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Ectoparasitic mites (Acarina) from small

mammals

in southern

Sweden

By Aunrns Eor,nn

Department of Animal Dcologv, l.lniversity of Lund

Contents

・・・・・・・・・・・・・・・・・・・・・・・・・・・・・・・・・・・・・・・・・・・・・・・・・・・・・・ 100 100

・・・・・・・・・・・・・・・・・・・・・・・・・・・・・ 106

l.

Introduetion

In 1963-1969 small mammals were collected at five places in Scania, the southernmost province of Sweden (Fig. 1). Two localities are situated on the coast, the others inland. The aim lvas to study seasonal fluctuations in numbers of rodents and their ectoparasites. This paper lists the mites col- lected.

2.

Material and methods

A total of 2,507 small mammals, r'epresenting ten species (Tab. 1), rvas collected at all seasons. 1,206 specimens, representing seven species, had 8,582 mites, an in- festation frequency of 48.1 o/0.

The mammals *'ere collected in live net cage traps (Edler 1971). The ectopara- sites rr-ere removed by tu'eezers and preserved in 80 0/o ethanol until mounting on slides.

3. The

ectoparasitic mites

The mite species are listed below, arranged according to Bregetova et al.

(1955) and Bregetova (1956), but in a few cases the names suggested by Evans and Till (1966) are used.

8,582 mites were recorded. 7,334 of them belong to the superfamily Gama- soidea Reuter (order Parasitiformes)

.

The others (l[gobia, Pggmephorus

Entomol. Ts. .lrs. 93. H, 1-3, 1972

Fool

100

108 108 10S

(2)

ECTopARAsrrrc r\{rrEs (lc,lnrN,L) FRor\r sMALL MAMMALS 101

Fig. 1. Southern Sweden

with the collecting locali-

ties. B:Barsebdck, H:

Hiillestad, K:Kullaberg, L:Lund and S:Stam- pen.

and .Neofro mbiculrr), all of them suborder Trombidiformes Reuter, are Acari- formes.

29 species were represented, 5 of them new to Sweden. There were 1,249 larvae, 100 protonymphs, 336 deutonymphs, 607 males and 6,290 females.

99.7 0lo of the 1,248 specimens of Acariformes were larvae (Tab. 2 and 3).

Order Acarilormes Reuter

Suborder Trombid,iforrnes Reuter, f9O9 Family Pyemotid,ae Oudemans, 1937 Pggmephorus sp.

Three specimens found on S. arnneus.

It has been recorded by several authors from T'alpu europeea L. and Apo- dernus spp. in Europe, but these are probably temporary hosts (Zumpt 1961).

Family Trombiculid,ae Ewing, 1929

lieotrombiculcr zcrchuatA'ini Schluger, 1948 (:Trombicula zuchuriftini Schlu- ger 1948).

1,244 larvae found on S. oraneus, C. glarcolus, trI . ugrestis and A. sgluoticus.

The larvae of this mite are parasites both on marnmals and man. They are known from many species of small mammals in Europe (Kepka 1gG4).

Entomol. Ts. Ars. 93. H. 1 -3, 1972

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102 ANDERS EDLER

Family Myobiid.ae iVl6gnin, lB77

Xlgobia musculi Schrank 1781 One specimen frorn A. sgluaticus.

The Myobiidae parasitize small mammals throughout the world and are strongly host specific (Hughes 1959 and Baker and Wharton 1952). M.

ntusculi is mostly found on .11. musculus and sometimes on A. sgluaticus (Bregetova et al. 1955 and Evans et al. 1961). Wharton (1960) showed that interstitial fluid and not blood is the food source.

It

is often responsible for dermatitis in laboratory rats and mice.

It seems to be new to Sweden.

Order Parasitifornr.es

(Reuter)

Zachvatkin Suborder Mesostigmata G. Canestrini,

l8l9

Superfamily Gamasoid,ea Reuter Family Parasitidae Oudemans, l9O2

Pergrmrusus crns.sipes L., 17 58 One female on A. flrwicollis.

Sometimes this free-living mite is found in nests of birds and small mam- mals (Nordberg 1936 and Mrciak et al. 1966). A few records from the fur of small marnmals are also known (Mrciak and Brander 1965 and Edler 1969). Accordin6; to l\Iicherdziriski (1969)

it

is common throughout Europe.

