olson, Chairman, University Publications
The Brigham Young University Science Bulletin, Biological Series, publishes acceptable papers, particularly large manuscripts, on all phases of biology.
Separate numbers and back volumes can be purchased from Publication Sales, Brigham Young University, Provo, Utah. All remittances should be made payable to Brigham Young University.
Orders and materials for library exchange should be directed to the Division of Gifts and Exchange, Brigham Young University Library, Provo, Utah 84601.
Brigham Young University Science Bulletin
ECOLOGY OF SCELOPORUS OCODENTAUS
LONGIPES BAIRD AND UTA
STANSBURIANA STANSBURIANA
BAIRD AND GIRARD
ON RAINIER MESA, NEVADA TEST SITE
NYE COUNTY, NEVADA
by
Wilmer W. Tanner
and James M. Hopkin
BIOLOGICAL SERIES — VOLUME XV, NUMBER 4 JANUARY 1972
Frontispiece. Rainier Mesa Study Plot.
TABLE OF CONTENTS
Ecology and Life Histor)' of a Population of Sceluporus i>. longipet: Baird on Rainier Mesa, Nevada Test Site,
Nye County. Nevada - -- - 1
REVIEW OF LITERATURE 2
Ph\sical En\ ironment 3
Biotic Em ironment 4
Parasites _ 6
METHODS AND PROCEDURES 7
Home Range 8
Bcliavior 13
Heat Requirements and Temperature Relations 14
REPRODUCTION 16
Size of individual eggs and the numbers in a clutch 19
Hatchlings '.: ^0
lusfniles -■-
Adults 25
Population Structure -6
Food Habits 27
DISCUSSION AND CONCLUSIONS 28
LITERATURE CITED 30
Notes on the Life Histor\ of Uta s. stamhtiriana Baird and Girard 31
DISCUSSION AND CONCLUSIONS 37
LITERATURE CITED 39
ECOLOGY OF SCELOPORUS OCCIDENT AUS LONGIPES BAIRD
AND UTA STANSBURIANA STANSBURIANA BAIRD AND GIRARD
ON RAINIER MESA, NEVADA TEST SITE, NYE CO., NEVADA
ABSTRACT
This paper constitutes the first in a series of tenniiKil reports to tlie United States Atomic Energy Commission for Contract AT (11-1) 1496. It includes a major study on the ecology and life history of Sccloponts occidentalis lon- gipcs Hollowell and a comparative study of three populations of Vta s. stansburiana Baird and Girard with special emphasis given for the Uta population on Rainier Mesa.
The study plot ot major concern in this
report is located on Rainier Mesa of the Nevada Test Site. Reports for other shidv plots ( French- man's Flat— four study areas— and Mercury Val- ley ) will follow in the near future and will include infonnation concerning the ecology and life history of at least the following species: Coleonyx variegtxis, Crotaphyfus coUarls, Crota- phijtus wislizeni, Saurornahis obesus, Callisaurus (Imconoidcs, Sccloporus mu<nster, Plirt/nosoma phityrliinos, and Cliiotuictis occipitalis.
THE ECOLOGY AND LIFE HISTORY OF A POPULATION OF
SCELOPORUS OCCIDENTALIS LONGIPES BAIRD ON RAINIER MESA, NEVADA TEST SITE, NYE COUNTY, NEVADA
by
Wilmer W. Tanner' James M. Hopkin-
Only a few life history studies have been concerned with lizards in the Great Basin, and most of these have not included a detailed study of marked lizards on a study plot. Most studies dealing with autecology have suffered from a lack of fjuantitativc data, which is per- haps a direct result of gathering data over a short period of time. It is true that much infor- mation can be accumulated in one year. How- ever, we have found the desert to vary from year to vear, primarilv the edaphic factors which in turn induce and effect changes in the biotic factors. We believe that in order to examine the fluctuations of the biotic cycles and rhvthms in a desert population several years of intensive study are needed if one is to understand the basic aspects of population ecology. Further- more, the roles of individuals, males and fe- males, and age groups are not easily understood
unless at least one generation is followed through a cycle.
There are a few studies of lizard ecology which are complete enough to be cited as ex- amples. A recent one by Tinkle (1967) con- cerning the life and demography of Vta stan-s- buriana stcjiwt^eri in western Texas is based on a large quantity of data gathered over a period of many years. Perhaps the completeness of the Uta study lies not only with the thoroughness of the investigator but also from the fact that utas are numerous and therefore a larger num- ber of individuals increased the mass of data available. In our studv plots in southern Nevada, We have been more successful with dense populations than with sparse ones.
The work of Fitch began in 19.3.5 and has continued to the present. His publications cover a wide area in vertelirate ecology and
M)ej)arliiient of 'Awilogy. Rri^ihiini Y'liin^ University. I'rdvcj. I'Lah MHiDl -Instniclftr nf BioIoev. Inilian .SprinKs HikH .Srhonl. Indiciii .SiirinRs. NcviiJii.
BntciiAM YoiNG University Science Bulletin
include detailed reports on Sceloporus (1940), Euinccei (1954 and 1955), Crotaphytus (1956), and Cneriii(loj)liorits ( 1958 ) and a report on li/.ard roproduction ( 1970). Tlie studies of Blair ( 1960), Carpenter ( 1959-1961), and Zweifel and Low (1966) represent other excellent studies.
The present studv was begun in mid-July 1965, and intensive fieldwork was continued through 1967. Since then field work has been done only for short periods during the period of lizard activity. We have been assisted bv graduate students in the field and laboratory. Others most closely associated with this study plot were Ronald L. Morris, Philip A. Medica, and Dr. John E. Krogh. Each has provided field notes and suggestions which have aided in preparing the manuscript. Dr. Henry S. Fitch, University of Kansas, visited our study areas during the summer of 1967. In September of 1968, Dr. Jeff Swinebroad of the U.S. Atomic Energy Commission also visited our plots. We
appreciate their many suggestions. For all this, we are grateful; however, the analysis of the data anil liiial pri'paration is that of the authors.
This study was supported by Research Grant number AT (11-1) 1496 between the United States Atomic Energy Commission and Brigham Young University, Provo, Utah. We are grateful for this support and for that given by the Civil Effects Testing Operation ( CETO ) base at Mercury, Nevada, under the direction of Mr. Erwin Schultz.
It is not the intent in this studv to attempt a report which would include all segments of the widespread species Sceloporus occidentalis, nor to even suppose that the population found on Rainier Mesa (Fig. 1 and 2) and reported below would properly or completely represent other populations of the subspecies under con- sideration. However, a general review of the literature is included as a basis for a general understanding of the species.
REVIEW OF THE LITERATURE
The Western Fence Lizard, Sceloporus occi- (lentalvi Baird and Girard, occurs over a wide range in western North America. It is one of the most commonly seen lizards in areas where it occurs because of its diumal habits which in- clude basking fully exposed on rocks tree trunks, poles of fences, and on other objects exposed to solar radiation. The range of the species extends from the state of Washington to Baja California and from the California coast through Nevada into the Great Basin area of Idaho and Utah. We agree with Stebbins (1966) that this species does not (X'cur in the low desert valleys but is found in habitats where there is adecjuate cover consisting of at least sagebrush and where basking areas are avail- able. The ability of this species to absorb sun- light and warm (juickly enables it to inhabit areas from sea level near the California coast to altitudes of oyer 9,000 feet in the mountains of California, Nevada, and Utah.
Taxonomic studies of Sceloporus occidentalis have been published bv Camp (1916), Smith (1939), and Bell (19S4). Bell recognizes six subspecies. It has not been possible to examine populations in other altitiulinal ranges, and it is recognized that populations may vary morpho- logically and ecologically because of differences in altitude, the accoinpan\ing differences in the physical and biotic cm ironments, and from the isolation impo.sed bv these factors in desert
areas. Because of the morphological variations and the fact that there are several recognized subspecies, we suspect that there may also be within the limits of this species variations in behavior, reproductive activities, size, growth rate, and other life history and ecological pat- terns. The Great Basin subspecies, Sceloporus occidentalis longipes Baird, occupies the larg- est range of the occidentalis subspecies and, ex- cept for S. o. taijlori which is reported to be of ecjual size, attains a larger individual size than other subspecies.
The subspecies Sceloporus occidentalis occi- dentalis has been studied rather extensively, probably because it occurs convenient to Berke- ley, California, and nearby research areas. Fitch (1940) studied S. o. occidentalis in the field near Medford, Oregon, and near Berkeley, Cali- fornia, with special emphasis on growth and behavior, and Davis ( 1967 ) made a detailed studv of growth and size at the Hastings Res- ervation, Monterey C'ounty, (California. Thermal relations of S. occidentalis have been studied by Dawson and Bartholomew (1956), Wilhoft (1958), Wilhoft and Anderson (1960), Larson ( 1%1 ), and McCinnis ( 1965). Talbot and Live- zey (1964) and Coleman and Livezey (1968) studied the closely related area of integumental reflectivity. Testicular histology and seasonal changes of the testes of S. occidentalis have been' studied by Wilhoft and ^uay (1961).
Biological Sehies, Vol. 15, No. 4 Sceloi'ohus Occidentalis and Uta Stansburiana
^^^^*L}
Fig. 1. A \ie\v of Rainier Mesa from the southwest showing the granitic cap rock exposed on the sonlhwesfem edge. Photo taken about 11:00 a.m. bv R. L. Colyer, AEC photographer.
Johnson (1965) .studied the diet of S. occklen- tahs. Other studies of anatomy and physiology have made use of this species (Ells 1954, and Stebbins and Eakin. 1958).
Sceloporus occidentalis occurs in many kinds of plant associations. The study areas of Fitch (1940) were described as "A stretch of rail fence . . . which passes through an overgrazed pasture . . .; a stretch of rail fence . . . through open woods of Garry oak . . .; and a board fence and a wooden bridge adjacent to pasture land, eucalyptus trees, Monterey cypress, and native chaparral." In the Great Basin, S. occi- dentalis tV'pically occurs in the pinyon-juniper plant fonnation (Fig. 1 and 2) but also occurs in areas of oak brush (Quercus <iavd)eli) and also in nearly pure stands of Artemisia tridentata if basking areas are available.
Physical En\'ironment
The high altitude of the Rainier Mesa study plot, 2,286 meters ca ( 7,500 feet ) above sea level, gives it a climate very different from the
surrounding low valleys (3,500-4,000 feet). The temperatures are much lower and the rainfall considerably higher, as indicated in Figures 3 and 4 (Swan, 19.54).
The cap rock of Rainier Mesa is lava ( Fig. 2), and the cap is covered by a shallow mantle of soil formed from the lava with the addition of organic matter from decaying plant materials. At the base of pinyons and junipers, there is a deep layer of decayed and decaying needles. There are many rock outcroppings throughout the study plot, and these, along with lava rock fragments and tree trunks, furnish basking areas and shelter for the lizards.
Glimatic conditions prevailing at the Nevada Test Site were summarized by Allred et al. (1963) for the years 1959-1961. Their reference to two weather stations at 7,480 feet gives a comparable elevation to the station near the Rainier Mesa study plot (7,490). They report 8.75 inches for 1960 and 6.74 for 1961. Weather Bureau reports for the Rainier Mesa station re- port precipitation for 1965, 1966, and 1967 to be 12.67, 7.34, and 11.13 inches, respectively
Bricham Young Universiti' Science Bulletin
. ■■ A
■<¥.
"^K,
i^'f '
Kij^. 2. A view looking north from tlie nortliwfst cornrr ol the stniK plot. The L'.S. weather tower is seen in the haeki'rounil. Photo hv R. L. Colyer.
(JuuL' tliroiigli Dcft'inhcr). This is considerably more than reported by Allred and indicates the great variability of the climate in this area (Fig. 3). Allred et al. ( U)63) reported that 136 percent of the precipitation occurred during the summer months (April through September). Available data for 1965-1967 averages 41.3 per- cent. There may be heavy precipitation occa- sionally during a given month; however, the most consistently moist period .seems to be July and August (Fig. 4). An average of the precipi- tation lor these months for the years 196.5-1967 is 1.4 inches per month. The greatest amoimt recorded for one month is 4.67 inches in Novem- ber 1985. October and June have been the driest months, with an average of onlv 0.29 inches each month.
Temperatures are moderate, rarely exceeding (85° F in summer and reaching 0° upon occasion during the winter. On 1 June U)67, temperatures were below free/.ing and a skiff of snow fell. A freezing storm also occurred on 16 September 1965. Rainier Mesa is therefore subject to the cold fronts which descend across the desert
from the northwest and also to the influences in the summer of the hot desert valleys which nearlv surround it.
BiOTic Envibonment
The following study covering nearlv five years of field work should add to our under- standing of the life history, food habits, home range, behavior, and general ecology of this common western lizard as it occurs in a pinyon- juniper commimity on a high mesa in southern Nevada.
The plant eanopv cover on the studv plot at Rainier Mesa (Nevada Test Site) is approxi- mately 60 percent, with Pintts inottojiliijlhi the dominant plant. Other woodv plants such as Artemisia tmlcntata, Quercus gambelii, C.titier- rczki inirroccjiluihi, Junijicnis astcospcrimi, Pur- sliia tridcntcit, Ejilwdra virulis. Rihcs vclitti'iiurn, Tclradiimid ciincsccns:. lA'])tod(irliihm punncus, and Linanllioshniin nuttallii make up the great- er part of the woody plant cover (Fig. 1). The percentage of annuals is not included because
HioLOGicAL Series, Vol. 15, No. 4 Scelopohus Occidentalis and Uta Stan.shuriana
1961
1962
1963
1964 1965 1966
1968 1969 1970
lit^ 3. Precipitation for the ten-year period 1961 to 1970. The dark cohirpns for Hock Valley. Data provided bv the U.S. Department of Commerce, courte.sy of Mr. Ralph (Quiring, Climalologi.st.
it Viirie.s con.siderahlv each vear depending on the moisture available.
Allred, Beck, and Jorgensen (1963) consider- ed both the plant and animal species of the pin\on-juniper community and comments here will deal onlv with the more common plant and animal species that seemingly are more directly associated with SceJojwrus occidentalis lon- gipes. For a complete list of plants and animals see Allred et al. (1963) and Beatley (196.5 and 1969). Only two other species of lizards, Via s. stan.shuriana and Eiimeces skiltoniamis utalien- sis, were found on the study plot. Both of these species are considerahlv smaller in size than S. o. longipes and therefore they may not exert much competitive pressure on S. occidentalis. On the lower foothills of Rainier Mesa, Scclo- porus magister and S. occidentalis overlap their ranges, and in this area of overlap the competi- tion for food, shelter, and basking sites is un- doubtedly more intense. No effort has been made to studv the area of overlap, but it has been observed in driving through the area that a basking site on a boulder was occupied by a member of one species one dav and by the other species only two davs later. It is also possible that Crotaphijtns coUaris and Crota)>hij- tus wi.slizeni occur on the fringes of the range of this population of S. occidentalis, but no specific observations have been recorded. If these species do overlap, it is almost certain that the two species of Crotaplujtus prey at least on the hatchling and juvenile of S. occidentalis.
Three species of snakes have been collected on or near the studv plot. One specimen of Masticophis taeniatus was taken on the plot in 1966. Another was sighted .50 to 7.5 feet north- west of the plot in 1967. \ \er\' large Pituophis
mclanolcncns dcscrticola was captured within 1.50 teet of the northeast corner of the plot in 1967. Three specimens of Crotaltis niitchelli stcphcnsi have been sighted on or near the study plot. One large one was captured, marked by scale clipping, and released in Ui65. He has not been recaptured. A large individual was seen on the west edge of the plot in 1966 and a female was taken on the plot in 1967.
A survey of the abimdance of arthropod species in the area goes beyond the scope of this study, but it is obvious to those who have attempted to rest in the shade of a pinyon on Rainier Mesa that ants are abundant both as to number of species and individuals. There are 28 species of ants known to occur on Rainier Mesa (Cole 1966). Other arthropods known to be common by their freijuent appearance in can-traps are beetles (mostly Tencbrionidae), Jerusalem crickets (Stcnoj)ehnatus ftisciis), scor- pions, and solpugids. A list of animals collected from within the pinyon-juniper plant communi- ty at tlie Nevada Test Site is indicated by All- red, Beck, and Jorgensen in their report "Biotic Communities of the Nevada Test Site" (1963). Tanner and Jorgensen ( 1963) listed the reptile species known to occur on the test site.
Of the possible predators of Sceloporus oc- cidentalis longipes on Rainier Mesa only one species Masticophis taeniatus, is known to feed primarily on lizards. A large A/, taeniatus in cap- tivity has been observed to eat four adult S. occidentalis in a period of less than four hours.
The gopher snake, Pittiophis melanoleuciis, and the Panamint rattlesnake, Crofalus niitchelli stepJiensi, are potential predators, but both species are believed to prefer mammals; and, because of the few individuals observed, it is
Bnicu AM VoiNo University Science Bulletin
I'ii;. 1. Prtcipitation for tlu- year 1965 plotted for each month. Dark columns represent Rainier Mesa and linctl columns are for Kock Valley. The symbol = iindicates onlv a trace of precipitation. Solid and broken liiiis plot the a\erage mavimum temperatures for Rainier Mesa for 1965 and 1966.
iiiiliktiv that tlit'V exert appreciable pre.s.siire on tile lizard population.
No evidence has been found to indicate that mammals prey on the lizards. However, several species of mammals known to occur in the area (Hayward 1965) have been reported to feed occasionally on lizards in other areas. Coyotes are known to eat vvcsteni fence lizards (Leach and Frazicr 1953; and Ferrel, Leach, and Tillotson 1953). Badgers are also known to eat lizards, although the records are for species other than S. occidentaJis (op. cit. ). Foxes, l)ob- cats, skunks, weasels, and possiblv even cougars may occasionally prey upon lizards. A shrew killed and ate one of the lizards while they were together in a can-trap. Perhaps shrews kill lizards in nature especiallv if the shrew sliould lind the lizard in its hiding place when it is cold and toipid. A shrew was taken alive to the (>ivil Effects Test Operations laboratorv in Mercury, Nevada, and kept alive for about a month. It showed the ravenous appetite for which shrews are famous, but would not kill and eat lizards as long as crickets or other
insects were axailable for food. We therefore doubt that shrews prev on lizards in southern Nevada.
Jerusalem crickets, Stcnojx'hnatu.s- fusctts, killed and ate several lizards in can-traps, and others were badlv injured bv this predacious insect. In Stenoj)eImatus\ we have a possible predator of S. occidentaUa in nature, but we have no evidence to show that this insect would kill and eat lizards under circumstances other than the unnatural confineinent of the two ani- mals in can-traps. Ants and scorpions also killed several small lizards w hile together in can-traps.