Eugttntrrsus krae pelini Berlese, 1904

Four deutonymphs, one from S. rtrttneus and three from A. flcruicollis.

Normally

it

is free-living, but in some cases

it

has been found on small mammals (Nlrciak and Rosicki 1956 and Ildler 1969).

It

is found in rnost parts of Europe (Micherdziriski 1969).

Eugtrrttttsus re ntbe rti Oudemans, 1912

Eight deutonymphs, five from s. nrnrreus and three from .Jl. crgrestis.

The deutonymph is found both in the nests and on the fur of small mam- rnals in many-paits of Europe. The adult is free-living (Mrciak and Tovornik

1966. Edler 1968 and 1969 and Micherdziiiski 1969).

Eugcnrnsus sp.

Deutonymphs from S. ctrnlteus, N. /odiens and C. gloreolus.

O/ogrrrttnsus sp.

One deutonymph on S. ttrrtnerrs.

According tb triicherdziriski (1969) the name of this genrrs should be Holo- parasitus Oudemans, 1936.

Purasitidae sp.

Five deutonymphs frorn C. glkteolus.

Entomol. Ts. Arg.93. H. l-3, 1972

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EcropARAsrrrc MrrEs (nc,LnlNe) FRoM sMALL

MAMMALS

103 Family Ascaid,ae Oudemans, l9O5

Eurgparasitus emarginafus C. L. Koch, 1839

23 deutonymphs and one male on S. nrrrneus, C. gkreolus, M. agrestis, A. flauicollis and 4. sgluciicus.

It is a free-living predator, common in nests of small mammals in Europe and Asia (Bregetova 1956 and Pirjanik 1962) but sometimes the nymphs are found on the fur of small mammals (Edler 1969).

According to Micherdziriski (1969) the name should be Parusitus oude- monsi Berlese, 1903.

Cgrtolaelaps mucronatus G. & R. Canestrini, 1881

35 deutonymphs were found on S. ar(neus, C. glareolus, M. agrestis, A.

flauicollis, A- sgluaticus and ,\y'. erminea.

It is known from the nests of small mammals and, as deutonymphs, from their fur (Willmann 1952 and Edler 1968 and 1969).

Cgrtolaelaps minor Willmann, 1952

32 deutonymphs found on S. araneus, S. gktreolus and A. sgluaticus.

This species is rare on the fur of small mammals and this is only the second record from Sweden (Mrciak 1959 and Edler 1968).

Family Laelaptid,ae Berlese,

IB92

Hgpoospis sardoa Berlese, 1911 (:Androlaelaps sardous Berlese, 1911 used by Bregetova 1956).

Three females on S. aroneus, C. glareolus and A. sgluaticus.

This mite is mostly found

in

nests and litter, but sometimes on small mammals.

It

is known from Europe (Evans and

Till

1966)

,

but new to

Sweden.

Hgpoctspis hgatti Evans & Till, 1966

One female on S. oroneus.

Evans and

Till

(1966) found it on Bombus muscorum L. in Great Britain.

It is new to Sweden.

Androlaelaps fahrenholzi Berlese, 1911

1925 in Bregetova 1956).

Three deutonymphs and nine females

(:Haemolaelaps glasgorui Ewing, on S. crnneus, C. glarcolus, M. qgres- tis, A. flauicollis and A. sgluaticus.

It

is found in most parts of the world on a variety of small mammals and in their nests (Evans and T'ill 1966, Solomon 1968 and Edler 1969).

It is important as a reservoir for some diseases (Mrciak and Tovornik 1966).

Heemolaeleps casolis Berlese, 1887

Two females on,4. sgluaticus.

It is cosmopolitan, usually found in birds' nests, where reproduction takes place, and rarely on small maurmals (Mrciak 1959 and Ifvans and Till 1966).

Edler (1969) stated that

it

was new to Sweden. However, Sellnick (1956)

Ento,nol. Ts. Arg. 93. H. 1 -3, 1972

(5)

104

ANDERS EDLER

recorded H. molestus Oudernans, 1929 from various localities

in

central Sweden. This is synonymous with H. custtlis (Evans and Till 1966).