There are several species of birds that may prey on lizards. Among them the shrike, hawks, and falcons are known to be present on Rainier Mesa. The roadrimner has been seen in the pinyon-juniper communit%- at lower elevations; however, we have not seen one on the top of Rainier Mesa.
Fah.\sites
()l 145 lizards' stomachs examined, 41 con- taiiu'd round-worm parasites. Many contained
Biological Series, Vol. 15, No. 4 Sceloi'ohus Occhjentalis and Uta Stansuhkiana
only 1 womi, whereas the maxiimiin was 30. The average for those parasitized was 4.09. Only stomachs were examined for wonn para- sites. The intestines undoubtcdh' contained ad- ditional parasites, but have not been examined. Intestines are still available for examination.
Mites are common on many species of liz- ards. Only one species of mite is known to infest
S. 0. Jongipes on Rainier Mesa. Geckohidla tcxana Banks has been taken from lizards during the months of May, June, and July.
Small ticks also have been taken from S. o. longipes. During a brief period in August (15-20) of 1966, we noted that about half of all lizards captured had one or more ticks, usually in the region of the neck and shoulders.
METHODS AND PROCEDURES
This stiidv was begun in July of 1965 and continued from 1 June to 1 September each summer through 196S. Limited data were col- lected in September of 1965, 1968, and 1969. Some sampling was done in October of 1967, 1969, and 1970 and also in April and May of 1967, 1968, 1969, 1970, and 1971. The lizards are in hibernation from late October until early April.
The study plot used for capture and release methods is on Rainier Mesa at an elevation of 2,287 meters (7,480 feet). The plot is 137.25 by 183 meters (450 x 600 feet) and has an area of 2.51 hectares (6.2 acres). It is in a pinyon- juniper community. The plot was staked out in a grid with 15.25 meters (50 feet) between stakes. The stakes sometimes had to be moved slightly because of rocks and trees but were always placed as near as possible to their proper location. When the plot was established it had 10 rows of stakes in each direction, but three rows of 10 were added to the south end of the plot in June 1966. A can-trap was placed at the base of each stake as near to the stake as pos- sible and on the south side. Here again it was sometimes necessary to compromise, but the can was placed as near its proper location as physical circumstances would pemiit.
There is a weather station about 100 meters north of the study plot and weather data ( Fig. 3 and 4) are available for use in this study.
The can-traps mentioned above are metal cylindricLil cans about 15 centimeters in diam- eter and 30 centimeters deep. The cans were set in the ground so that the top of each can is level with the soil surface. The opening of the can is then covered with a small sheet of masonitc with two-inch wooden blocks so fas- tened to it as to form tripod legs which keep the cover two inches above the ground surface. The covers prevent direct sunlight from reach- ing the animals and also serve to keep out dir- ectly fallinsi water from rainstorms which are
much more frcrjuent on Rainier Mesa than in the surrounding desert flats. The covers prob- ably attract lizards by offering shelter and thus increase the chances of their capture. The cov- ers may have an effect also in keeping out larger predaceous animals. A lizard is relatively safe in this type of can-trap for up to 56 hours. The cans were checked at least every 48 hours except on two occasions when AEC testing activities prevented us from entering the area. We did lose a few animals in the can-traps as a result of water flowing into the traps during heavy rains and by other animals killing the lizards.
In addition to can-trapping, lizards sighted on or near the edge of the study plot were captured whenever possible. Some were cap- tured by hand, but most were caught with a small noose of braided nylon tied to the end of a fishing pole. This procedure will be called noosing throughout this report.
Captured lizards were marked by toe clip- ping following the system used by Tanner (1965). Each lizard was also marked by paint- ing a number on its back with a water-base poster paint. This made recognition in the field possible and reduced the necessity for repeated captures since lizards only needed to be cap- tured occasionally to check for growth and u'eiglit change. The paint was especially helpful in sighting lizards clinging to tree trunks where they are usually difficult to see.
Records were kept for each lizard including place of capture, toe-clip number, snout-to-vent length (s-v), total length, tail regeneration, weight, sex, and approximate age. From hatch- ing until the first hibernation, a lizard is called an hatchling; from emergence in the spring to the second hiberation, the lizard is called a juvenile; and on emergence from the second hibernation the lizard is considered an adult. When the age of a lizard was in ([uestion, a s-v length of 70 mm was arbitrarily considered to be the dividing line between juveniles and
8
HiuciiAM YoiNO University Science Bulletin
adults, although marked juveniles have been knov^Ti to exeeed that length in late August, and a few adult lizards are known to measure less than 70 mm s-v at their emergenee Irom the second hiberation in April and May.
If a lizard was captured by noosing or some other means that would not greatly change body temperature, its cloacal temperature was taken with a Schultheis (juick-reading thermom- eter. If the lizard was c-aptured in a can-trap where it was prevented from regulating its temperature by its usual behavioral means, the temperature was taken only when it would be useful for comparison with lizards free to regu- late.
Special notes were made of sears or other abnonnal marks, rate of tail regeneration, repro- ductive condition (i.e., gravid or not gravid during the I'gg-laying season), and behavioral observations.
A series of more than 200 lizards was col- lected at season intervals outside the study plot for histological examination of the gonads. Since most of the mating activity occurs during April and May when it is not possible to spend more than perhaps a weekend at the study area, it is especially important to have other means of determining the peak period of mating. Some of the lizards were captured and then frozen, others were taken to Provo alive for fixing and sectioning of tlie gonads. The latter method was very satisfactory, whereas the testes that were frozen were badly disrupted and did not show cellular detail to the desired degree. The testes wiTc fi.xed in Houins fluid, imbedded, sectioned at 7 microns, and stained by Delafield's hemo- toxylin and counter-stained with eosin. All go- nads were weighed before fixing, and the fat- bodies associated with the gonads of both sexes were weighed and discarded.
The stomachs of some of the lizards collected for the reproductive sample were examined as to volume and kinds of food they contained. Stomach volmue was measured by placing the stomach content in a 10 milliliter graduated cylinder and then adding alcoliol to the cylinder from a burette until the stomach content was immersed or at least floating. Both the burette and graduated cylinder were then read to de- termine^ the volume of the stomach contents.
tor a li/ard of its size. The home range that is believed to be a maximum estimate for 9.5 per- cent of the adult male lizards oi this population is 1.67 acres (Fig. 5). The home range maxi- mum estimated for females bv the same method
Fig. 5. A polvgoii reprcsfiitiiig tlir actual home range of a malt" Sccloporiis as determined hv the eapture- ree.ipture niethcid for lizard inimber 2-6. There are 32 eaptnre^ plotted dnring three summer seasons, 196.5-19()7.
is ().4S acres (Fig. 6). The average home range area as estimated by the minimum polygon method is 0..59 acres for adult males and 0.09.5 for adult females. As explained above, it is be- lieved that the actual home range area is be- tween tliese two estimates and probablv varies greatly with the individual lizard and the physi- cal and biotic makeup of his home range.
Home range estimates have been made by two methods. Thev will be referred to as (1) the minimum polvgon method and (2) the re-
Home Hange
Home range means the area utilized by a lizard in its daily pursuits during a year and in some individuals a lifetime. Srrloporus o. longipes utilizes a relative small home range
15.25 meters
(soft.)
I'ig. (1 A poKgon ^llo\^ing the outer hmits of a female hzard liome range, number 1-14, as dctcnnined by 20 e.iptures.
Biological Series, Vol. 15, No. 4 Sceloi'ohus Occmdentalis and Uta .Stan.smuhiana
capture radiu.s method. The ininiiiium polygon method eon.si.sts of plotting capture .sites on a graph and connecting the outennost sites with lines to form a polygon ot three or more straight sides. Only lizards with four or more captures not in a straight line were used in estimating home range. The polygon figure is then divided into rectansiles and triansiles so that the area of the polygon can be calculated.
For the recapture radius method the geo- metric center of all the points of capture and oh,ser\ation ( considered hereafter as a recap- ture) is located with the capture points dis- tributed around the point representing the center of acti\ity. A figure for each lizard is produced, as in figures 5 and 6. If many in- dividual lizard capture-recapture figures are superimposed with the center of activity for each set above a single point on a graph ( Fig. 7 and S), a figure of recapture radii for a species or population is theoretically the result. A circle can now be drawn to include 95 percent of the capture recapture points. The radius of this circle is then the observed home range radius thcoreticalK' possible for an individual at the 95 percent level. This radius can now be used to calculate the area of a hypothetical circular home range.
It is almost certain that circular home ranges are rare or do not e.xist in nature, but it is a
15.25 meter (5 0 ft.)
Ki}^. S
The ohst-rved rt-capture radii for all females the populatiiiii, with the 9.5 percent level in- dicated by the circle. There are 26 lizards captured and recaptured 270 times. Each lizard has a minimum of at least three recaptures.
Fig. 7. The observed recapture ratio for males in the popul.ition, with the 9.5 percent ]ev<'I indicated bv the circle. There are 244 captures of 19 lizards each with a minimum of 3 recaptures included in the fii^nre.
convenient wav to estimate the approximate size of a home range from a few captiires. The minimum polygon method may give an indica- tion of the actual shape of the home range if there are a large number of recaptures. The area estimated by the minimum polygon method is probably less than the actual size of the home range area. It is likely that the actual home range, if it could be accurately determined, would have an area between that estimated by the recapture radius method as a maximum and that estimated by the minimum polygon method as a minimum. Furthennore, our data indicates that some lizards have larger or smaller home ranges than do others. This is perhaps an indication of the variations occurring in the terrain of the habitat, availability of food, or other factors such as individual variations in behavior. Turner, Jennrich, and Weintraub (1969) also concluded that the home range is intennediate between the minimum polygon as indicated by Tinkle, McGregor, and Dana (1962) and the recapture radius inethcxl as in- dicated by Jorgensen and Tanner (1963). F'ur- thermore, there is variation in the size and shape of home ranges. This variance is seemingly the
10
Briguam Young University Science Bulletin
result of the uih-vcii terrain and the age of in- dividuals (size being a factor), and we believe that there is a variation in the aggressive be- havior of individuals, particularly in males, which increases the area under sur\'eillance.
There are records of a distinct change of home range between two summer seasons. Liz- ard :;:;:3-16, an atliilt female, occupied a home range of about 2.5 feet in radius during the 1966 and 1987 seasons but was captured 150-200 feet east of that range on 3 June 196S. Another adult female, it 19, was captured six times in
1965 in an area about 200 feet into the plot from the east side and 150 feet in from the north. She was not captured or observed during the
1966 season. In August 1967 she was captured approximately 1.30 feet east of A-5 on the edge of the plot. She was recaptured three days
later in the same general area. Apparently, she had moved her center of activity approximately .55 feet between 1965 and her recaptures in 1967. There are no records of a complete change of home range in adult males; however, the males have a much larger home range, and the polvgon figure drawn for anv one of several of the males would i-ncompass the enlarged home range areas of both ot the females that changed the center of activity' in their home range or a new home range if two were estab- lished.
There is considerable o\'erlap of home ranges, as indicated by polygon figures drawn from capture and recapture data over a three-year period from July UJ65 through IfJ67 (Fig. 9). Based on the considerable overlapping of the polygons for males, one is (juite uncertain as
^>..
■>-^ + +
1 /
V
Tig. 9 \ scries (it lioinc r.iiii;c polN^oiis tor mairs (luriiii; llic prricKJ trcim 1965 tlirDiigh 1967. Tlic Jiffcrent t\|)cs of iiiu's .ire tor lomcniciuc ot lollowiiii; the o\ rrl.ippiiit; liiu's.
Biological Series, Vol. 15. No. 4 Scelopohu.s Occident.-m.is .\ni) Ut.\ St.\n.siilihi.\na
11
to the territorial relation.ships between males. A few tentative concliision.s .seem justifiable. Males do li\e in close proximity to other males but do not share tlie same basking areas and are not found together during the spring and summer months. They occupy relatively large areas, which apparentlv thev cannot constantly protect; thus, other males frequentiv cross what appear to be boundary lines and are recorded as being in anotlier home range or having a large area of their own which is entered by other nearbv males. In a rough terrain where vegetation is plentiful, home ranges may not be as protectable as in desert flats. In spite of the apparent overlapping of home ranges, we note that males do remain in a restricted area for several years. Data does not invalidate the home range, but it does indicate a reduced tendency for males to set up and virgorously protect a large territorv to the exclusion of other males. On the basis of the polygons ( Fig. 9 and 10) and our field observations, we believe that there is a sort of peck order among males
in which the more dominant males essentially rule within their territory, with younger and less dominant males being submissive. Thus the dominant males move freely within a large territory but protect with vigor a smaller area which we refer to as the center of activities and wliich includes the principal basking and hiding areas within a given home range.
We have not observed any social order among the females, although it may occur. Fe- male lizards are often seen basking together, and a male and a female lizard together on a rock is not an uncommon sight.
The large overlapping home ranges which are not totally defended may account for the few encounters betwi'cn males observed by us. The home ranges plotted for 16 males, each of which was observed for three years (Fig. 9), shows considerable variation in size and indi- cates that some males enter three or four other home ranges. One wonders if there is any territorialism, and yet we note some evidence of it in that males are not found together on
Fij^. 10. Hdiiic r^mgt's for male Sceloporus occidentalism H.iiiiicr Mesa, l(ir llic siimmiT of lWj7. Twelve home ranges are plotted. 'I'lic different oiitline.s are u.sed only for eoriM-nienee of lollow ini; the boundaries.
12
BniciiAM Y'ouNC Univkhsitv Science Bulletin
the .same rock, log, or in the same tree. Wc have ol).st'rvc'd a fi-vv hrict encounters; however, these encounters have not indicated that males go much beyond tlieir center of activity, or basking area, in pursuit ol anollier uiale. This apparently leaves large areas of the home range free for other males to move, and our polygons indicate a considerable freedom for movement.
At first we surmised that males changed their home range each year. We therefore plot- ted 12 males' home ranges for 1967 (Fig. 10) and cami' up with about the same size and variety of polvgons as tor the three-year plot- tings. We have concluded that little change occurs except that older males may have large areas. Females, as previously indicated, also have overlapping home ranges. Their home ranges are much siuallir ( Fig. 1 1 ) and give the distinct impression tliat they may be cluster- ed, althouirh this mav be a coincidence.
Males bask side by side, at least in the early spring, as they emerge from hibeniation but are not seen together after the mating season
begins. Oniv a few instances of a lizard's de- fending its territory or challenging the approach of another lizard have been recorded under natural conditions. As an example, on 5 June 1967 an adult male (—3-15) was sighted bask- ing on the can-trap cover at stake H-12. He was noosed, weighed, measured, his location of recapture noted, and was released on a large boulder a few vards awav from the stake. As is tlie custom in this spi-cies and almost all lizards, he soon disappeared on the opposite side of the rock, where he was challenged and soon return- ed across the rock, passed the observer, and rushed into a nearby r<K-kpile. The pursuing lizard, rr6-7, stopped at the edge of the boulder where he displaved bv flattening himself later- ally ( Noble and Bradley, 1955 ) and bobbed sev- eral times. He showed no fear and was noosed. Occasionally as the study plot is being worked, males are temporarily displaced into an adjoin- ing home range. This same type of response mav then occur.
The fact tiiat ft'w instances of territorial
I N \
I'lij. II. Ildiiu' r.iiii;('s lor tcmalcs Sccloponis occidcfi talis, li.iiiiicr Mesa, for the Mimmcr of 1967. Seventeen
Moinc rariiics
.ire plotlecl. The different outlines
are iiset! onu
ominienee of folliming tlie Ixiundarics.
Biological Series, \'ol. 15, No. 4 ScELor()HU,s 0(;c:ii)entali.s and Uta Stansuhriana
13
defensive behavior have been ob.served, in spite of the nuinerous man-hours spent in tlie study plot, suggests that the territories (of males) are established and that trespassing by other males may oceur, but if observed they may be chal- lenged bv a more dominant male. Our data was gathered primarily during June, July, and Aug- ust, with only limited observaions during April and Mav during the breeding season when ter- ritories mav be more meaningful. However, our limited data for the early spring does not appear to be different from that recorded for June, al- though intensive field work for an extended period of time has not been done.
Attempts have been made to determine how the home range is utilized by a lizard. It ap- pears that Scelo}>orus occidentalis Ionaij)es is a rather sedentary lizard, spending most of its time basking in sunlight or loafing in the shade according to its temperature rojuirements at the time. If a suitable prey is seen, the lizard moves (juickly to capture it. If no prey appears, the animal may mo\e only a few feet in several hours. No active foraging or scratching in the soil for prey has been observed. Lizards have been observed to run rapidly for 10 feet or more, snatch an insect or spider, and then run rapidly back to a basking place. A lizard suddenly disturbed and caused to flee in a direction that will lead away from its home range soon be- comes confused. Its apparent unfamiliarity with new terrain induces an immediate return to the home range and the familiar area where hiding places are available.
A most interesting phenomenon is the great consistency of nearly all individuals to stay in a given area for three or more years without any indication that they have changed in the least their center of activity. Those few who have made a noticeable change seem to have made the move to a new center of activity early in the spring and certainly before the mating period begins. This consistency of homing is also remarkable inasmuch as some home ranges are made available for readjustments by the death of old lizards. Presumably, these areas are taken up or expanded into by the rising generations of hatchlings and juveniles, but we have no evidence that established adults ap- precial)Iy expand their home ranges.
Behavior
The home range of an animal is generally defined as the area utilized by an animal in its search for food, mates, and shelter (Tinkle 1967). Capture and observation localities have
been recorded for many lizards during the three seasons of intensive study, but accurate mapping of all of the boundaries of each home range has not seemed practical. Figure 9 does give the home ranges of males. A good estimate of the general areas in which a given lizard may be found can be made, usually with reasonable accuracy. The common practice of showing capture locations and connecting the outermost sites with lines gives some idea of home range, but this by no means shows its precise bound- aries or the extent to which the various portions of the home range are used (e.g., basking, loafing, foraging, ect. ). The latter information is very difficult to obtain because the animals are so sluggish and may remain on a single rock three feet across for several hours. Most at- tempts at observing daily activities have been frustrating. Watching a single animal for the recjuired amount of time to describe a daily schedule is tedius. Attempts to keep two lizards under observation at one time have not been successful. One can watch a lizard for two to three hours and then take his eyes away for ten seconds only to find the lizard gone when he again turns his eyes to the lizard's location. We have noted upon many occasions that a basking lizard does not take flight until a mov- ing object approaches. Rapid movement is more apt to frighten than a slow-moving object. Therefore, a lizard may respond to a movement of the head that had been motionless since its appearance. When a lizard moves down from a basking place, one loses sight of it between rocks or in vegetation and often cannot spot it again without approaching so closely as to frighten the lizard into taking cover. Brief notes have been made of observations on individ- ual behavior. Examples given below are taken directly from our field book (August 1967).