Ololaelups sp.

One male on S. nrnneus.

Species of this genus prefer darnp localities.

Eulaelups stabularis C. L. Koch, 1836

421 speciemens, most of them females, found on S. nraneus, N. f odiens, C. glareolus, M. agrestis, A. f lauicollis and A. sgluaticus.

Knorvn from Europe, Asia and North America on various host animals and in their nests (Bregetova 1956 and ilIrciak 1959). Nordberg (1936) found

it

in Finland in nests of Riparia riparia L. Iidler (1968 and 1969) recorded the species in north and central Sweden especially from C. glareolus. In this collection, the highest infestation was found on A. syluoficus, 13.6 0/o (Tab.

4), compared with about 8 0/o on A. llctuicollis and ,11. agrestis. Only 5.3 0/o of the C. glloreolus collected had this mite.

Laelups hiluris C. L. Koch, 1836

926 specimens found on S. eruneus, C. glureolus, M. agrestis, A. flaui- collis and A. sgluaticus.

This species is common throughout Europe. The main hosts are species of ]licrotus and Pitgmgs, the latter not occurring in northern Europe (Mrciak 1959). The infestation frequency of II . ugrestis was 63.6 0/0. On the other host animals it rvas always less than 2 0/0.

Luelops rrgilis C. L. Koch, 1836

4,284 specimens found on S. rtrrtneus, N. fodiens, C. glareolus, )l . agrestis, A. llauicollis and A. sglunticus.

This was the most frequent species forrning 49.9 0lo of the collected mites.

It

is known from most parts of Europe. The main hosts are A. flrtuicollis and A. sglwiicus (Solomon 1968). In the present investigation 69.7 0/o of all collected A. f lauicolli.s and 17 .8 0lo of A. sglurtlicus rvere infested lvith Z.

tqlilis. The other host animals had an infestation frequency below

3

0/0.

Edler (1971) found that significantly more A. flauicollis, ca. 75 0lo, than A.

sgluuticus, ca.50 0/0, were infested by L. agilis in southern Sweden. In central Slveden the infestation rvas about the same, 25 0/0, otr both the species. The nunrber of mites per infested host rvas, however, higher on A. flauicollis than on A. sglurtficrrs both in southern and central Sweden. Similar obsen ations

hzrve been madeby several authors, e.g. Pirjanik (1962) found the infestation frequency on A. flauicollis to be 55.6 0/o and on A. sgluaficus 38.1 0/0. The mean values of mites per infested host were 8.8 and 7.6 respectively.

Hgperloelups nrunlis Zachvatkin, 1948 \:Hllperlaelups nticroti Ewing, 1933 in Evans and Till 1966)

496 specimens found on S. rtrruteus, C. gloreolus, )l . agre.stis, A. fluuicollis nnd .{.. sgluuticus.

55 0lo <tf the .11. uclrestis collected rvere infested by this mite. Other host animals had an infestation frequency below 1 0/0. According to some authors

lI . arualis Pallas is the main host, though species of Pitplmgs also are heavily infested at many places (Mrciak 1959). H. uroalis has a wide ecological

Entonol. Ts. ,lrg. 93. Il. 1 -3, 1972

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ECTopARASrrrc MITEs (,lc,lnrr.u) FRoM sMALL

MAMMALS

105

amplitude and is found in most parts of Europe. In most parts of Scandi- navia and Finland, where M. arualis is not found, M. agrestis is the main host. M. oeconomus Pallas may also be infested by this mite (Edler 1969) .

Mgongssus rossicus Bregetova, 1956

111 specimens on C. glareolus, A. flauicollis and A. sgluaticus.

8.7 olo of the A. flauicollis collected, 1.7 0lo of A. sglucrticus and 0.7 0lo of C. glareolus had this mite.

It

is found in some other European countries, viz. USSR, Czechoslovakia, Bulgaria, Albania and Roumania on species of Apodemus (Bregetova 1956, Mrciak 1958, 1959 and 1960 and Solomon 1968).

It

lives in nests of small mammals, where the development takes place. All

stages may parasitize small mammals (N{rciak 1959).'fhis species is new to Sweden.

Family Haemogamasid,ae Oudemans, 1926

Haemogumasus ftorridus Michael, 1892

25 specimens on S. aruneus, N. fodiens, C. glareolus, A. fluuicollis and A. sgluaticus.

This mite is known from many small mammals in Europe. though

it

is

rnore common

in

their nests (Nlrciak and Brander 1965). The specimens found on the mammals are mostly deutonymphs, 52 0/o of all specimens in this collection. This confirms the observations of Mrciak (1958), l{rciak and Brander (1965) and Edler (1969).

Haemogamasus nidi Michael, 1892

818 specimens on S. aroneus, C. glureolus, M. ugrestis, A. flauicollis and A. sgluuticus.

A mean of 4.0 H. nidi were found on infested C. glareolus, but only 2.1, 2.0 and 2.3 on ll . ugrestis, A. flcruicollis and A. sgluaticus. The infestation frequency on C. glureolus, 15.8 0/0, was a little less than on ,J1. agresfis and A. flauicollis, about 18 0/0. Edler (1971) showed that even if it prefers species

of ]Iicrotus in many countries, e.g. Czechoslovakia, Yougoslavia and Rou- rnania, the main host in Sweden is C. glareolus. Microtidae are more common than Muridae as hosts.

This mite is very common not only in the fur but also in the nests of small mammals. In southern Sweden

I

have found that about 60 0/o of the mites from some rodent nests were 11. nfdi. Furman (1968) recorded the same frequency of this species 1 in rodents' nests in the USA.

H.

nidi is known from Europe, Greenland, the USA and Japan.

It

is a

vector of certain diseases, e.g. tick encephalitis (Mrciak 1960).

Haemogumasus ftirsatus Berlese, 1889

40 specirnens found on S. araneus, C. glareolus, M. agrestis, A. flauicollis and A. sgloaticus.

This mite is found in the nests as well as on the fur of small mammals.

1 Furmau used the name H. antbulons 'Ihorell, giving European authors use the name IJ. ambulans Thorell for

1956 and Evans and Till 1966).

as a synont'rln H. nidi Nlichael.

another species (cf. Bregetova

Entomol. Ts. )rs.93. H,1-3,1972

(7)

106 ANDERS EDLER

The nymphs prefer the fur and the adults the nests (see Mrciak and Tovor-

nik 1966 and Edler 1969).In the present study 21 of the 40 specimens were deutonymphs.

It

has a wide ecological amplitude and is found on a variety of small mammals in all parts of Europe. It is a vector of tick-borne enceph- alitis (Mrciak 1960).

H uemogttntrrsus ft frsutosimilfs \\'illmann, 1952

2l females on A. f lauicollis and A. sgluaticus.

This species is known from many countries in Europe. It is recorded from various mamrnals and especially from their nests (Mrciak 1959, Mrciak et al. 1966 and Evans and Till 1966). This species is new to Sweden.

Family Liponyssid,ae Ewing, 1923

Ifirstionys.s us isubellinus Oudemans, 1913

34 specirrrens found on S. arnneus, C. glareolus, M. agrestis, A. flauicollis, A. slllucrticus and II. erntinea.

Although this collection is small,

it

is evident that species of Microtidae are the main hosts (cf. Mrciak and Brander 1965 and Edler 1969).30 of 34 specimens were found on C. glareolus and M. agrestis.

It

is, however, found in less numbers on a variety of host animals and also in nests (Mrciak 1g5g) .

It is known from many parts of Europe, Asia and North America.

It

is im- portant as vector of tularemia (Mrciak 1960).

Ilirsfionyssu.s soricis Turk, 1945 (:HirstionAssus eusoricis Bregetova 1956) .

I'welve females cln S. rrrnneu.s.

It

is restricted to the Soricidae (Bregetova 1956) . According to literature

it

is very rare.

It

is known from USSR (Bregetova 1956), Czechoslovakia

(Mrciak 1958). Roumania (Feider et al. 1965), Great Britain (Evans and

Till 1966) and Sweden (Edler 1969).

I{trstiony.s.sus ntusculi Johnston, 1894 (:g;.sffonyssrrs kiiscutcttus de Nteil- lon & Lavoipierre, 1944 in Evans and Till 1966).

Ten specimens found on S. araneus, A. flcruicollis and A. sgluaticus.