.Sc<7()/)ori(.v occidentalis #5-15 (a large fe- rn lU- ) sighted ba.sking on a .small rock nnrth- iMst of H-12 at approximately 9:50 a.m. (1 then continued eheeking ean-traps). At 1:05 p.m. she was at the southwest eonier of a nearby large boulder.
I walked too elose to her at 1:40, and she moved to the northeast faee of the bonlder. She had mosed less than six ft'et by 1:.50 (i.e., from her 1:05 location).
At 2:24 she mo\ed south 1-2 feet into the sunlight for two to three minutes and then ran across tlie top of the boulder toward the west. She stopped on the southwest comer of the boulder in a shaded spot. 2:32 p.m. Moved about one foot — still in the
shade. 2:36 p.m. Moyed about six inches into sun 2:39 p.m. Moved north along west side out of sight.
14
Hkicmam VouNf: University Science Bulletin
2:41 p.m. .Stopped dm the northeast comer
of the boulder in the shade. 2:5.5 p.m. Moved into sun. 2:56 p.m. Moveil around to the nort]iea.st
face of houldcr whieli is in the shade. .3:15 p.m. Moved onto the tree nearest the big boulder on the east. This li/ard is extremely wary, and although 1 was about 40 feet away she still may have been disturbed bv my presence. She was never observed to e;it or show interest in fcxxl during these oliserv.itioiis.
There was ample tiiiii- for her to warm up and feed prior to the first sis^hting at 9:.50 a.m. Slie may also have fed between 9:50 a.m. and 1 :()"3 p.m. The following is another note cover- ing tliree hours of observation of i±5-13 on 5 August 1967.
Two days later another behavioral note was made of the same lizard:
19 July I()()7 — arrived at the plot at 7:00 p.m. in an attempt to obstTXe lizards preparing to spend the night.
S. occidctitdlis #5-15 sighted in tri'e .30 feet northwest of H-12 at 7:15 p.m. S:0() p.m. Tlie sun is going down and #5-15
is still li.ingiiig, head down, on the
trunk. S:2(i p.m. She turned .iround so her head
is up. 8:35 p.m. She moved (I heard her. It is too
dark to see her). 1 shinetl a liglit on the
tree to find that she had moved up
about one foot. 9:00 p.m. It is much too dark to follow her
if she moves. The flashligfit shows her
to still be clinging to tlie free trunk.
Apparently she is going to spend the
night in the tree.
On 20 Jnlv a female lizard ii3-17 was ob- served to take slielter under a roek and re- mained there as darkness fell. She was well under the rock when it was necessary to use a light to .see her.
On 7 August a lizard it5-20 wa.s observed under tiie edge of a flat rock at S:07 p.m. No attempis were made at ()l)taiiiiiig c-lose observa- tion or observation by a flashlight in hopes that she would remain as .she was and might be seen emerging the following morning. At 6: 2.5 a.m., 8 August, the sun was not up and the tempera- ture was 58° P^ at the nearbv weather station. The sun appeared at 6:.'i0 a.m. At that time the soil temperature was 14.4°C. By 8:45 a.m. the soil temperature was 20.2°. At 8:50 a.m. the head and toreleg of a lizard were seen emerging from imder the rock where ±:5-20 was seen the previous evening. It was not until 9:20 a.m. that the lizard emerged eompletelv and was seen to be zr5-20. At 9:55 a.m. she moved up to the
top of the rock and basked, even though direct simlight was interrupted somewhat by cloud cover. She was observed again ;it 10:32 a.m., 12:18 p.m.. and 1:15 p.m. still b;isking but hav- ing moved about two to four feet at each ob- servation.
The behavior of adult lizards toward each other if thev chance to meet in the course of normal activities is of interest. Soon after emerg- ing from hibeniation males set up a home range or center of activity and become antago- nists. Thus, ;ui encounter between males results in a bobbing contest with the blue belly patch exposed by each. This display of bobbing and flattening of the bodv to expose the blue may i:ist for only a few moments or may be carried on for several minutes. We did not observe a phvsical combat. In all cases, one of the antago- nists fled. In contrast, encounters of females and of females with males did not produce any outward excitement e.xcept during the mating season when males would go into their bobbing of head and body push-ups. Only rarely did we observe such activities on the part of females.
The aggressive displays of males toward each other is not the same as in the case of the supposed courtship displays of the male toward the female. In the fonner, a greater flattening of the body, an arching of the back, and an exposure of the blue on the belly is most apparent. .Although males through their bobbing before a fcm;de do expose the blue, it does not appear to be done with the same intensity. The aggressivi- behavior of males as well as the sex recognition response seems to be tied to a sight recognition of the bodv color and/or color patterns.
Heat Recjuirements and Temperature Relations
The ability of ScelojHinis occidentalis long- ipes to absorb sunlight and warm the body to its active (optimal) temperature while the air remains cool is noticeable. The range of body temperature for all active lizards measured was 21.6° to .37.0-C. The average for adult males is .3.3.0 and adult females .321) (Table 1). These temperatures were recorded for lizards that had been free to regulate their temperature and
Table I. Range of temperatures recorded for :utive lizards (Sceloporus) on the Rainier study plot.
Minimum M;L\imum Average
Male .\dults 21.2 36.2 32.9
Female .\dults 21.6 37.0 32.9
Male Juveniles 19.2 36.9 .34.0
Female Juveniles 22.2 36.2 34.0
Biological Series, Vol. 15. No. 4 Sceloi-ohus ()c:c:ii)Entali.s and Uta Stan.suuriana
15
were captured hv means that would not ijreatlx' alter their body temperatures. The time at which temperature was taken was recorded, and usu- ally the temperature of the air and substrate were recorded tor the same time (as nearly as one thermometer would pennit). Occasion- ally, body temperature was taken on a lizard in a can-trap for comparison purposes. Usually a lizard found in a can had been there over- night. .■Mthougii the cans are only a foot deep the temperature at the bottom of the can is usually much cooler than the air or the soil sur- face. It is not unusual for the temperature of a lizard in a can-trap to be 10-20 degrees below that of a lizard free to bask and absorb solar energy. On 7 June 1967 two lizards from can- traps showed temperatures of 17 and 17.4° at appro.ximatelv 10:.30 a.m. \ lizard noosed nearbv at 10:1.5 a.m. had a body temperature of 32.4°. During the three summer months we found that most lizards emerging from their oyernight hiding places ha\e body temperatures yarving between 15 and 20 \ and this accounts for the low temperatures recorded from lizards noosed soon after thev appear on rocks.
Temperature obseryations made on lizards found in cans and then noosed between one-half and one hour later indicate that at this study plot body temperatures can be increased or lowered 10 to 15 degrees in a short period of time. As an example: A juvenile female (62 mm) rr 1-2-9 was found in a can-trap at 11:03 a.m. Cloacal temperature at that time was 24.3 degrees. She was released and after being free to bask was noosed at 11:47 a.m. At this time the cloacal temperature was 35.6 degrees. In 44 minutes the body temperature had increased to near the ma.ximum temperature voluntarily tolerated by this population. At 12:45 the lizard was sought for a further temperature observa- tion and was again found in the can-trap and with a cloacal temperature of 2.5.4. It is not known how long it had taken the lizard to lose that much heat, but it is known to be less than an hour.
We have observed that earlv in June and after the middle of .August the lizards are more often seen on the ground. During the warmer part of the summer, lizards are seen in trees more often than earlier in the spring or later in the summer and fall. It is also significant that can-trap captures drop off noticeably dur- ing the wanner part of the summer.
On Rainier Mesa the air temperature seldom if ever exceeds .32°C which is within the ac- tivity range of S o. low^ipes. The lizards warm themselves to the desired body temperature and
then simply move into a shaded area. This is not possible for other lizard species in the nearby desert flats where air temperatures may and soil surface temperatures do exceed toler- ance. They must go underground or find heavy shade where it is cooler. Many observations of such actions have been noted. During late June and through July and early August the fence lizards spend a great deal of time in trees. They typically move from areas of sunlight to shade to maintain their desired body tempera- ture and feed on insects that are on the trunk and branches of the tree. Our observations as reported above definitely indicate the possibility that lizards probalxilv do spend the night in trees during the warmest part of the summer. Thev may return to the ground to spend the night in the safety of crevices in the rocks; but, if they do, thev appear to go directly to the trees in the morning with very little, if any, basking on the ground or on rocks.
S. o. Ion<ii])cs exhibits a distinct change in color with changing temperature and light con- ditions. Lizards emerging from their hiding places on cool mornings appear to be nearly a solid black. During the warmer part of a sum- mer day they are lighter in color and the dark pattern markings on the back can be seen in contrast to the lighter ground color.
Rasking lizards, usually observed in the morning hours, are darker in color than those which have reached their optimum tempera- ture and do not need to remain continuously in the direct sunlight. Lizards observed in shade or broken shade or those taken from can-traps are invariably lighter in ground color than those observed while basking. Although some excep- tions may occur, it is thought that the darken- ing of the skin is brought about by a combina- tion of low body temperature in the presence of light and that it serves a useful function in increasing absorption of solar radiation so that the lizards removed from can-traps (while they are still cool and still have the light color pat- tern) and placed on a rock for basking require only a few minutes to change to the darker or "basking" color phase. Although there is a mini- mum of conditions that effect such changes ( such as air temperature, intensity of light, body temperature), lizards permitted to bask for about five minutes have developed the basking color phase.
The length of time a lizard basks or retains the basking color phase varies with the season and the fluctuations of daily weather. In .'\pril and into May basking may be an all-day activi- ty. However, as the days become longer and
16
BniciiAM Young University Science Bulletin
wanner there is a rediiction in tlie time that one may expect to see them in full light on the top or side of a rock or tree trunk. A lizard usually does not leave its basking area but merely seeks the shaded side of a rock or the shade of an ovcrluiiii'liis/ hush. We soon dis-
c-overed that basking lizards were more easily noosed than those who were warm and in shaded areas. The highest temperatures record- ed came from backing lizards. Those noosed later in the day from partially shaded areas were usually between 30° and 34°'C.
REl^liOlJUCTION
Male Cycle: We haye not been able to collcel large series during the months of April and May, but did collect Scelojwnis occiclentalis from Rainier Mesa during the first week of April 19aS and on 12 April 1^)69. Tlie tempera- ture was 7.2°C at noon (4 April) and there was snow in patches on the nortli side of trees and rocks. The April samplings produced only male lizards, indic'atirig that males emerge from hibernation earlier than the females. Woodbury and Woodbury ( 194.5 ) state that the males of Scelo])orus f^raciou.su.s emerge about two weeks earlier than the females and establish territories during that period. If this is the case with S'. occidcntiilis, they may haye just emerg- ed, for there was no indication that territories had been established. It was found that several males were basking side by side; and, on turning over a small rock to capture a lizard known to have taken shelter under it, a total of four male lizards was found.
The males, on emerging from hibernation, have greatly enlarged testes when compared to the ,size of the testes during the summer months. They are sufficiently enlarged as to be easily detected by palpation. The testes of some lizards collected in April and early May constituted as much as 4-5 percent of total body weight. His- tological examination of the testes of lizards taken in .April shou- spennatogenesis with many spennatids, but no spermatozoa.
On 5 May 1968 a series of seven lizards was collected, including one female. The testes or the May series were larger but made up a small- er proportion of the body weight because the lizards were older and had a greater average body weight. The testes and epididymi in this scries contained many mature spermatozoa; in fact, the epitlidvmi were greatly enlarged and compacted with spennatozoa. Mating therefore must occur in late April and May, with some late mating possibly (x-eurring in early June. No mating has been observed in June, although some courting behavior was observed.
In adult male lizards the testes decrease in size from their inaxinmm in ,\pril to ;i minimum
during July and August (Fig. 12). The testes begin to enlarge in late .\ugust and continue to enlarge until the lizard goes into hibernation. The limited data available for late September and October indicati's that an increase in the size of the testes continues to the time of hiber- nation. An examination of the histological slides of the gonads shows a progressive thickening of the seminiferous tubules from late August into October. In earh' September, tubules have approximately 9 or 10 circles of cells; however, by early October several more rows have been added, and it appears that most of the cells which are to be used in the fonnation of mature spennatozoa the next spring are fonned. We note 14-16 rows of cells in the tubules in mid- April. The maximum size is reached in April, although a late spring such as 1969 may push this part of the cycle into early May. We did not find mature spermatozoa in the epididymus until early May (196S). A.ssuining that 1968 was at least an average spring if not earlier than most, we believe mating occurs primarily dur- ing the month of May. For exceptional years it may occur in late .\pril or be continued into early June.
Correlated with the enlargement of the testes is the enhirgement of the fat-bodies. Tliis paral-
e60O .
Imi;.
12. rhf weight of testes plotted in milligram.s.
I liese ilata were olitaiiied from lizards taken in .\|)ril and through tlie sniniiier and into October. .\ disirepanev of ;il)ont ()..'> milligram occurs be- luceii till' (Xtolier .uid .\|)ril wciglits.
Biological Series, Vol. 15, No. 4 Scei.opohus Occidentalis and Uta Stanshoriana
17
lei cknelopiiR'iit begins in late Augu.st and con- tinues into April. A.s the testes reacii their nia.xi- mum size, so also do the fat-bixlies. They begin their decrease at this point and are greatly re- duced by the end of tlie mating season. There is a general belief that fat-bodies function as a storage of energy for the deyelopment of re- productive products. This may be true in some species of lizards; howeyer, our data indicates that the fat-bodies remain large until the sper- matozoa begin to mature. At this time, the epi- didymus becomes enlarged and the testis begins to lose volume. Our in\estigations have not been extensi\e enough to preclude the possibility of some energy flow from fat-bodies to gonad; however, the anatomical changes and physio- logical responses have indicated that much of the stored energy is used by males during the mating period. It is during this time that both the testes and fat-bodies begin to decrease in volume and the epididymi are filled with ma- ture spermatozoa. After the first of June, the volume of viable spemi is greatly reduced and the testes rapidly become smaller. July through early August is a period of testicular (juiescence, and the fat-bodies are small if at all present. By late August an increase in the size of the testes is obscr\able, and the fat-bodies are again beginning to develop. Thus a new cycle has begun.
Female Cycle: As noted above, males emerge earlier in the spring than do females. We have observed that until approximately the first of May considerably more males are seen than females. Our observations indicate that few females appear until late April or early May. This delay seemingly pemiits the males to es- tablish or reestablish a territory before the emergence of the females.
The earliest spring date for the examination of the female cycle i.s' 5 May 196S. At this date, two adults (s-v 74 and 77) were examined. They had follicles measuring 5.3 and 2.7 mm in diameter, respectively. Data available do not indicate the extent of egg volking, if any, nor the amount of volking, if any, that may occur in late September and shortly before or by the time they enter hibernation. However, we have detennined that volking is not extensive in the fall nor in females we have examined in early spring. It is soon after emergence in late April, during May and June, and until the time of egg laving in early July that a continuous yolking and growth of the eggs occur.
We also note that some follicles which are included in those clustered around the ovary do not yolk. These are usually retained as small
tojlicics; however, in some females a few may begin \olking and remain in the ovary after the yolked eggs are laid. Such a phenomena caused us to speculate that perhaps more than one clutch is laid per year. We have found no data to support such a supposition, neither by sacrificing lizards continuously during June through mid-August nor from our data gathered from marked individuals in our study plot.
In the females the fat-bodies increase in size during the fall months and are large when thev emerge from hibernation in the spring. The reduction in size occurs during the period of egg yolking. We have concluded that the energy stored in fat-bodies is used primarily to provide a rapid yolking of the eggs and is perhaps occurring at the same time the males are expending considerable energy in the pro- tection of territories as well as in the (juest for mates and mating. The reduction in the size of fat-bodies occurs simultaneously in the two sexes, and by July both the gonads and the fat-bodies have been greatly reduced in size. .Although we have not calculated the weight differentials for the fat-bodies during the year, they appear to have a cycle which is not only similar to but parallels the seasonal changes ob- served in the gonads (Fig. 12).
Two methods of detennining the time of egg laying were used. Whenever female lizards were captured on the study plot, they were palpated to determine whether or not they were gravid. If eggs could be detected, the fact was recorded. If a female was captured during the season of egg laying and was found not gravid, a note stated that she was not gravid. In this way succeeding captures gave fairly precise infonnation as to the time of egg laying. Time of egg laying is known within two days for sev- eral females. For others, the time between cap- tures is longer, and only close approximation of the date is possible. The other method used to give an indication of laying time was the capture of lizards from outside the plot to be sacrificed for examination of the reproductive tract or placed in terraria until their eggs were laid. For example, from a group of four gravid females captured 14 June 1967, two were sacri- ficed for examination and two that appeared to be nearer to laying were placed in terraria and fed until they laid their eggs. Of the two that were opened, one had 15 oviducal ova and the other had 12. Of the remaining two, one laid 13 eggs on 28 June 1967. These eggs were placed in a glass jar containing about an inch of moist silica sand, and a mason jar lid with the central disc inverted was tightly screwed
18
KmcuAM VouNc Univkhsity Science Bulletin
on it. Tliis provided a c-ontainer that would retain nioisturc but was not c-omplett'ly air tight. Tlu- jar containing the egg.s wa.s then placed on a shelf in the laboratory where all eggs hatched 25 August 1967. The room tem- perature although not constant ranged l)etween 70-75° F. The weights and measurements of the eggs are shown in Table 2. The other lizard did not lav her eggs until 14 July 1967. The clutch consisted of 14 eggs and weighed 7.48 grams, with an average weight of 0.53.5 grams each. This compares with 0.6.30 grams each in the previously mentioned clutch. The second female apparently retained her clutch much longer than she would have in nature. It mav also be that she was undernourished at a time when nutri- tion was critical in the development of the eggs since she did not ade(juately feed during her confinement. None of the eggs hatched, al- though each showed the pink area that indi- cates a developing embryo at the time of laying.
Talilr 2. Number and weight of ova at time of laying (1967) for three clutehe.s laid in the labo- ratory.
|
Number of e in clutch |
KK-- |
W |
eigl |
it ot chitc |
.\\eraire weight h per egg |
|
13 |
8.19 |
0.630 |
|||
|
14 |
7.48 |
0.53 |
|||
|
10 |
4.22 |
0.42 |
|||
|
Aver. 12..3.3 |
6.a3 |
0.538 |
The earliest date recorded for egg laying in the field was for li/ard #5-14, captured 5 June 1967. At the time of capture this li/ard was gravid. When recaptured on 10 June l(-)67, pal- pation revealed no eggs. During this time, she had also suffered a slight injury. A cactus spine had punctured her lower evelid and was still in the lid when captured. The spine did not damage the eye but did indicate that she may have been digging in the loose soil around a cactus plant for the purpose of laying eggs. This same li/ard laid eggs sometime after 24 June 1966. The 1966 season was, in general, about two weeks later than the 1967 season. Aside from the case of lizard #5-14, the earliest records of egg laying in the field for 1966 and 1967 wcTc 14 June and 24 June, respectivelv. There are two records for each date. The latest record (±t.3-16) of a female still gravid is 10 July 1967, and the eggs were laicl before her next capture on 12 Jul v.
Fat-bodies taken from female lizards during the .\pril to .September period show a gradual
increase in size and weight beginning about one month after egg laving (in August) and con- tinuing until the next spring (late April to earlv June), when the fat-bodies are gradually reduced in size until the eggs are laid. At the time of egg laying, the fat-bodies are small or coiupletelv spent. The reduction in si/e is cor- related with the \c)lking ot the eggs and ap- parentlv serves as a ready source of material for yolking. The fat-body buildup begins again after egg laving and is a parallel development with the gonadal cvcle.
A comparison of the fat-body cvcle in males and females indicates that in males the stored energv is used during the mating period for such activities as aggressive defense of his ter- ritorv and perhaps seeking females and mating. During these activities, males may not feed normallv; thus, the stored energy is available to provide for or iiLsure reproduction. In con- trast, the more passive female, during this time, is a ravenous feeder and thus uses not only stored fat-bodv energy but also food energy to volk the eggs. The above explanations, al- though perhaps not a complete analysis of fat- bodv function, fit generally our data and ob- servations.
Twenlv female li/ards were collected near the studv plot on 1.3 June 1968. Sixteen were gravid and four appeared not to be gravid. Three of the later four, however, were yolking small eggs of approximatelv 3-4 mm in diameter and the fourth contained what appeared to be 9-10 atretic follicles. Those considered to be gravid were fed in the laboratory. Nine laid eggs lip to 12 July U)68. It is suspected that females kept in the laboratorv hold their eggs longer than would be the case in nature.
Based on the data concerning egg laying and ONulation, it appears that the female cvcle compares favorably to that of the male in that the yolking of eggs begins at approximately the time of the fonnation of mature spennatozoa in the male. .Assunu'ng that there is some delav in fertili/ation, the female cvcle is at the ap- propriate stage for mating during late April and especiallv so during the month of Mav. Accord- inglv, ovulation takes place in late Mav and earl\' June, with the first records of egg laving on 14 June 1966 and 24 June 1967. Egg laying in 1968 corresponded closelv to that of 1967. Thus, we mav expect that eggs will be laid between 10 Jvme and 10 Jiilv. In a small popu- lation such as the studv plot on Rainier Mesa, seasonal fluctuations will affect these dates; however, we have lound that, (or a single si'ason, egg laving is completed in about three
Biological Seiues, \'ol. 15, No. 4 ScELorouu.s Occidentalis and U ia Stansirihiana
19
weeks; the dates given above represent the extremes.
Size of individual eggs and the numbers in a chitch
Sixty-five ova were measured soon after they were laid in the laboratory in 1967. The average length for all was 1.3.96 mm with an average diameter of 8.165 mm. Three clutches were weighed soon after they were laid. A total of 37 eggs averaged 0.53S g (Table 2).
Clutch size has been determined by retaining gravid females in the laboratorv until the eggs have been laid and by sacrificing obviously gravid females and counting the yolking or oviducal eggs. These methods were necessary because eggs in nature are most difficult to find. Although we have looked for eggs in the field, we ha\e not found a single clutch. The largest clutch layed contained 15 and the small- est 7 eggs. There is a correlation between the number of eggs produced and the size of the female. Small females arc presumed to be the younger ones in the population and to produce fewer eggs their first reproductive year. Two lizards S3 and 85 mm s-v, laid 14 and 15 eggs, two, 76 and 77, respectively laid 11 and 10, and two, 72 and 73, each laid 9 eggs (1967). Data concerned with clutch size, dates of egg laying (in the laboratorv) and the size of some females are presented in Table 3.
Data summarized from 15 clutches laid and from oviducal eggs examined in sacrificed liz- ards indicate that we might expect an average of 11 to 12 eggs to be laid by each adult resident
Table 3. Clutch size and date liid. Females were from Rainier Mesa, near the study plot, and all eggs were laved at the Mercury base (CETO) laboratory-.
Table 4. Population stnieture based on marked lizards 1965-1967.
Year
|
Eggs in |
Size of female |
Date |
eggs |
|
clutch |
were layed |
||
|
14 |
83 |
28 June |
1967 |
|
15 |
85 |
29 June |
1967 |
|
11 |
78 |
25 June |
1968 |
|
11 |
76 |
27 June |
1968 |
|
10 |
77 |
.30 June |
1968 |
|
9 |
73 |
26 |ime |
1968 |
|
9 |
72 |
30 June |
1968 |
|
14 |
82 |
.30 June |
1968 |
|
10 |
-- |
4 July |
1968 |
|
15 |
4 July |
1968 |
|
|
7 |
-- |
13 July |
1968 |
|
9 |
-- |
14 |uly |
1968 |
|
15 |
16 June |
1969 |
|
|
10 |
17 |une |
1969 |
|
|
9 |
17 June |
1969 |
|
|
168 |
Marked males Adults Juve.
Marked females Hatchlings
Adults Juve. Total marked
|
1965' |
14 |
17 |
22 19 |
72 |
0 |
|
1966= |
29 |
4 |
40(31) 6 |
79 |
76 |
|
1967' |
26 |
19 |
31 23 |
99 |
25 |
'Study plot not opened until 12 July ■'First hatchliug marked 13 August First liatchling marked 2-3 August
female on the study plot. Although we have found a yearly variation in the numbers of females in the study plot, as noted in Table 4, we believe that these fluctuations in numbers can be explained partly on the basis of our inability to mark the total population and also from the fact that tliere is some immigration and emigration. In 1965 we marked the first lizards on 13 July while we were setting up the plot. Bv mid-July most of the large adults are in the Pinyon or Juniper trees; they thus spend less time on the ground and were not caught as often in our can-traps; also while in the trees they are much more difficult to see or catch.
The tact that some immigration and emigra- tion occur, particularly among juveniles and fe- males, may influence numbers present during a summer si'ason. We invariably marked more adult females than males during each year. Those marked lizards found from 30 to 50 yards from the edge of the plot were with one exception females or juveniles. Furthermore, in 1966 we recaptured 17 of each sex that were marked in 1965; however, only 11 other males were marked while 23 unmarked adult females were marked. We also noted that a number of females were seen in the plot (1966) for only a short time, usually during the egg laying season. For example, #4-7 and #4-17 were marked at stations [-7 (edge of plot) and B-9 (50 feet inside plot) on 2 and 12 June 1966; it was noted that each was gravid. They were not seen again until 29 and 27 June 1967 at which time they were caught at J-9 and A-8 (west and east edge of plot), and again both were noted to be gravid. Other examples of gravid fem;des captured on the plot once or twice during a short period of time, usually less than a week, are as follows: 3-19, 4-5, 4-8, 4-16, 5-11, 5-12, and 5-17. This would account for nearly half of the increase in females observed on the plot in 1966. We, of course, expect that some of the females living on the plot may have left the plot for egg laying. However, we know
20
Bhioham Y'ouno Univkrsitv Science Bulletin
reasons, we observed more adult females than males on the plot. Assuming that the tcrritorial- ism of males keeps them within a home range and tlnis with much k'ss slutting ahout. we can arrive at a near balanee between males and females by subtracting 9 from the 40 adult fe- males known to be on the plot at some time during the summer, giving us a near balance of 29 males to 31 females. In 1967 we did not find a large number of females (3) coming onto the plot during June. Our only explana- tion of this is that there was a large population of juveniles in UJ67 and tew in 1966. Whether this increase in population at this level of the population affected their nesting habit is onlv speculation.
If we assume that the adult temak' popula- tion on the study plot ranges from 25 to 35 females, then we may estimate that during an average year approximatelv '30 clutches of eggs will be laid. In other words, between 30<) and 400 eggs are produced to insure a continuous population on a si.x to seven-acre area.
Hatchlings
The tcmi hatchliiig is used to designate that part of the life of a lizard from the time of its hatching in August and September until it enters hibernation for the first winter. In order to designate them as separate entities, the next spring they are referred to as juveniles.
The earliest record of a hatchling in the field is 9 August 1966. (There were no hatch- lings found on the plot in 1965; plot closed 9 September.) Two more were taken 13 August, and newly hatched lizards were still being cap- tured when the can-traps were closed 4 Septem- ber 1966. In 1966 we marked 77 hatchlings. The first hatchling recorded for U)67 was marked on 23 August, which seems to corroborate an earlier statement that the season was later in 1967 than in the previous year. New hatchlings continued to appear until September 11 when the plot was closed. In 1967 we marked 26 hatchlings. A brief sampling trip to the plot on 7 October 1967 produced 12 hatchlings during a 24-hour periexl with the can-traps open. One of the hatchlings measured only 29 mm snout to vent and weighed 0.72 g, indicating that hatching does continue into late September during favorable years.
In U)66 there were less precipitation and slightly warmer temperatures (Fig, 4), fai-tors which not only lead to an earlier date for hatch- ing but also a longer period, thus increasing the of none. For this and perhaps other less obvious
possibility for eggs laid toward the end of tin- ni'sting season to be successfully hatched.
Till' minimum size recorded for a hatchling in the fii'ld was s-v 26 mm and a weight of 0.33 g. From several clutches hatched in the laboratory, the smallest was 22 mm s-v, and with a weight of 0.48 g. The largest hatchling known to have been less than 24 hours old was hatched in the laboratory, measured 28 mm s-v, and weighed 0.90 g. Part of this weight can be accounted for by the fact that the um- liilical cortl was still attached. It is seemingly a fact that hatchlings less than 24 hours old and from eggs hatched in the laboratory are generally smaller in snout-xent length and heavi- er in weight than the hatchling lizards captured in the field. This suggests that young hatchlings with a (juantitv of yolk still in the mid-gut may remain relatively inactive for a few days before digging out to the surface; during this time, those hatched in the field may grow in length and decrease in weight. .\t least, such a con- clusion seems plausible based on the weights and measurements ot hatc-lilings hatched in the laboratory in contrast to those examined at the study plot. Tlie size and weight of 20 hatch- lings measured shortly after hatching are listed in Table 5. A summarization of measurements for hatchlings observed on the study plot is recorded in Table 6. In order to indicate gen-
Tahle 5. Size ;iinl weight of labonitorv hatchling.s — mea.surements taJcen 24 hours ;ifter hatching.
|
Number and |
s-\- |
Tot..! |
Weight |
Date |
|
ehiteii |
lengtli |
length |
hatched |
|
|
First Clutch |
||||
|
1 |
24 |
51 |
n.,58 |
8-22-69 |
|
o |
24 |
47 |
0.51 |
8-22-69 |
|
3 |
26 |
55 |
0.76 |
8-22-69 |
|
4 |
25 |
56 |
0.69 |
8-23-69 |
|
5 |
25 |
54 |
0.63 |
8-23-69 |
|
6 |
25 |
52 |
0.63 |
8-23-69 |
|
T |
26 |
54 |
0.68 |
8-23-69 |
|
8 |
24 |
52 |
0,54 |
8-23-69 |
|
9 |
2.5 |
53 |
0.71 |
8-23-69 |
|
Second Chitt |
■li |
|||
|
I |
2:3 |
49 |
0.58 |
8-23-69 |
|
2 |
24 |
50 |
0.67 |
8-23-69 |
|
3 |
25 |
51 |
0.72 |
8-24-69 |
|
4 |
24 |
50 |
0.,54 |
8-24-69 |
|
5 |
26 |
55 |
0.67 |
8-24-69 |
|
6 |
25 |
54 |
0.76 |
8-24-69 |
|
i |
27 |
56 |
0.77 |
8-24-69 |
|
S |
24 |
,54 |
0,72 |
8-24-69 |
|
9 |
24 |
49 |
0.61 |
8-24-69 |
|
Ihiril Clutili |
||||
|
1 |
23 |
48 |
0.38 |
8-26-68 |
|
.1 |
22 |
48 |
0.37 |
8-26-68 |
|
A\erages |
24.55 |
51.90 |
0.626 |
BioLooiCAi. Series, \'()i,. 15. No. 4 Sc:elopohu.s Ocx;ii)eni ai.is .xnd Via St.vn.shuhiana
21
Table 6. A summ;iry of mea.siircnients taken from hatchlings marked on the Rainier Me.sa study plot 1966-70.
|
Number |
S-V |
Total |
Weight |
Date |
|
|
15 |
26(29.73)34 |
52(64.36)73 |
.33 (.653) 93 |
9 Aug. thni 16 Aug. |
1966 |
|
18 |
27(28.83)30 |
57(63.05)71 |
.65 (.787) 96 |
19 Aug. thm 23 Aug. (5 days) |
1966 |
|
23 |
27(29.43)35 |
57(65.52)79 |
.55( .788) 1.2.3 |
25 Aug. thni 29 Aug. (5 days) |
1966 |
|
23 |
27(30.0)37 |
55(65.52)83 |
.50( .886) 1.60 |
31 Aug. Iliru 4 Sept. |
1966 |
|
4 |
29 (29.5). 30 |
63(65.5)68 |
.67 (735). 80 |
23 Aug. thru 29 Aug. |
1967 |
|
9 |
29(30.66)32 |
60(67.11)72 |
.60(862)1.11 |
7 Sept. thni 11 Sept. |
1967 |
|
12 |
29(32.,5«)36 |
63(71.5)80 |
.68(1.06)1.66 |
7 Oft. 1967 |
|
|
14 |
30(33.36)39 |
63(73.79)87 |
.62(1.18)1.97 |
4 and 5 Oct. 1968 |
|
|
8 |
30(34.13)38 |
65(73.0)80 |
.78(1.33)1.75 |
9 iuid 10 Oct. 1970 |
enil ^routh rati'.s, averages for different periods are reported. Data on the growth of hatchlings before their first hiliemation is limited. In all cases of hatchling lizards captured in the field, the exact time of hatching is unknown. Often a hatchling is not captured until it is nearly a full grain in weight, which is not surprising considering their small size and the heavy cover at the studv plot. The fact that we closed the plot by or before 10 September limited our observations of hatchlings. Other data, however, pro\ides a basis for dctcnnining growth (Table 6), and from these data we offer the growth curve (Fig. 11 and 12). The length of time available for late summer and fall growth varies. Some vears pcmiit feeding until mid-October. In 1967, 196.8, and 1970, small series were taken between 4 and 10 October; however, on 17 and IS October 1969, no lizards were seen. Only a few hatchlings recaptured in the season of their hatching showed a gain in weight or length. The need to close the plot in early Sep- tember at the height of hatching gave only limited time for observation. Of those marked in 1966, 21 (27.6 percent) were observed the next spring. There were several hatchling re- captures in 1966 in which no change in length, but a loss of weight, occurred. One lizard show- ing weight gain was numbered 6-16 and was the first hatchling captured in 1966. Her cap- ture dates, measurements, and weights are listed in Table 7. Unfortunately, this lizard was not seen the next spring, as was the case for many other marked hatchlings. Although years vary, it is obvious from the data available that there is a high hatchling mortality each year in this population and presumably in the species.
Since few other lizards were recaptured showing a weight gain in the season of hatching to give concrete data, we conclude from meas- urements taken that the lizards hatch with snout-vent measurements of 24-28 mm, total length of 45-60 mm, and body weights of 0.35-
0.90 g. They may, if hatched early, attain s-v length of 38 mm or more and a weight of 1.6 g before they hibernate, presumably, by mid-Octo- ber. The young remain active later in the autumn than the adults and emerge from hiber- nation about as early in the spring. Of 15 lizards captured on 7 October 1967, 12 were hatchlings with s-v length of 29-38 mm and weights of 0.68-1.66 g, 2 were marked juveniles, 1 male and 1 female, and 1 a marked adult male. On 9 and 10 October 1970, 7 hatchlings (s-v 30-38 and weight 0.78-1.75) and 6 juveniles were examined.
In 1963 we marked 76 hatchlings. Of these, 12 were recaptured in 1966. Twenty-one of those marked as hatchlings were recaptured in 1967. This is a 27.63 percent survival of those marked. However, we marked an additional 45 as juven- iles in 1967. If we assume the same percentage of survival for these, we can account for approx- imately 2.50 hatchlings on the plot in 1966. An attempt to estimate the hatchlings on the plot per year has not been entirely successful. Table 8 gives our estimations based on hatchlings marked the next spring and summer.
The variation in the estimated yearly output of hatchlings is a direct result of the variations in climate. Few summer seasons provide the same weather conditions. There are early falls, late springs, and an occasional cloudy wet sum- mer (Fig. 3 and 4) which provide some of the fluctuations observed in the populations. Such climatic conditions may induce factors which increase or decrease food sources which in turn may increase or decrease the numbers of eggs
Tal)Ie 7. Measurements and weiglits for hatchling number 6-16.
|
Date |
S-V |
Total |
length |
Weight |
|
|
8-9-66 |
28 |
mm |
65 |
mm |
.69 |
|
8-17-66 |
28 |
mm |
65 |
mm |
.65 |
|
8-29-66 |
30 |
mm |
66 |
mm |
.90 |
|
9-4-66 |
33 |
mm |
73 |
mm |
1.00 |
BmciiAM VouNO University Science Bulletin
Table 8. Ksliination of hatcliliiigs procliicrd mi stiiily plot for four years.
|
Year |
Hatchlings |
Juveniles |
Estimated number |
|
|
marked |
marked |
of |
hatchlings |
|
|
1964 |
150 |
|||
|
196.5 |
0 |
36 |
42 |
|
|
1966 |
76 |
10 |
250 |
|
|
1967 |
25 |
45 |
194 |
|
|
1968 |
15 |
22 |
|
prodiici'd a.s well a.s the time of hatching and the succcs.s of the yearly hatcli.
In 1965 the senior author .spent 6 and 9 September at the Rainier Me.sa .study plot in an attempt to Find hatchlings. Although there are undoul)tedlv some Iniman errors in the dates of hatchling emergence and the yearly num- l)ers observed, the fact still remains that there is coiisidi'rable variation. Our observations in- dicate that no two years are the same, with extremes such as 1965 and 1966 occurring at least in high elevation desert habitats.
Juveniles
The period of greatest growth tor voung li/.ards begins with their emergence from their first hibernation, as can be seen in Table 9. As indicated previously, lizards emerging from their first hil)eniati()n are referred to as juve- niles. A lizard remains in this age group until it enters the second hibernation.