Species of Muridae are the rnain hosts of this mite, which only rarely para- sitizes Nlicrotidae (I{rciak and Brander 1965). Nine of the ten mites were found on species of Apodemus.

It

is known from Europe and South Africa

(Evans and Till 1966). It is a vector of tularemia (Mrciak 1960).

4.

Frequency

of

various mite speeies on host animals 'l-he frequency of the common gamasid mites on common host animals is found in Tab. 5. Neotrombicula zachuatkini is not included, because of

1,2-1-1 specinrens 1,186 were found on 130 specimens of C. glareolus.

Although .325 S. ereneus rvere collected, only 67 were infested, with a total

of 137 rnites. This is partly due to the rapid death of these hosts in the traps causing the par:rsites to leave the host. In her investigation of fleas Brinck (1966) found that the infestation frequency of S. aranerrs fell from 61 0/o to

Eillottol. Ts- ..lrg. 93. H. 1-3, 1972

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ECTorARASTTIC MITES (eclnrN,l) FRoM sMALL

MAMMALs

107

6 0/0, some time after death.

It

is wellknown that fleas leave a host rather quickly after its death. However, mites are not so mobile as fleas and re- present different ecological groups of ectoparasites (Sosnina 1967 and Edler 1971). In this collection the infestation frequency by mites was 28.2 olo on live specimens of S. craneus and 18.5 0/o on dead ones (Tab. 1). The differ- ence is not significant, 0.05 < P < 0.1. Since the traps were placed in the late afternoon and examined next morning, the dead specimens may have been dead only a few hours.

Two parasites, not mentioned in Tab. 5, Hirstiongssas soricfs, specific to Soricidae, and Cgrtolaelaps mucronatus made up 8.8 0/o and 14.6 0/o respec-

tively of the Gamasoidea on S. nraneus. H. soricis was found only on this host. Less than 1 0/o of the mites on the other hosts consisted of C. mucto- natus. Feider et al. (1965) found C. mucronatus to be the predominant mite, 28.57 0lo, on S. araneus in Roumania.

C- glareolus had 666 gamasid mites, 69.7 0lo of which were Haentogtmasus nidi. This confirms the results of Edler (1971), who showed that C. glareolus is the main host of this mite in Sweden. In Roumania Feider et al. (op cit.) also found H. nidi predominant,51.07 0/0, among the gamasid mites of this host, followed by Laelaps clethrionomgdis, 27.66 0/0, which 'was not repre- sented in the material from southern Sweden. It is found in forest and moun- tain zones, but so far not in the coastal land (Mrciak and Tovornik 1959 and Edler 1968 and 1969). 32.6 o/o live and 26.1 0lo dead C. glureolus wete infested.

Of 1,447 mites found on M. agrestis, 60.5 0/o were Laelaps hilnris, and 31.9 0/o Hgperlaelaps arualis.In Sweden M. agrestis is the main host of these

mite species (see sect.3).81.3 0/o of the live specimens and 81.8 0/o of thc dead ones were infested.

1,966 gamasid mites rvere found on A. f lauicollis. 85.4 010 were Laelaps agilis As mentioned (see sect. 3) it is restricted to species of Apodemus. 79.6 0/o of the live specimens were infested, and three of the six dead.

The greatest number of gamasid mites, 3,109, was found on A. sgluaticus' 82.1 0/o were L. agilis. 57.8 0lo of the live specimens and 33.3 0/o of the dead were infested.

Feider et al. (op cit.) found the same high frequency of L. agilis,86.15 0/0, on A. flauicol/is in Roumania, followed by 11. nidi, 4.58 0/0. In this Swedish collection 5.8 0/o were EL nidi. The predominant species on A. sgluaffcus in Roumania were Haemolaelaps glasgowi

(:

Androlaellps f uhrenholzi), 46.94o10, and Z. agilis,30.16 0/0, followed by 11.nidi,8.39 0/0, and E. stlbu- Iuris, 7.94 0/0. In this collection from Sweden 9.6 0/o were E. stabularis and 5.5 0lo H. nidi. Only on A. sgluaticus was there a significant difference be- tween the infestation frequencies of live and dead hosts. Similar calculations

for ,4. flauicollis were impossible because only six dead animals were col- lected.