As they emerge from the first hibernation, many have not vet reached a snout-vent length of 35 mm, and all are less than 40 mm. By
earlv June, half of the juveniles have reached 40-45 mm, and this growth continues at a rapid rate during June, July, and August. By late August or early September, a few have attained a snout-vent li'ugth of 70 mm and are small adults in size. It is also important to note that bv this date even the smallest juvenile is ap- proaching a snout-vent length of 55 mm and this generation of the population ranges be- tween 5.3 and 71 mm in the snout to vent length. During late August and September, their growth continues but at a slower rate. This reduction in growth rate is particularly true for larger individuals, and this tends to produce a more uniform size for this age group in October. The data for the variations indicated above and for figures 13 and 14 are based primarily on observations made during the summers of 1985, 1966, and 1967. but with data obtained in 196S, 1969, and 1970 used to supplement the previous years, particularlv that for late fall and
Table 9. Measurements and ueight for juveniles in May and June compared \Nith August and Sep- tember.
|
Minimum |
Miiximum Average |
|
|
May and June |
||
|
.Siu)iit-\('nt |
32 mm |
54 mm 43.9 mm |
|
Total Inigth |
72 mm |
118 mm 99.0 mm |
|
Weight |
0.7 g |
6.0 g 2.36 g |
|
August and September |
||
|
Snout-vent |
.52 mm |
71 mm 67.4 mm |
|
'I'otal Icngtii |
100 mm |
165 mm 140.6 mm |
|
Weight |
4.9 g |
14.0 g 8.7 g |
80 .
t70.
«
6
^60J
C
-C O)
c
j; 5o_
>
I
40_
30
• • •
Jun (
July Aug
Fig. 13. (irowtli (i( ju\fTiilcs plotted from data gathered from June 1. to inid-Se]>temi)er.
Sept. 1 -12
Biological Series. Vol. 15, NO. 4 Scei.opokus 0<:c:ii)en iai.is
AM) Uta Stansbuhiana
23
25
100
2 0 80
15 60
E o
6
c
a>
„. IIIIIIUI.I"''*""""
|
o gt h in Mil |
40 |
....■•••••••' |
»»» |
c O 'o c 0) -Q X |
'''/^ |
|
5 I |
20 |
•^ |
/ |
||
|
-J |
/ |
||||
|
> 1 |
/ |
||||
|
0 |
0 |
— -' ' |
Aug.toOcT. June to 0«t. June June June SOM0.+
1966 1967 1968 1969 1970
Fig. 14. A .sumnuirization of data .showinj; i^rowth in the Rainier Mesa population from liatchlini; (1966) to juvcniie.s and adnlts throut;li 1967 to 1970. The broken line indicates snout to \ent lengths and the solid line growth as determined hv weight in grams. The tliscrepaneies following the first and seeond hihemations represent growth during April and May.
early spring. Because this data represents a summarization of three years, there may be a greater variation in size than would be encoun- tered during anv one vear. The effect of an early or late autumn or spring would either favor or retard growth during the earlv periods of the hatchling and juvenile growth cycle. How- ever, by mid-September, or by the end of ap- proximately one full year, the lizards of this population have reached the size of small adults. It is further believed that with few exceptions, none of which we are aware of, the juveniles, now young adults, will reproduce the next spring for the first time.
A further examination of figures 13 and 14 indicates that the growth curve after emergence from the first hibernation would be about 35 mm s-v and would slope steadily upward into mid-August at which time a flattening of the curve occurs. It is also ol)vious that the average size at this time is approximately 64-65 mm. We are unable to note any significant difference in the rate of growth between the males and females.
In Table 10 a summarization of growth and rate of growth is given for 25 juveniles examined during the summer of 1967. Although the rec- ords are not complete for each individual for each month listed, there are data available to indicate that these individuals had a net growth of 428 mm in s-v length and 1.30.76 g weight in a total of 1633 days. For this year, at least, the approximate daily growth from 1 June to 10 September was 0.2530 mm in s-v length, with a weight increment of 0.08052 g.
In tenns of percentage of growth, lizards 7-13 had a 50 percent increase in s-v length, 14-15 had 70 percent, 16-19 a 65 percent, and 17-20 a 68 percent. Data available for other years also indicate a snout-vent increase rang- ing between 50 and 70 percent, with the ap- proximate averages ranging between 60 to 65 percent. The variation in the percentage of s-v growth from 1 June until 10 September is ap- proximately .50 to 70 percent for 1967.
A series of hatchlings secured between 9 and 16 October 1970 had .s-v lengths of .30(.34.13)38 (Table 6). The series of 10 juveniles taken in
24
BmciiAM Young Univeksity Science Bulletin
June 1967 (1-15, Table 10) measure out at 32(40.1)43, whereas a series of 8 taken in June (1-13) 1968 measured ,34(44.6).50. On the basis of these and confirming data for otlier years, we conclude that most hatcldings liibernate before reaching 40 mm in s-v length and emerge in April ( 1-15, depenchng on year) at about
this same length. We recognize that these fig- ures will vary from year to year and suspect that few juveniles reach 50 mm in s-v before mid-June. During favorable springs, growth be- gins in April and continues to accelerate during \lay. Juvenile growth is rapid and uninterrupted until they reach at least 60 to 65 mm. Thus
Table 10. Growth during .summer (1967) for marked lengtli; .second figure — weight in gram.s.
liz.ird with ,i known age. First figure — snont-vent
|
.Specimen |
I).\TES onsEBVED |
Total |
Days of |
% |
||||
|
numlier |
June 1-15 |
June 16-30 |
July |
August |
September |
growth |
growth |
Day |
|
7-13 |
42-2.60 |
5.5-5.51 |
60-7.44 |
64-8.54 |
22-5.94 |
100 |
.220 .0594 |
|
|
8-17 |
51-3.79 |
60-8.81 |
67-9.75 |
69-10.45 |
18-5.94 |
75 |
.240 .0792 |
|
|
10-11 |
43-2.69 |
48-3.74 |
64-8.81 |
21-6.12 |
89 |
.236 .0688 |
||
|
14-15 |
.37-1. .58 |
43-2.62 |
.53-4.67 |
.57-6.38 |
63-9.09 |
26-7.51 |
94 |
.276 .0799 |
|
16-18 |
44-2.59 |
53-4.58 |
62-8.34 |
18-5.75 |
65 |
.277 .0884 |
||
|
16-19 |
40-2.26 |
45-3.31 |
60-7.61 |
65-9.98 |
66-10..54 |
26-8.28 |
91 |
.286 .0909 |
|
17-20 |
37-1.80 |
42-2.40 |
51-3.70 |
59-7.70 |
62-8.63 |
25-6.83 |
95 |
.263 .0719 |
|
18-19 |
32-0.70 |
44-2.,50 |
12-1.80 |
33 |
.363 .0545 |
|||
|
18-20 |
42-2.42 |
51-4.25 |
9-1.83 |
39 |
.231 .0469 |
|||
|
19-20 |
41-2.42 |
46-3.28 |
.5-0.86 |
19 |
.263 .0453 |
|||
|
1-2-3 |
43-2.64 |
.59-7.06 |
66-9.65 |
2.3-7.01 |
88 |
.261 .0796 |
||
|
1-2-7 |
45-3.00 |
62-8.14 |
63-9.31 |
18-6.31 |
88 |
.205 .0717 |
||
|
1-2-9 |
44-2.95 |
61-7.46 |
17-4.51 |
55 |
.309 .0820 |
|||
|
1-2-10 |
45-3.12 |
61-7.86 |
64-9.84 |
19-6.72 |
84 |
.227 .0800 |
||
|
1-2-12 |
43-2.83 |
46-3.50 |
.57-7.15 |
14-4.32 |
69 |
.203 .0626 |
||
|
1-2-13 |
4.5-3.24 |
6.5-9.18 |
20-5.94 |
55 |
.363 .1080 |
|||
|
1-2-1.5 |
45-2.74 |
.5.5-5.56 |
58-6.68 |
63-9.75 |
18-7.01 |
83 |
.217 .0845 |
|
|
1-2-17 |
43-2.26 |
63-8.10 |
20-5.84 |
62 |
.323 .0942 |
|||
|
1-2-18 |
44-3.29 |
63-9.13 |
19-5.84 |
67 |
.283 .9872 |
|||
|
1-2-20 |
44-2.52 |
62-8.80 |
18-6.28 |
62 |
.290 .1012 |
|||
|
1-3-4 |
46-3.27 |
6.5-9.21 |
19-5.94 |
49 |
.387 .1204 |
|||
|
1-3-9 |
46-2.98 |
.54-5.59 |
61-7.93 |
63-9.30 |
17-6.32 |
78 |
.218 .0810 |
|
|
1-3-10 |
51-5. 16 |
56-5.89 |
65-9.95 |
14-4.79 |
53 |
.264 .0904 |
||
|
1-3-14 |
52-4,82 |
62-7.89 |
10-3.07 |
40 |
,2.50 .0767 |
|||
|
25 |
32(40.1) |
42(45.7) |
44 (.53.7) |
46(61.19) |
57(63.63) |
428(17.12s-v) |
1633 |
days |
|
44 |
51 |
60 |
67 |
69 |
130.76 (5.23) |
growth |
per day |
|
|
1.. 58(2.17) |
2.66(3.73) |
2.5( 5.27 ) |
3.5(8.09) |
7.15(9.3) |
s-v 0 |
.2.530 ' |
||
|
2.69 |
5.16 |
8.81 |
9.98 |
10.54 |
wt. 0.08052 |
Biological Series, \'ol. 15. No. 4 Sceloi'OHUS Occidentai.i.s and Uta Stan.shuhiana
25
from Juno to Septoinher 15 the juvenile lizard increases its s-v k'ugth b\' about 65 percent and it is about four times as heavy.
As with adults, juveniles spend considerable time basking in the spring and fall and during the earlv morning hours throughout the summer. Temperatures taken indicate tliat this age group may maintain a higher average body tempera- ture for more Iiours of the dav than is the case for adults (Table 6). By controlling the tempera- tures tlirougli basking, and with an abundance of food as indicated below, the rate of metabo- lism is perhaps greater, thus providing for the rapid growth rate observed in juveniles. We also note that the greatest rate of growth (in 1967) occurred during June, Julv, and the first part of August but with a noticeable slowing down during late August, September, and October (Fig.' 13 and 14 and Table 10).
Sexual matiiritv is reached by September or October just before the juveniles enter their second hibernation. We note that in males the gonads arc increasing in size as indicated in figure 10 and that such males do produce mature sperm the next spring. Although most juveniles measured during September and Oc- tober are not yet 70 mm in s-v length a few have reached the size of small adults, and most reach 70 mm bv or before June of the next spring.
Adults
Lizards that are sexually mature are con- sidered to be adults. This stage in the life cycle is reached in most males as they enter the second hibernation. We have no data to indicate that anv remain as juveniles after they emerge from the second hibernation. Size is also a factor and, although we have previously in- dicated a size of 70 mm s-v to signify adult- hood, some juveniles, particularly males, emerge as sexually mature individuals with a size slight- ly less than 70 mm.
Two males taken on 9 April 1968 had s-v lengths of 64 and 67 mm; yet both had greatly enlarged testes and were considered to be breeding males. By late April and certainly be- fore eggs are laid in June, few if any sexually mature lizards are less than 70 mm in s-v length.
Adults reach a s-v size varying between 70 and 92 and in total length from 150 to 199 mm. Lizards of this size have a \ariation in extreme weights of approximately 10.0 to .31.0 g. The longest lizard measured was a female; however, the heaviest was a male.
Many adults (18 percent in 1967) have re- generated tails. This remarkable phenomenon
of regeneration pemiits the lizard to duplicate almost perfectly tlie lost part, at least insofar as size and outward appearance is concerned. The real value of the tail to these lizards is perhaps not entirely known, but the fact that it is regenerated so completely and at a rapid pace suggests that its presence is of value. Tinkle (T967) in his considerations of tail re- generation in Uta believes that tail breakage is a part of an escape mechanism as well as an organ for balance. Similar functions for the tail are perhaps also true for Scclo])oms. We note that the tail may also be used by these lizards for the storage of food. At least lizards deprived of food lose considerable substance from the tail before there is a noticeable reduc- tion in the vigor of the individual. Other func- tions may yet become apparent.
The rate of regeneration is variable. Our data suggests a more rapid rate during the summer when the rate of metabolism is generally high and the days are longer.
Figure 14 indicates a growth curve for five years. The curve includes a large number of individuals for the first two years but a con- tinuallv reducing number thereafter. These curves are also based on averages. Size variation among individuals of a given age group even after two years is considerable and is known to vary at least 10 mm in s-v length. The fact that averages of age groups show a continuous, although slower, increase in size suggests that we might expect most individuals to continue to grow as long as they arc in good health.
We have noted that the older lizards seem to hibernate before the hatchlings and juven- iles. However, two adult lizards taken in Octo- ber 196.S and 1970 were not healthy. The latter, ii:7-L5, was marked as a hatchling on 17 August 1966 and recaptured for the last time on 16 October 1970. At the last capture it was 85 mm. in s-v length and at least 50 months old. Al- though vigorous, it was emaciated and in poor health. An autopsy revealed an obvious intes- tinal infection.
Our study did not cover sufficient years for us to state with certainty the longevity of the oldest individuals of this population. It does indicate that not many live beyond the fifth year and relatively few are known to have reached an age of five years. Dat;i for lizards marked as hatchlings or juveniles (where an age can be estimated within a few weeks) have given us several females known to have been gravid for four consecutive years. Such individu- als would be at least six years old and we suspect some to be older. There is, therefore,
26
BnicHAM Young University Science Bulletin
suFficicnt data to demonstrate a lone;evity of five to six years hut with a progressive and rapid thinning of individuals of the age groups older than four vears.
Young adults hreed and lay the first cluleh of eggs when they are 21 lo 23 months old. This is in contrast to I'ta who lay eggs the first year. Thus, when lizards are approximately two years old, their first generation of hatehlings are emerging (August and September). We have indieated that a pressure is seemingly exerted on the adults by the large numbers of hatehlings and juveniles which appear in the population each year. If such pressures are assumed lo be responsible for maintaining a relativelv unih)rin balance in the size of the population, we would expect that anv reduction in the numbers of the vounger age groups would be reflected bv slight variations in the numbers of older lizards in the population. Such general conclusions fit well except for 1965 and 1966, two years in which considerable variation in numbers occurred in the vounger age groups (Table S). However, we did not record a no- ticeable reduction in the numbers of the older age groups in the spring of 1966, and tlie larg- est number of hatehlings was observed that summer (1966) than tor anv other vear. This mav not necessarilv be tlu- result of more and or larger adults; liowever, it does suggest an inherent population resilience which permits the population to maintain an adult breeding population capable of withstanding adverse conditions for short periods of time and thus to maintain a breeding population.
In Table 1 1 the approximate date of egg laying and the differential in weight of females before and after egg laying is given. We at- tempted to correlate these weights and meas- urements with thost- taken from actual clutches laid and measured in the laboratory. Although we can see some similarity in weights, it is not possilile lo cslimale accuratelv the number of
eggs laid based on the difference in weights of a female before and after the eggs are laid in the field. Several factors interfere, such as the difference in the size of eggs, the loss of fluid at ihe time of laving and general reduction in bodv weight which may have resulted trom energv spent in "nesting" or the reduction of or abstinence from feeding during the period im- mediatelv preceding and following the laying of eggs. In two instances (Table 11), females weighed two and seven days between egg laying have shown a loss in weight of more than 11 grams. We do not have records of entire clutches weighing more than 8.26 grams, and it is therefore considered to be more reasonable to assume that the three plus grams in weight loss resulted from other causes as indicated al)o\c However, both females were large and may have laid more than 14 or 15 eggs, which is the largest number observed bv us in anv one clutch. If larger clutches are laid this mav also account for the discrepancies in weight. Fitch ( 1970) reports the largest clutch for this species to be 13 eggs.
Population Structure
The structiuc of the population \aries from vear to vear ( Table 4 ) . Theoreticallv, one would suppose that the numbers of hatehlings, juven- iles, and adults could be estimated if the ap- proximate population stnicture is known for a tew vi'ars. This mav be possible within broad limits it the adaphic factors of the environment were less variable. However, with the consider- able variation not only in annual rainfall but also with the considerable fluctuation in tem- perature, particularly in some of the more criti- cal months, estimations are seemingly not suf- ficientlv meaningful to be worth the effort, par- licularh' it one atti'uipts to evaluate the entire popukition. Perhaps the most stable part of the population is the adults; at least we have found less variation each summer in the number of
T.il)lc 11. Field data showiiig uoight difffrcntials of female liz.irds liefiire and after eggs are laycd.
|
Date |
Weight |
Date not |
Weight |
Days |
Difference |
S-V |
|
|
Toe number |
gravid |
gravid |
gravid |
after laying |
between |
in weight |
length |
|
.3-17-18 |
6-18-67 |
29.23 |
6-27-67 |
20.79 |
9 |
8.44 |
90 |
|
.3-16 |
7-10-67 |
25.80 |
7-12-67 |
16.88 |
."> |
8.92 |
80 |
|
■!-!() |
6-20-67 |
28.86 |
6-27-67 |
17.21 |
7 |
11.65 |
86 |
|
4-14 |
6-18-67 |
25..56 |
6-24-67 |
17.83 |
6 |
7.73 |
87 |
|
4-18 |
6-20-66 |
21.01 |
6-22-66 |
15.02 |
.-) |
5.99 |
83 |
|
1-.5 |
6-7-67 |
19.27 |
6-29-67 |
14.61 |
->o |
4.66 |
83 |
|
5-1.5 |
6-7-67 |
20.74 |
7-12-67 |
17.68 |
35 |
3.06 |
80 |
|
.5-20 |
6-16-67 |
23.67 |
6-28-67 |
18.99 |
12 |
4.68 |
85 |
|
6-9 |
6-20-67 |
19.79 |
7-12-67 |
1 5.08 |
.-).-) |
4.71 |
80 |
|
1-2-19 |
6-22-67 |
29.92 |
6-24-67 |
18.02 |
2 |
11.72 |
89 |
Biological Series, Vol. 15, No. 4 Sc;ELoroHus Occhjentalis and Uta Stansduhiana
27
adult.s, with a. greater variation in the number of .surxiving juveniles and liatehiings.
Based on our can-trap counts, five small terrestrial vertebrates occur on the study plot. Lizards are most often caught in the following order: Scelojwnts, Uta, and Etinwces. Two mammals are occasionally caught, with Pero- mtjscus most often seen, but with a shrew seen occasionallv. Only a few snakes have been seen and we ha\e not made a bird census. Thus, by walking tlirough the plot, the most common small vertebrates seen would be lizards.
The size of the lizard population varies not only as to numbers but also as to age group. Thus, if one is to detennine the biomass of SceIoj)onis on the plot he should be specific as to the date because there is a continual growth of juveniles and adults plus the usual adding of hatchlings in late summer which pro- vides for a constant fluctuation of the biomass.
In Tables 12, 13, and 14, the average weight for adults known to be resident on the plot dur- ing 1966 and 1967 is tabulated. It is noted that both males and females averaged heavier in 1967. We account for this primarily because of the differential in the dates of the survey. In 1966 the weights were mostly taken in early June, whereas in 1967 they were taken in late June and early July. The differences are par- ticularly noticeable for the females. This is un- derstandable if we realize that nearly half of them were gravid when weighed in 1967.
Food Habits
The stomachs of 150 lizards were opened and their contents examined. Of those with large quantities of food in the stomachs, the volume
Table 12. Biomass of Sccloporus on Rainier NIesa studv plot for fulv 1966.
Table 14. Bioma.ss of Sceloporus on Rainier Mesa studv plot for August 1967.
Sex
M
Number Avg. in grams Biomass Total biomass
29 31
19.48 19.66
584.40 609.46 1193.86
Table 1.3. Biomass of Sceloporus on Rainier study plot for June 1967.
Sex
Adults M F
Juveniles M F
Number 26 31 19 24
Avg. in grams 19.97 22.09 3.08 2.73
Biomass .519.22 684.79 .58,.52 6.5..5.5
Totals 1204.01 124.07
|
Sex |
Adults M F |
Juveniles Hatch- lings M F |
|
Number Avg. in grams Biomass Totals Total biomass |
26 31 19 24 47 20..56 21.03 8.67 8.18 0.735 534..56 651.93 164.78 196.32 .34..55 1186.49 .361.10 34.55 1582.14 |
Total biomass
1328.08
was measured. We have not attempted to ex- press the food items in terms of percent of volume or in temis of numbers found, but do indicate in general terms the food items ob- served and the volume and weights occurring in what appeared to be filled stomachs.
The major source of food for Scelojwrus on Rainier Mesa is ants. Perhaps this should be expected since there are 2.8 species of ants known to occur there (Cole, 1966). Ants are also extremely abundant as individuals, espe- cially in the organic matter under the pinyons and junipers. Every stomach examined which contained food material contained several ant heads. Some of the fullest stomachs contained only ants. One stomach contained ants with a volume of 1.34 ml, and in this one 68 ant heads were counted.
It appeared in several instances that lizards had found the winged reproductive forms of one of the ant species swanning and had gorged themselves to capacity. There is little doubt but that ants make up a large percent of the diet of S. o. Ion<iij)es on Rainier Mesa. At least we found the selerotized parts of ants (e.g., head, legs, and thorax) to be abundant in the digestive tracts of all lizards examined.
Beetles of several species and in a wide range of sizes are eaten by these lizards. Lady- bird beetles and weevil are the more common ones observed in stomach contents.
Lepidoptera larvae and perhaps some beetle larvae were commonly found. Some of these were relatively large, and, since they are soft- bodied and readily digested, they are undoubt- edly a major food source when available.
Flies are abundant on Rainier Mesa, and several species have been found in the stomach contents. Lizards have been observed to catch flies, but because of their relatively soft bodies as compared to ants and beetles, and since there was often a lapse of time between the lizards' capture and the time they were examined, we are not certain of their identification. Based on the feeding activities as we observed them in the field, we anticipate that flies would con-
28
Bhigh.\m Young U.NivEKsm' Science Bulletin
stitute ;i greater percent than we fonnd in the stomach contents. We have observed lizards to run after Hying insects and to actually jninp for them, and yet, in spite of our seeing the lizards eat flies more fretjuently than any other insect except ants, they have not proved to be a major food item based on stomach contents.
Several species of Hemiptera, both adult and immature, were found in the stomachs. They, like beetles, varied greatly in size. Bugs were recognized in the stomachs at least as often as flies. They are more easilv identified than flics because of the sclerotized portions of the fore- wings and the head and mouthparts which seem to resist digestion.
Orthoptera are common on Rainier Mesa. Camel crickets are commonly found in can- traps, and other crickets and grasshoppers are also seen but are not common. Jerusalem crick- ets are abundant and are occasionally found in the stomachs. The adults with their large, heavily sclerotized heads and powerful mandi- bles are probably not eaten by S. o. lon^ipes. They may be too large and vigorous to be sub- dued and eaten. As previouslv mentioned, im- mature lizards have been killed and eaten by Jerusalem crickets in can-traps .Crickets in general are large prey for S. o. longipes and thev are chewed, shaken, and crushed to a great extent before they are swallowed.
Several other orders of insects undoubtedly are eaten by the lizards in small numbers, among them Neuroptcra, Isoptera, and Homop- tera.
Spiders were found in several stomachs, but.
again, their soft bodies are quickly digested and easilv broken up, making an accurate count difficult. One .scorpion was found in a lizard stomach.
.\s pre\iousiv mentioned (in connection witli home range), Scclopoms occidcntaU.s lono. ipes does not appear to move about foraging for food. More commonly they bask or loaf in the shade and eat whenever an arthropod comes close enough to attract their attention. They appear to eat almost anything of suitable size that moves enough to attract their attention and with almost no discrimination. The fact that they eat tenebrionid beetles and also that a feather and a piece of red plastic ribbon were found in stomachs tends to corroborate the idea. Tliev often ingest small rocks and such items of plant matter as sagebrush leaves {Arte- mviia tridentata) and fragments of needles of the pinvon and juniper trees. A Sceloponts oc- cidental is longipes collected near Milford, Heaver Countv, Utah, was noticeably thin and refused to eat when food was offered. It died and examination showed a large lump in the abdomen. The abdomen was opened and an ingested pinvon needle punctured the intestine, blocking it. In front of the needle was a pile-up of sclerotized insect parts which made the lump easily detectable bv palpation. Opiintia spines and other threatening objects were found in the stomachs of Sceloponts from f-lainier Mesa showing again that the lizards are not, in our opinion, discriminating in their selection and not dexterous in their capture and ingestion of food.
DISCUSSION AND CONCLUSIONS
Few studies of high altitude populations of saurians have been done in western North America. Stebbins (1944) and Stebfnns and Robinson ( 1946) stiidied SceJoporus oraciosus gracilis at 6,000 feet elevation at the Chaos Jumbles area in Lassen National Park. This study revealed certain mixlifications in the basic ecology of this population of Sceloponis graciosus gracilis such as: (a) a larger percent- age of adults in the population; (b) a greater longevity; (c) a reduced number of predators, both as to numbers of species and individuals; (d) a much shorter annual period lor activity; and (e) fewer young.
Most of their comparisons were made with Sceloporus occidenUilis (Fitch 1940), and we
are presently studying Sceloporus graciosus graciosus in central Utah and Sceloporus undu- lattt.s in southeastern Utah, both of which will serve as comparati\e studies.
The present reports on Scelojwrus and Vta from Rainier Mesa have certain facets that are similar to those presented by Stebbins and Robinson (loc cit). In the case of Sccloj)orus occidcntalis, we note the following differences when contrasted with the study of Fitch ( 1940):
I ) A substantial increase in the maximum size of adults. This may be the result of various factors other than altitude. We are dealing in Nevada with a different subspecies and such differenci's as size may represent genetical varia- tions in the gene pool coining from their adapta-
Biological Series, Vol. 15, No. 4 Sceloporus Occidentalis and Uta STAN.'iiiuHiANA
29
tions to a dryer, a.s well as a more elevated, environment.
2) A slightly longer lite span with the ma- jority of adults 3-4 vears old, but with somi' living for six years. In June 1967, 57 percent of the population were breeding adults (22 months or older). This percentage was much higher in 1966 (85.71), owing to a very poor reproduction in 1965. We consider 1967 to be more nearly average but recognize that there will usually be a vearlv fluctuation in all age groups.
3) A shorter season for normal activity. In southern Nevada at 7,500 feet, cool weather limits adults and most juveniles by or before November, and spring rarely comes before 1 April.
4) A larger number of eggs are laid. We have had laid in the laboratory or have observed oviducal clutches totaling 15 eggs and, on the basis of weights before and after egg laying, which exceed the maximum for a clutch of 15, suspect that the larger females lay more.
5) Growth is rapid, particularly during the first full summer or as juveniles. We note that juveniles on Rainier Mesa grow from about 14 mm to an average length of 70-73 mm in 22 months, most (51 percent) occurring from 1 June to mid-August while they are at an age of 9-12 months. Even more remarkable is the gain in weight from approximately 2 to about 9 g in the same length of time. The months of June, July, and August are seemingly ideal weatherwise, and there is an abundance of food —two factors which seemingly contribute heavily to the rapid growth rate.
6) Diet consists largely of ants but with beetles, bugs, flies, and other available arthro- pods included. We do not believe that ants are any more desirable to these lizards than other small anthropods. The fact that they are avail- able in abundance is the important factor. Ants are represented on the mesa by 28 species ( Cole 1966) and have been noted to occur in all nit- ches where the lizards live. Therefore, a lizard may be relatively sedentary, feeding on those food items (ants primarily) as may chance to come nearby.
7) Home ranges of males are relatively large; however, they apparently are not defended in their entirety. Our data suggest that the area really defended is much smaller and consists of that part of the home range used primarily for basking and hiding. Other parts of the home range mav be used for foraging or for other purposes which are not clear to us. Female
home ranges are much smaller and, in contrast to males, may be altered by changing the center of activity. We have not observed any defensive behavior on the part of females.
8) During the warmer parts of the summer, there is a considerable reduction in the amount of time lizards bask. Males climb into trees near rocks or other objects used for basking and may remain in the trees at least overnight and per- haps longer. Lizard activity on the ground is reduced to such an extent that it becomes obvi- ous in the numbers of lizards found in can-traps. Although accurate percentages are not available, we did find a greater percentage of juveniles than adults in the can-traps during the warm periods. Although we did not attempt to record the lengths of time the lizards spent in trees, our observations do indicate that the trees are used mostly from mid-July to early August and that adult catches in the traps gradually increase as the evenings begin to cool from mid-August into September. We did not note any difference between males and females. Both used the trees; however, there was a larger percentage of males in the trees in early July.
An examination of figure 4 indicates that the daily maximums are above the mid-eighties during much of the period in which the lizards are in the trees. Although we did not check care- fully the differential in air temperatures on the surface and in the trees, it Ls seemingly true that temperature regulation would be more easily controlled a few feet above the surface. We have concluded that the true climbing behavior is done primarily for the purpose of temperature regulation. A tree may afford full or part shade or sun as needed, but, more appropriately dur- ing the warmer periods of the year, air circula- tion pennits for a proper temperature balance.
9) On the basis of laboratory hatchlings and those caught in the field, it appears that hatch- lings have an initial or early growth coming largely from yolk remaining in the mid-gut. This increase in s-v length is accompanied by a decrease in weight. We note a difference of approximately 15 percent in the s-v length of laboratory hatchlings (22-26) and have yet to find in the field one less than 26 mm in s-v length.
10) The climatic factors of this desert study area dictate to a great extent much of the eco- logical variaion which we have noted. Popula- tions in this general area rarely have the same weather conditions for two years. We have con- cluded that each year is in some way different, more rain, dry, cool springs, late or early falls, cloudy (reducing radiation and thus affecting
30
Bmr.iiAM Young Univehsity Science Bulletin
growth and/or cinhryonation of the eggs) or a conihination of these aiul otliers.
In general tliis study dein()nstrat<'s that tlie western fence hzard is an adaptable reptile. Because of its basking habits, which pennit it to wami early in the morning even on cool days, this lizard increases its rate oi nietabolisin per- mitting a longer period of activity than would otherwise be possil)le. This allows longer periods when feeding is possible, pennitting an increase in the rate of egg volking, ( thus, an earlier laying of eggs) insuring reproduction. Its utili- zation of available arthropods gives it a food
supply during the season of its activity. The latter, parlicularlv the utilization of ants, is of consideral)le importance in the lizard's ability to siuvive in high altitude habitats.
Because this species occupies many diverse habitats, has a wide geographic distribution, and coTisists of several subspecies, we would hope that future ecological studies might be made in a more northern habitat. Such comparative data would give a more precise understanding of the ecology of this species and perhaps indicate if longitude has an effect similar to what appears to be the case for altitude.
LITERATURE CITED
Allheu, UoHALi) M. .\Ni) D Klden Beck. 1962. Ec()loi;itaI di^tril)lltioIl of mites on lizards at tlie Nfvaila Atomic Test Site. Herpetologica 18:47-51.
Ali.kei), Dohald M., D Eli>en Beck, and Clive D. J()I«;ensen. 1963. Biotic commimitifs of tlie Nevada Test Site. Brigham Young University Science Bulletin, Biol. Series, 2(2):l-.52.
Beatlev, J. C. 196,5. Ecology of the Nevada Test Site 1. Ceograpliie and ecologic distributions of the vascular flora (annotated checkUst). UCLA 12-.553. Lai), of Nuclear Med. & Rad. Biol., Univ. of Calif., Los Angeles. 69 p.
Beatley, J. C. 19{i9. Vascular plants of the Nevada Test Site, Nellis Air Force Range, and Ash Mead- ows ( Northern Mojave and Southern Great Basin Deserts, South-central Nevada). UCLA 12-705. Lah. of Nuclear Med. & Rad. Biol, Univ. of Cilif., Los Angeles. 122 p.
Bell, E. L. 19.54. A preliminarv report on the sub- species of the western fence lizard, Sccloporui ociidcntalh, and its relationships to the eastern fence lizard Sccloporita uiidtilntus. Heqietologica 10:31-46.
Blaui, VV. Frank. 1960. The rustv lizard: a popu- lation study. Austin, Universitv of Texas Press. 178 p. 31 fig.
Boceht, C. M. 1949. Thcrniorcgulation in reptiles, a factor in exolution. Evolution 3( 3 ) : 19.5-21 1.
CANn>, Chahles Lewis. 1916. The subspecies of Sccloporus occidentulis with description of a new form from (he .Sierra Nevada and .systematic notes on oilier (^.ilifornia lizards. Univ. of Cal. Pub. in Zool., 17(7):6.3-74.
Caiu'knteh, C. C. 19.59. .\ popul.itiou of Sccloporus uiitliiliitus connohrinus in sonthcentr.i! Oklahoma. •Southwest. Nat., 4:110-111.
Cahi'enteu, C. C. I9fil. Patterns of social behavior of Merriam s Canyon Lizard {Sccloporua rii. vicr- ri<inii-l<^U(iiii(l(ic). Southwest. Nat.. 6:138-148.
Cole, .'Vhtuuh C, |h. 1966. Ants of the Nevada Test Site. Brigham Young Universitv Science Bul- letin, Biol. .Series, 7(3): 1-27.
CoLENL\N, PlULLU- H. AM) RoilKHT LiVEZEY. 1968.
Infrared rellection of the integument of live S((7- oponi.s occidcntdlis occidciitali.s Baird and Girard from three habitats. Journal of Hen)etology 1(1-4):
71-75.
C<)WLp;s, Kavmoni) B. 1939. Possible implications of reptilian tlHrm:il tolerance. Science 90 (2342): 465-466.
Cunningham, John D. 1966. Additional observations on the bodv temperature of reptiles. Herpetologica 22(3): 184-189.
Davis, John. 1967. (Growth and size of the western fence lizard Scchiponis iiccidcntulis. Copeia 1967 (4):721-731.
Dawson, \V. R. and G. A. BAnriiOLOMEW. 19.56. Re- lation of oxygen consumption to l)odN weight, temperature, and temperature acclimation in liz- ards I'ta stiitishiiritnui and Sccloporus occidenUdis. Physiological Zoology 29:40-51.
Ells, H. A. 19.54. The gross iuid microscopic anat- omy of the liver and gall bladder of the lizard Sccloporus occidcnialis blscrUitus Hallowell.
Ferhel, Cahol M., Howard R. Leach, and Daniel F. TiLLOTsoN. 19.5.'3. F(M)d habits of the covote in California. Calif. Fish and Game, .39(3) :.3dl-.341.
Kiif II, H. S. 1940. A field study of the growth and behavior of the fence lizard. tJniv. Calif. Publ. Zool., 44:151-172.
Fitch. H. S. 1954. Life history and ecology of the fi\e-line skink, Eumeces fasciatus. Univ. Kans. Publ. Mus. Nat. Hist., 8:1-156.
FiTCM, H. S. 19.55. Habits :md adaptations of the CIreat Plains Skink Euinccr\ ohsolctus. Ecol. Monogr., 25:.59-83.
Fitch, H. S. 1956. An ecological studv of tlie col- lari'tl lizaril (Crolaphi/lus colliiris) . Uiii\. K:ms. Publ. Mils. Nat. Hist., 8:213-274.
Fitch, H. S. 1958. Natural history of the six-lined raceriiiiner {Cnemidophorus scxliucutus). Univ. Kansas Publ. Mus. Nat. Hi.st., 11 (2): 11-62.
Fitch, H. S. 1970. Reproductive Cyeles in Lizards and Siiiikes. Univ. Kansas Publ. Nlus. Nat. Hist., .52:1-247.
JOHNSON, Hay (^i.iiroiui. 1965. The dirt of the Pa- ci'ic lence lizard, Sccloporus occidcnl(dis occidcu- Irdis ( B.iird and Giraril) from northern California. Herpetologica, 21(2 ): 1 14-1 17.
I.MisoN. Mkuvin W. lt)61. The critical thrniia] in.ixi- inuni of the lizard Sccloporus occidcutdlis occiden- tulis Baird and CJirard. Herpetologica 17(2): 11.3-122.
Biological Series, Vol. 15. No. 4 Scelopohu.s Occidentalis and Uta Stanshuhiana
31
Leach. H. R. and \V. H. Khazieh. 1953. A shiily of tlie pos.sible extent of predation of heavy con- centrations of \alley (liuiil with .special reference to the bobcat. Calif. Fish and Came 39(4): 527-538.
McCiNNiN, S.\muel M., |h. 1965. Thermal ecology of Scctoporua occidciitalif^. Unpublished Phi). Dis- sertation Univ. of Calif., Berkeley.
Noble, G. K. and H. T. Bradley. 1933. The matinj; behavior of lizards; its bearing on the theory of sexual selection. Ann. N. Y. Acad. Sci., 35:25-100.
Smith, Houaht M. 1939. The Mexican and Central American lizards of the genus Scclo)>orus. Field Museum of Natur;J Historv. Zool. Series, Vol. 26. 397 p.
Stebbins, Robert C. and R. M. Eakin. 1958. The role of tlie "third eye" in reptilian behavior. Am. Mus. Novit. No. 1870:1-40.
Swan, L. W. 1952. Some environmental conditions inlluencing life at liigh altitudes. Ec-ology, 33(1): 109-111.
Talhot, H. E. and R. L. LrvEZEY. 1964. Integii- mental reflectivity of live Sceloponis occideiituluK occidentalis from four liabitats. Herpetologica 19(4):269-274.
Tanner, \V. W. and Clive D. Jorcensen. 1963. Reptiles of the Nevada Test Site. Brigham Young University Sci. Bull., Biol. Series, 3(3): 1-31.
T.\nner, VV. W. 1965. A comparative population study of small vertebrates in the uranium areas of the upper Colorado River Basin of Utah. Brigham Young University Sci. Bull., 7(1):1-31.
Tinkle, Donald VV. 1967. The life and demography of the side-blotched lizard, IHci stdtLshuridnu. Misc. Publ. Mus. of Z<H)1. Univ. of Mich., 1.32:1-177.
Turner, F. B., R. I. Jennrich, and J. D. Weintbaub. 1969. Home ranges ami body size of lizards. Ecology 50 ( 6 ), pp. 1 076- 1 08 1 .
Tubner, F. B., G. a. HoDDENiiAcii, P. A. Medica, and Joseph R. Lannom. 1970. The Demography of tile lizard, Utd stunnhiiriaua Baird and (iirard, in southern Nevada. J. Anim. Ecol. .39, .505-519.
Wu.noKT, D. C. 1958a. Observations of preferred bo(b temperature antl feeding habits of some selected tropical iguanas. Herpetologica 14(3): 161-164. 19.58b. The effect of temperature on tliyroid histology and survival in the lizard Sce- loponis occidentalis. Copeia, 1958:265-276.
WiLHOKT, D. C. AND J. D. Andebson. 1960. Effect of acclimation on the preferred body temperature of the lizard, Sceloponis i>cci(lcnt(di\. Science, 131: 610-611.
WiLHoiT, D. C. AND VV. B. Quay. 1961. Testicular histology and seasonal changes in the lizard S«'- loponts occidentalis. Journal of Morphology, 108 (1):95-106.
WOODIIUBY, M.\RIAN AND AnCUS M. VVooDIlURV. 1945.
Life-history studies of the sagebnish lizard Scelop- onis g. gracio.ftis with special reference to cycles in reproduction. Herpetologica 2(7) : 175-195. ZwEiEEL, Richard G. and Charles H. Lowe. 1966. The ecology of a population of Xantiisia vigilui, the desert night lizard. Am. Mus. Novit. No. 2247:1-57.
NOTES ON THE LIFE HISTORY OF UTA S. STANSBVRIANA BAIRD AND GIRARD
bv
WilnuT W. Tanner
In July of 1965, a study plot was established on Rainier Mesa for the purpose of examining the basic biology of the western fence lizard, SceIo]>oni.s occidentalis Baird and Girard. The area selected is in a loose stand of pinyon-juni- per with Artemisiu and Qtiercus interspersed (Fig. 1). A more complete list of animals is pre- .sented bv Allred et al. ( 196.3) and a list of plants by Allred et al. (1963) and Beatley ( 1965 and 1969).
Associated with Sceloponi.s in this pinyon- juniper community are two other .species, Eti- mecefi s. utahensus and Uta s. staivibxirkina The population density of these species is rela- tively low when contrasted with Uta in the lower valleys of southern Nevada and Eumcccs in areas of California and Utah. In spite of the
lower density and the fact that the major effort at this studv plot was to investigate Sccloporus, some ecological data have been gathered which add to our understanding of Uta. Particularly, we have been able to add infonnation concern- ing longevity and growth. Unfortunately little infonnation was obtained for Exinveces.
The data were gathered by the following graduate student assistants: James M. Hopkin, Ronald L. Morris, Phillip Medica, and Dr. John E. Krogh. The work was performed under con- tract AT (11-1) 1496 bet^veen the U.S. Atomic Energy Commission, Division of Biology and Medicine, and Brigham Young University.
Data pertaining to Vta were gathered for the most part during the summer months of 1965-1967. During the.se years seventy-eight in-
M)ep.Trtiiifnt iif '/.m)loBy. Bnphani Voung Universily, I'rnvn, llLih H4^)()l.
32
BmcnAM YouNO Univejisity Science Bulletin
^.-- ,.,. . . 'i^&^:M
Fig. 1. A loose stand of Pinyon-Juniper with Artcmiiki and Qucrnis interspersed. Study plot Riiinier Mesa. nu .. u. r. . Colyer A. E.G. Photographer.
Photo hv H.
dividuals were marked. In 1968-1970, lizards were ob.served and data gathered, but none wa.s marked. Forty of those marked were recap- tured at least once and some were recaptured several times over a period of four years.
The numbers assigned to the digits are as follows: inner right front, no. I; inner left front, no. 6; outer risiht liind, no. 11; outer left hind, no. 16 (Tanner, 1965). Ages are given to the nearest month. In some instances these dates may be off as much as a month because we cannot detennine the exact date of hatching; however, these slight discrepancies do not alter the age significantly. Table 1 presents data which indicate size and longevitv in this popu- lation.
In 1965 we marked only 2.3 I'tci. Tiiis reflects our late start in marking lizards at this plot during the initial vear ( 18 Julv 1965). However, of this group, seven were recaptured and at- tained at least 50 mm (av. 52.2) in snout to vent length ;md reached at least three years
of age. We have few records of Uta less than 24 months old attaining a snout to vent length of .50 mm. The extremes of size range from 21 mm in small hatchlings to 56 mm in old adults. Based on marked lizards, three age groups can be recognized in April and May. The first year or last vear's hatchlings are 35-40 mm, the second vear, 20-22 months old, are usuallv 42-47 mm but with a few 48-50 mm, and the older adults, nearing three years, 50-55 mm. Tliere may be more overlapping of size in some years than in others. Size in the spring reflects the growth pennitted and attained by hatchlings during the late summer and fall months. By mid- June the larger juveniles mav overlap the small- er indi\iduals of the second age group, and the larger individuals now t^vo years old are indis- tinguishable from the older lizards. During July, .\ugust, and September, only two age groups are casiK' recognized: hatchlings jirst emerging and adults. Some of the vounger adults, one or two vears old, are still onlv 45-49 mm in snout to
Bioi.ociCAi, Sehies, Vol. 15. No. 4 ScEi.oponu.s Occidentai.is .vnd Uta Stansuuhiana
33
vent longth, and \vv suspect tliat few reach 50 mm until thev are past two years of age, that is, by August or September of the second sum- mer ( Table 1 ) .
Records for marked two-vear-old Uta on Rainier Mesa indicate that they have a snout to vent length of 46-52 mm with those less than 50 mm being the exception rather than the rule. We suspect that some of the adults caught in August are of this age group. The following examples indicate that some individuals do reach 50 mm bv the time thev are 13 months old: No. 3. F. 28 Julv 49 mm; 25 August 52 mm; No. 4, M, 23 July 4S mm; 29 August 51 mm; No. 5, M, 30 July 45 mm; 15 August 47 mm. We believe, however, that the majoritv of liz- ards reaching .50 mm or more are in their second year, that is. they are 13 plus months of age. An apparentlv rapid growth by a few young indi\iduals and a longer life span has made this population unicjue when compared with Uta populations in the lower valleys. Our records of age groupings seem to justify the categoriz- ing of lizards which are 50 mm or more in s-v length as old adults, that is, two or more years old.
A more precise record of size and growth during September and October is needed. We suspect that all age groups grow during these fall months, albeit the older ones at a slower rate. We observe hatchlings from late August to earlv October to be 28-40 mm, and by early June of the next year they have grown to 40-44 mm. Six male hatchlings were marked in late July and early August and recaptured in late .'August or early September. Table 2 indicates the growth in these individuals.
Altliough there is variation in the rate of growth, the data for 1966 and 1967 do indicate a relatively similar rate for those which show growth. Where there is some growth, such in- dividuals may represent those with a chance for survival. If we consider the natural vicissi- tudes under which most hatchlings emerge per- haps the problem of food getting is a major problem, at least in the low valleys where it is hot and dry. However, the fact that some may survive for 20 or more days (1-19, Table 2) without growth mav also suggest that even with- in the first few weeks those that will not survive the first winter are indicated. Furthermore, many hatchlings do not survive more than a few weeks, making it difficult to mark more than part of the hatchlings before they die. The heavy mortality of hatchlings may be a function of feeding, cold weather, predation, a late hatch- in<' date, or a combination of these and other factors. In 1965 this plot was closed on 2 Sep- tember. Although we were most careful in our observations for hatchlings, we did not see one; yet, in early June of 1966, we marked five Uta that were in the size range of hatchlings from the preceding summer. Even though a few were marked the next spring, they represented a small proportion of a normal hatch as we observed it in 1966 and 1967. In 1966 we marked 33 hatchling Uta on Rainier Mesa. Only seven (21.2 percent) were recaptured in 1967. However, nine juveniles were marked in June 1967; that is, their size (42-46 s-v) corresponded to those marked in 1966 and recaptured in 1967. A de- crease in the number of surviving hatchlings does effect the percent of juveniles and old adults in a population. In June of 1966 we had
Table 1. Data for Uta recaptured in tivo or more years (1965-1968) at Rainier Mesa study plot.
|
Toe |
Date |
Snout to |
Weiglit |
Times |
.\g(' in |
Maximum |
Weight |
Last date |
|
|
No. |
marked |
vent mm |
in grams |
recaptured |
months |
Se,\ |
size |
in grams |
captured |
|
4 |
28-7-65 |
46 |
4.0 |
3 |
35 |
M |
51 |
3.50 |
24-6-67 |
|
6 |
5-8-65 |
50 |
3.0 |
8 |
48 |
F |
52 |
3.98 |
10-8-68 |
|
7 |
5-8-65 |
44 |
3.0 |
1 |
36 |
F |
50 |
5,10 |
6-8-67 |
|
10 |
7-8-65 |
.54 |
5.0 |
2 |
.37 |
F |
.54 |
3.94 |
23-8-66 |
|
12 |
9-8-65 |
47 |
2.0 |
12 |
36 |
M |
52 |
4.24 |
12-8-67 |
|
13 |
11-8-65 |
44 |
3.0 |
3 |
48 |
F |
51 |
3.69 |
1-8-68 |
|
16 |
1,5-8-65 |
50 |
4.0 |
1 |
35 |
M |
.52 |
4.40 |
6-6-67 |
|
1-13 |
23-7-66 |
50 |
3.0 |
2 |
35 |
F |
51 |
4.41 |
3-6-67 |
|
2-5 |
11-8-66 |
30 |
.62 |
1 |
13 |
F |
49 |
3.32 |
28-8-67 |
|
2-7 |
1.3-8-66 |
24 |
.32 |
1 |
11 |
M |
45 |
2.96 |
3-6-67 |
|
2-14 |
2.5-8-66 |
31 |
.80 |
1 |
11 |
F |
43 |
2.62 |
18-6-67 |
|
2-18 |
29-8-66 |
23 |
.45 |
3 |
23 |
F |
49 |
4.35 |
3-6-68 |
|
3-9-10 |
4-9-66 |
26 |
..50 |
3 |
12 |
M |
48 |
3.47 |
26-8-67 |
|
3-11 |
4-9-66 |
31 |
.60 |
3 |
11 |
]•■ |
46 |
2.80 |
10-7-67 |
|
3-19 |
.5-6-67 |
.50 |
2.70 |
3 |
36 |
M |
51 |
4.17 |
4-8-68 |
|
4-5 |
.5-6-67 |
42 |
2.20 |
5 |
.36 |
M |
52 |
,5.11 |
8-7-69 |
34
Brigham Young University Science Bulletin
Table 2. Observed jjrowlli in Uta hatchlings Rainier Mesa, 1966.
|
Toe |
First |
S-V |
Last |
S-V |
Time |
Growth |
|
No. |
capture |
length |
capture |
length |
in days |
m mm |
|
1-11 |
17-7-66 |
2.3 |
4-9-66 |
32 |
49 |
9.0 |
|
1-14 |
27-7-66 |
24 |
4-9-66 |
34 |
38 |
10.0 |
|
1-16 |
9-8-66 |
26 |
23-8-66 |
29 |
14 |
3.0 |
|
1-18 |
9-9-66 |
26 |
29-8-66 |
29 |
20 |
3.0 |
|
1-19 |
9-8-66 |
29 |
29-8-66 |
29 |
20 |
0.0 |
|
2-6 |
1.3-8-66 |
29 |
29-8-66 |
32 |
16 |
3.0 |
ill tliis population approximately 65 percent old adult.s, whereas in 1965 and 1967 our .sample indicated only 3S and 36 percent, respectively.
We interpret these percentages to mean that this population has the ability to respond to a low reproduction by carrying over a larger per- centage of the older adults. It is also noteworthy that the percentage of older lizards dropped considerably in 1967, as a result perhaps of greater competition from the increased number of juveniles and because of the imbalance of old individuals in the population.
During the months of September and Octo- ber, weather conditions progressively limit lizard activity. By October, daily ma.ximums have dropped into the 70 or 60 degree range and the nights are cold with average daily mini- mums for October ranging from 32-47 degrees for the years 1965-1970. For October 1965-1970 the average daily temperature ranged from 51- 66° F. On 9-10 October 1970 maximums at 3:00 p.m. were 60^ and 58° F. At these temperatures, only three Uta (two hatchlings and one adult) were seen during tour hours of observation (2:00-4:00 p.m.).'
Data on the rate of growth indicate that most hatchlings continue to grow as long as weather conditions permit them to be active. On Rainier Mesa during the fall months, only the approximate amount of daily activity can be determined. Each year is apparently differ- ent, and growth rates are reflected by such con- ditions as ( 1 ) a late spring in which full activi- ty is not possible until mid or late April. An example of a late spring is 1969 in which a foot of snow covered the stuth- plot on 5 April. On S April 193S, the temperature was 45"F at 12:. 30 p.m.; however, two lizards were seen. We have noted that there is as much as two weeks' dif- ference in the time of egg-laying which is pri- marily because of a late or early spring. (2) Summers are at times cool on the mesa. In 1965 and 1969 cloudy days and rain kept tem- peratures down and delaved egg-laving anil particularlv hatciiing in 1965. Should either of
these weather variations occur, hatchlings ap- pear late in the summer and a reduced period for growth occurs before hibernation. Wc have not fxperienced a late spring followed by an early fall nor have we seen an early fall followed bv a late spring. Here as in most desert areas, particularly in steppe deserts, there is consid- erable variation not only in amounts of rainfall but also in temperature fluctuations. (See fig- ure depicting weather data in the preceding study of Sceloporus.)
The largest hatchlings measured in late Sep- tember and up to mid-October ranged between 37-40 mm in s-v lengths. Therefore in favorable vears when considerable activity occurs in Sep- tember and October, utas may achieve a sub- adult size and thus be sexually mature at an earlier date the next spring.
Shortly after setting up the study plots at NTS we noted that larger adults and a larger percentage of large adults were present in the Rainier Me.sa Uta population (Fig. 2). The fact that three populations separated by only 20 or .30 miles occur in different valleys and show a variation in the average size of individuals seemed to warrant additional study. We are aware that size variation in populations is not uniijue in utas (Tinkle, 1967; Pack and Tanner, I97()); howcNcr, these populations are in dif- ferent types of desert habitat and at different elevations. At the outset, we can recognize that the valley populations (Mercury and Yucca) are more ne;irlv the s;mie average size than either is to the Rainier utas. Furthennore, the valley populations have a longer period for ac- tivity (April through November) than those on the mesa. According to Tanner and Jorgen- sen (1963), some activity was noted in Yucca Flat during the wanner days of all months. This is not the case on Rainier Mesa where there is little or no activitv from late October to e;irlv .^pril. Howe\er, little difference in growth of hatchlings is noted in the populations and, therefore, we suspect that other factors are more important than the length of the period of si'asonal ;icti\it\'. Such factors as ;i\ailabilitv of food during most or all ot the seasonal ;ictivi- ty period, higher humidity, lack of competition (only two other species of lizards), and lack of prediition may mean that the mesa popula- tion, when active, grows faster, and our data suggest that a larger percentage, when active, grows faster, and that a larger percentage of the population lives longer.
riic (tiaphic factors seemingly contribute to a greatiT lood supph'. More rain in the summers |irodu((s a grciitcr plant cover which in turn
HioLociCAL Series, Vol. 15. \o. 4 Scelopohu.s Occihent.m.is .\ni) Uta Stanshuhianji
35
S-V Length in mm.
40 50 60
No. W t. in gr .
|
Mercury Va 1 1 e y 3,367 ft. elev. |
A B |
44.76 52.00 |
111 |
2.85 4.22 |
|||
|
^a^mmmm i |
|||||||
|
Yucca Flats 3.950 ft. elev. |
45.70 52.00 |
168 |
2.98 4.10 |
||||
|
A B |
|||||||
|
Rainer Mesa 7,480 ft. elev. |
48.44 56.00 |
45 |
4.49 5.78 |
||||
|
A B |
|||||||
Fig. 2. A bar graph .showing; the average ;uid niiUiimum ^ize of three popiilatiDiis of Vtu stiin.sliurUnid on the Nevada Test Site. Maximum size is based on adult m.iles.
supports more insects. In early September of 1967, Dr. Vasco M. Tanner and I spent one week collecting insects in an attempt to supple- ment infonnation published previously under contract AT (11-1) 786 (Allred and Beck, 1964; Tanner, 1965). Our efforts were centered in Frenchman's Flat and on Rainier Mesa. There was still enough residual moisture on the Mesa to keep the late summer shrubs and hebaceous plants green, and in the open areas between the pinyons some plants were still in bloom. In contrast the flats were dry with the plants in a state of dormancy. Under these conditions, we found far more insects, of all kinds and particularly small beetles and leaf hoppers, on the mesa. Considering the availability of food at this important season when the hatchlings are active, one is inclined to place great weight on the availabilit)' of food as a factor in growth rate and size.
There is also a higher average daily humidity on the mesa than at the low valley flats. We have reasoned that tliis may affect the lizards in such biotic communities by reducing the evaporation and thus permitting the habitat and its biota as well as the lizards to be active. In the low desert flats, the heat is intense with soil surface temperatures up to 140'^F at mid- day; soil moisture is ver)' low and relative hu- midity may fall to a level of less than 10 percent
during the heat of the day. Physical conditions permit lizard activity on the mesa during all or most of the day, but often prevent full activity in the low valleys. From the first of June until at least Tiiid-September, daytime maximum tem- peratures on the mesa range from 65 to 8.5° F. This permits lizards to be active throughout the day. In contrast, few lizards are active in Frenchman's or Yucca Flat from noon until .5:00 p.m. and often during July and August until much later.
We have therefore reasoned that the mesa utas are able to be active for nearly as many hours during four months as the low valley utas are in si,\ months. Our reasoning is based on the fact that the valley utas are active during most of June, July, and August, only during the cooler morning and evening hours; whereas the mesa population is active from soon after sunup to sundown and with an abundance of food. If days of activity and not hours determine growth and thus size, then the valley utas should grow faster and perhaps become larger (in snout to vent length ) than those on the mesa. This, however, is not the case. Hatchlings marked in August are about the same size in June in all three study plots. We are inclined to assume that the hours of full activity for mesa utas must be considered and not the number of days or months. Particularly during July and
36
Bbk;ham Young UNivEHsin' Science Bulletin
August, mesa utas have more time (6-10 hours per dav) than do utas in the low valley which have fewer hours of full ac-tivity per day during the summer months wlieii residual heat keeps the surface soil hot during most of the sunlight hours.
The entire prohlem ot growth and size is seemingly one of mctaholism in which the amount of food available and the length of time at which individuals can be at or near full activity in getting food play an important role in the populations under consideration. We be- lieve that tlie larger si/e of individuals in the population and the apparentiv more rapid and sustained growth of the mesa utas is the result of an al)nndance of food and a greater number of hours during the summer when full activity is possible.
In contrast, the utas in thi' low valleys are not active or have a greativ reduced activity during the hot middays of most of June, July, August, and part of September. Furthennore, the attendant heat reduces the available soil moisture and humiditv which in tuni affects the plants, slowing their growth and forcing them into donuancy. This reduces the available food, based on our sweeping samples. Such untoward conditions become intense in late August and September and are undoubtedly responsible for much of the late summer mor- tality in hatchlings, at least in the lower vallevs. Table 3 shows the data for hatchlings marked in 1985 and 1966 at the Mercury study plot.
We .saw no hatchlings in 1965 on Rainier Mesa. In Table 4 the data for hatchlings marked in 1966 are reported.
The data summarized in Tables 3 and 4 in- dicate a few discrepancies when compared with our discussion above. First, there appears to be a greater daily growth for hatchling utas at Mercury Valley than on l^ainier Mesa. This in- Table 3. Data for hatchling.s marked in 196.5 and 1966 on the Mercury Valley study plot.
Average Rate of Percent
Numl)er.sizc(inm) growth(g) recaptured
Marked 1965
(July 8-.Sep. 6) 201 2.5.18
Day 0.196 Kecapturcd 1965 38 28.19 Week 1..372 18.91
Recaptured 1966 26 44.59 12.93
M.irk.cl 1966
(June2,5-Aug. 31) 185 25.62
Day 0.202 Recaptured 1966 .37 28.32 Week 1.414 20.00
Recaptured 1967 17 43.0 9.19
T.ihle 4. Data for liatcliliugs marked in 1966 on the Rainier Mesa .study plot.
Average Rate of Percent Number size (mm) growth(g) recaptured
Marked 1966 (July 17-Sop. 4)
33
Recaptured 1966 8
Recaptured 1967 5
25.51
Dav 0.178 31.00 Week 1.250 24.24 44.40 15.15
consistencv is understandable if we note that hatchlings appear appro.ximately three weeks earlier there than on the mesa. Thus there is included in these data a longer period for growth. Our conclusion that there is a more rapid growth on the mesa is based in part, then, on the facts that the average size for hatchlings in these plots is about e(jual to the mesa hatch- lings, in spite of the fact that they hatch later ( first hatchlings 1966-Mercury Valley, 25 June; Rainier Mesa, 17 Julv) and have fewer days included in the data. The difference in average size for those recaptured the next spring or summer (0.024 mm) can be explained onlv on the basis that the l^ainier Mesa population is rarely active from November 1 to April 1, whereas utas in the low valleys are at times active during each of these months and pre- sumablv continue to grow but at a reduced rate.
Our data were not gathered to determine preciselv tlu' size for one-vear-old lizards; how- ever, the information available dm's indicate that indi\iduals of the three populations are approximately of equal size at one year in the longevitv cycle. Rainier Mesa utas average larger after the first year.
The question of genetic differences in these populations has not been explored, nor have we extended our efforts into a studv of meristic variables. However, there are obvious reasons, size tor example, to recognize that these popula- tions may be establishing some gene pool dif- ferences, unless we reason that all differences thus far noted are the result of more and per- haps more readilv available food during a criti- cal time in the life cycle.
In 1963 Tanner and |orgensen. using the data obtained at the 'V'ucca Flat study plot, esti- mated the density of utas to be 3.3 adults and 5.0 juveniles per acre in June (15-.30). In 1967 we emplo\ed a teneing techni(|ue in which one- (|uarter-acre plots were fenced with hardware cloth. This was done just before dark after lizard activitv had ceased. We set up three fenced plots (over a period of 10 davs ) and
Biological Sehies, Vol. 15. No. 4 Soelopouu.s Occident.'^li.s and Uta Stansiiukiana
.57
caught all lizard.s found in.sidf the enclo.sure. The fenced areas were .selected at random about 200 yards from the study plot. The fence was four feet high with the bottom buried in several inches of soil. Tal)le 5 gives a summary of our results with reference to Uta. Data for other species will appear in our report on the Frenchman's Flat study plot. We tried the fencing techni(|ue in Mercury \'alley in mid- July hoping to obtain exact data on hatching and hatchlings. This did not succeed because the mesh of the wire was too large to hold the small hatchlings. However, tor adults of all species of lizards on NTS we believe that the small fenced plot will give rather reliable den- sity intomiation in a matter of a few days.
T;ible 5. Vtti density in Frenchman ,s Flat as deter- mined bv three 1/4-acre fenced study plots ( 10-20 June 1967).
|
Est. no. Species Nimiher Adults Jineniles per acre |
Est. biomass |
|
Uta stansburiana 8 8 0 10.40 |
23.09 g |
Population density for Vta on Rainier Mesa has not been estimated for June as in Yucca (1962) and Frenchman's Flat (1967). However, on the basis of capture-recapture data for 1965, it is indicated to be approximately 4.5 lizards per acre (25 uta on 6.2 acres or a biomass of approximately 20.20 per acre). Data for 1966 and 1967 reflect a slie;ht reduction. This we expected and explain on the basis of a low re-
production on the mesa for 1965. An average for four years based on capture-recapture data (1965, 1966, 1967, and 1968) sets the biomass per acre at 19.98 grams. The one discrepancy is that the 1968 survey is ba.sed on an August rather than on a June-early July weight. This may influence the weights to be slightly higher but should not be any more disruptive than the yearly variations.
Thus far we have made only limited com- parisons between populations of utas on the test site. These, however, have indicated a dif- ference in snout to vent size, an adjustiuent to high altitudes to pennit a more rapid growth rate, variation in longevity, and a variation in population density. We may expect other eco- logical variations to occur and assume that a more detailed study of each study plot com- munity will offer additional reasons for some of the differences thus far noted in the popula- tions of utas.
We expect the data from our Mercury Valley plot to be similar to that being presented by Turner (1970) for the Rock Valley plot, since they are at approximately the same elevation and only three miles apart. A comparison of data gathered from the several populations of utas occurring in the low valleys of NTS and that presented for other populations should be made with a high altitude population such as that on Rainier' Mesa (7,.50O feet). The data now available on the ecology of the genus Vta should pennit an analysis of the species stans- buriana, at least of its three subspecies, stans- buriana. s^/iK'gt'r;, and uniforniis.
DISCUSSION AND CONCLUSIONS
An extensive population study of utas at an altitude above 4,.500 feet has not been done be- fore. Therefore, all comparisons refer to uta populations in habitats with a different set of physical and perhaps biotic factors. A study by Stebbins (1944) and Stebbins and Robinson (1946) considered Sceloponis graciostis graciUs in the Chaos Jumbles habitat at 6.000 feet in Lassen National Park, California. Some of their conclusions are similar to those to be made for utas on Rainier Mesa, namely that:
1) Utas live longer at higher altitudes, and thus there is a different population structure than that reported for populations at lower elevations.
Rainier Mesa is somewhat uiiinue since it is
a high but rather narrow isolated uplift in an otherwise relatively dry desert. From Yucca Flat to the east, the mesa rises abniptly from about 5,000 to its summit at about 8,000 feet. We recognize that a more massive mountain or plateau area may not have a population of utas at 7,500 feet. In spite of these unusual physical conditions, we believe that this high altitude population does offer important clues to popu- lation structure and altitude distribution in Uta. In previous studies. Tinkle (1967) found that in Texas adult utas 19 or more months of age made up only approximately 5-10 percent of the population ( percentage will vaiy from year to year in all populations). Tinkle (1967) and Pack and Tanner ( 1970) found the percent-
3H
Bhigiiam Young University Science Bulletin
age in the upper Colorado Basin of Colorado and Utah to be 20-25 percent, and Turner et al. (1970) indicated approxiinatilv 25 percent (1966-30.4%, 1967-17.4%, 196S-29..3'r ) for Rock Valley in southern Nevada. Tanner and Jorgen- .sen (1965) indicate about 50 percent for old adult.s in Yucca Flat. In this study the per- centages range from 36 to 65 percent. Fifty percent for Yucca Flat seems high. By noting the annual variations in the population studied, we must recogni/e that one cannot just drop in on a population for a year and expect to get a true picture of its population structure for more than that one year. The studies cited above show clearlv that vear to year variation in population structure occurs. The Yucca Flat population may, in a given year, have a large carry-over; however, it is not likely that several vears would indicate an average of much above 30 percent. The Texas studv areas were at 2,900 feet; those from Colorado, Utah, and southern Nevada (Rock X'allev) are between 4,000 and 4,.500 feet; Yucca Flat is at about 4,6(X) to 4,800 feet and Rainier Mesa study plot at 7,480 feet. It is of some interest to note that as the eleva- tion increases so also does the percentage of older adults in the population. On the basis of the studii's thus far reported, it would appear that in Uta altitude has a verv significant effect on the life span and that it affects the popula- tion structure to such an extent that in some (Texas) the older adults are so limited in num- bers as to contribute little to the annual repro- duction of the population (Tinkle, 1967). How- ever, in the other population studies done at moderate to high elevations, the 20-25 and .36-65 percentages of older adults do contribute to the annual reproduction and are an important factor in the population structure.
It is not clear to us just whv altitude may elfeet longevity. It has been suggested that the following may he factors which influence it:
a) A possible reduction in the yearly ;ictivity period.
b) A shorter period of intense heat during the activity season or perhaps a longer period of optimum temperatures.
c) Less competition from other utas (less dense population) and Irorn other species.
d) An inherent genetical difference which has resulted from adaptation to the varied habitats.
These factors and others with various com- liinations in;iv aid in answering the (piestion; however, it would seem pertinent also to investi-
gate a population on the northern edge of the species tlislribution to ascertain if there is a longitudinal as well as an altituilinal increase in longevity.
2) Utas attain a larger size than those oc- curring in the populations of the adj;ieent lower valleys.
This generalization mav not be tnie in all sections of the wide distribution of the genus Utii. We note that the Uta of western Colorado and southeastern Utah are noticeably smaller than those from western Texas, in spite of the increase in altitude. It has been suggested that size is dependent on the availability of food. This may be a factor; however, it is our belief that variation in size is associated with local groups and may be related to populations with- in subspecific groups.
A speculation based on the above observa- tions suggests that the average size of adults in a population such as on Rainier Mesa may be used to detennine the approximate percent- age of adults in the population. Variations in the yearly cycle should be considered and the survey made at a given time such as during the first week of egg-laying or a week after the first oviducal eggs appear. If such percentages can be derived a biomass can seemingly be readily determined.
3) With a larger average size for adults in the Rainier Mesa population there are more eggs per clutch. Turner et al. ( 1970) also found this to be generally true in Rock Valley, and it ap- parently is a fact to be considered in the esti- mation of lizard fecunditv. Our data do not pro- vide information on the numbers of clutches laid, however, we do find the increase in the number of eggs per clutch to appear significant. Twelve clutches laid by females taken at the Mercurv \'alley plot produi'cd clutches ranging from 2(3.34)4 eggs per clutch (females ranged from 42-47 mm s-v), whereas five clutches from Rainier Mesa ranged from 4(4.85)6 eggs per clutch (females range from 48-51 mm, s-v). Turner ( loc. cit. ) shows only averages ( Table 7), all of which are less than four eggs per clutch.
It would appear that in our eagerness to understand the basic ecology of this species we have not taken into account sufficiently the fact that ecological variations in habitat intro- duce new parameters into the basic ecology of popukitions. Therefore, we should not expect utas Ix'longing to different subspecies and locat- ed in vastly different habitats such as eastern I'tah, southern Nevada, or western Texas, to
BiouiciCAL Series, Vol. 15. No. 4 Scklopoiu'.s OcrinENTALis and Uta Stansburiana
39
behave the same ecologically anvinore than we should expect them to exhibit the same morpho- logical characteristics. An analysis of data thus far presented by man\' authors ( Jorgensen and Tanner, 1963; tanner, 1965; McKinney, 1971; Tinkle, 1961; Tinkle, McGregor, and Dana, 1962; and Turner, 1970) and the data ayailable for this stud\' suggest yer\' strongly that it is not desirable to consider in detail the demography of Uta staiisburUina unless one subdivides this species into its various subspecies. Behavorial,
ecological, and morphological characters appear to have produced basic variations in the biology of subspecies, some of which provide for varia- tions even within the populations of a subspe- cies. Perhaps, therefore, if in future studies sub- species variations are recognized, such factors as size and longevity would be considered as variables based on adaptation to habitat factors. If subspecies and or populations are to be con- sidered as incipient species we would expect variations to occur.
LITERATURE CITED
Allhei). D. M., D E. Beck, and C. D. Jorgensen. 1963. Biofic- Communities of the Nevada Test Site. Brigham Young Uni\ . Sci. Bull., Biol. Ser., 2(2): 1-52.
Allred, D. M., and D E. Beck. 1964. Arthropod associates of phuits at tlje Nevada Test Site. Brig- h;im Young Univ. Sci. Bull., Biol. Ser., 5(2): 1-16.
Beatley, /. C. 1969. Vascular plants of the Nevada Test Site, Nellis Air Force Range, and Ash Mead- ows. UCLA 12-705. Lab. Nuclear Medicine and Radiation Biolog\', L"niv. Calif., Los Angeles. 122 p.
. 1971. Ecologic and Geographic distributions
of Ihi- \ascular plants of southern Nve Countv. and adjacent parts of Clark. Lincoln, and Esmer- alda Counties, Nevada. (Supplement to UCLA 12-705.) Lab. Nuclear Medicine and Radiation Biologv, Univ. Calif., Los Angeles. 49 p.
Jorgensen, C. D., and \V. W. Tanner. 196.3. The application of density probability function to de- termine the home range of Vta stan.sljuruina .sffliw- hurUina and Cncmidophorus tigrk tigrii. Herpeto- gica 19(2): 105-15.
McKinney, C. O. 1971. Individual and intrapopula- tional \ariations in the push-up displav of Vta sttiml>uruitui. Copeia 1971( 1 ): 159-60.
Pack, L. C, Jr. and W. W. Tanner. 1970. A taxo- nomic comparison of Uta stansburiana of the Great Basin and the Upper Colorado River Basin in Utah with a description of a new subspecies. Great Basin Nat. 30(2):71-90.
Tanner, W. W., and C. D. Jorgensen. 1963. Rep- tiles of the Nevada Test Site. Brigham Young Univ. Sci. Bull, Biol. Ser., 3(3):1-31.
Tanner, W. \V. 1965. A comparative population studv of small vertebrates in the Uranium areas of the Upper Colorado River Basin of Ut:di. Brig- liim Young Univ. Sci. Bull., Biol. Ser., 7(1):1-31.
Tinkle, D. W. 1961. Population stmcture and re- production in tlie lizard Vta stanxhtiriuna stcjnegeri. American Midland Nat., 66( 1 ) :206-2.34.
. 1967. The life and demographv of the side- blotched lizard, Vta stansburiana. Mus. Zool., LIniv. Mich., Misc. Publ. 132:1-182.
Tinkle, D. W'., Don McGregor, and Sumner Dana. 1962. Home range ecology of Vta stansburiana
stcjnegeri. Ecology, 43(2) :223-29.
Turner. F^. B., G. A. Hoddenbach, P. A. Medica, and |. R. Lannoin, Jr. 1970. The demographv of the lizard, Vta .stansburiana Baird and Girard, in Southern Nevada. J. Anim. Eco. 39, 505-519, June 1970.
645
<=ncme
Bookbinding Co., Inc.
100 Cambridge St. Charleston/n. MA n?i?q
3 2044 072 224 645
I-^#.I
4 t
r'l.* «
|
i.i 1«* |
-r |
1 1 ^ ^ |
|||
€ t
4K m
M.^*-
m .A .
»•' * * * » "' ^ • J 11 ? il « it
4 M
*...#
.1 ;^;i;;f 5^' • ■ ■ - ■ _.l ^^
. -.. ■^. :^. '^^^ ^- t -'■■
•^ fit.
i i
, t f * «, -
i'i^
%. ■•*&
. * 'i-
-« •■•.«. A
«'^"j 'i& # «
|
..^.-ff S-, <^-. iJF |
|
|
,t ^- :* » |
|
|
..♦^**^.*^*^» |
|
|
■'.- ^^ # ^■• |
|
|
^' #*^^ ■ |
|
|
»^^-' # |
|
|
. li%%*J»-»*»*,''9* |
|
|
" il; '' |
^* •* |
|
■^A*fi. |
* f- |
|
^c.* ^,*, »• ii i' ■ |
|
|
^#»>^<*;.. |
|
|
'.> A Af .. |
^^^^*,*,*AJ'
f »
h^-n
|
:*? ' > |
r |
|
|
%%^' |
1 1 |
t |
|
,^r |
?• '*^ |
''S |
|
1 |
.^*^ |
|
|
# t |
f * |
yr^
,^^i^..A
.^'f*^:^-^^^^^^^^*,!^
^^^.^.i^lii
|
>-^ |
|
|
■ ■■ |
|
|
^^^ ^^^ ^^^ |
|
|
t |
■ ■■■ |
|
"i'\ |
|
|
KX\ |
■ ■■■ |
|
IBHB |
|
|
1 1^ |
|
|
3 |
■ ■■■ |
^^KKK'^^k.k.K%
,^^j^mK^^^^
^ W^'^it^^'^^r
. *i #1 *■ •
^■. IlI:*"
5*^^"
■*'i
Wi
jXi'^
.»:>:•
I |-|fc
iTf^
»^^.*.*^.ri;
r $ $ k
% $' t h k
y,>t^^^s^iH<'XK<<<':^:>ym^m.
* .* %:$:
M m
1:1 -1-^1:0:%%
f.«: I
X. trS 'C\ ,ti>. * -ij? ^- ..'"■, W.-
& - ■ ' ■ ■ ■■
^^.^i^:...*!. m. « t. ^ .«; "^ 'i^"v.
'• :^ ^i£. ,iJ^ '# -■# .((» .# .^ji '.