The frequency of total infestation differed significantly between the five host species, except M. agrestis and A. flauicollis. The difference between them was only 2.5 0/o (0.5 < P < 0.6). This makes

it

possible to rank the host species as regards frequency of infestation. About one fifth of the S.

eranelts were infested, one third of the C. glareolus, one half of the A. sgl- ucrticLts and four fifths of the A. flcruicollis and M. agrestis (Tab. 1).

Entomol. Ts. ,(rs. 93. H. 1 -3, 1C72

(9)

108 ANDERS EDLER

5.

Acknowledgements

The rvork was carried out at the Department of Animal Ecology, University of Lund.

NIy thanks are due to Professor Per Brinck, Lund, and Professor llilan l{rciak, Brati- slava, Czechoslovakia, for valuable advice, to )Ir. Lennart Hansson, Miss Gunilla Lind- quist and IIr. Anders Nilsson for help with the field and technical work and determination of the host animals.

The work was supported by grants frorn the Swedish National Science Research Council;

the Faculty of llathematics and Natural Sciences of the University of Lund, the Research Institute of National Defence and the Royal Swedish .{cademy of Engeneering Sciences.

6.

Abstract

8,582 mites of 29 species rvere collected from small mammals in southeln Srvedeu.

Five species \\'ere ne\y to Srveden. The distribution and host relationship is discussed.

A fifth of the Sorcr oteneus, a third of the Clefftrionomgs glareolus, half the Apo- dentus sylu(ficus and four fifths of the A. llauicollis and llicrotus ogresfis x.ele infested. There rvas no difference in the infestation of live ancl dead specirnen,;

excel)t for .{. .sylaatictt.s.

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Fac. Rerum Nat. Univ. Comen. 12

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- Acta Fac. Rerum Nat. Univ. Comen. 12

- Zoologia 13:117-139.

Bratislava.

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Specimens Infestation frequency

infested total 品鳳 1盟i燎

.Sorec minufu.s L. . .

S. araneus L. . ....

Neomgs fodiens Penn.

Ctethiion'omgs glareolus S;;;;;." : : :.

Microtus agrestis L. ....

Apodemus /lauico1lis N'Ielch.

A. sgloatictts L. ....

,[Iu.s musculus L.

Rattus noruegicus Berkenhout . .. .. . ..

Mustele erminett L. ....

26 325

5 735 151 310 917 5 2 1

2,507

0 67 3 235 123 245 532 0

20.6

320815

79.0 56.2

28.2 32.6813

79.6

18.5

26.1 81.8

57.8 33.3 0 -―

1 - 1,206 48.1

Tab.1.Frequency of infestation of the sma11 lnanlmals collected.

Σ

(11)

Stampen,

Blentarp Hlllestad Lund Barse‐bick Kullaberg

89

︲2

︲2 84 48 03 34

・4 00 22

︲9 99 78 89

39

︲2 75 75

110 ANDERS EDLER

Tab 2.DistributiOn of nlite species by locality.

1,827 2,078

03

2,569 3,460

ιοれο′.rs./r夕.93.″.r-3,1972

(12)

Tab. 3. Distribution of mite species between infested host species.

.ヽ

.で ヽヽ.く

.ヽ

- 9 -―

Mgobia musculi 1 -

Pergamasus crassipes

Eugamasus kraepelini I

-

1 -

3 -

E. remberti 5

- 3 -

1,244 -― ――

―― - 4

︼≦

38 62

・7 5︲

6。7 6。7

Eugamasus sp. . .. 2 I

Ologatnasus sp. . ... I -

Parasitidae sp. ...

Durgparasitus emarginatus .... 6 -

Cyrtolaelaps mucronatus ... 20

C. minor 4 -

Hgpoaspissardoa. I

H. hgatti I --

Androlaelaps lahrenholzi ... I -

Haemolaelaps casalis

Ololaelaps sp. . .. I -

Eulaelaps sfabulnris I 2

.■frs“fus.… .….… .…・…・・ 17 - ' . frsIIrOsfrlliris ...・・・・・・・・・

frsffοnyss″ s lsαι

`′

in“s .….… 1 -

″。sοricis ...… … … … … …・・…・ 12 -

4 - ―― 5 -― ―― ―― ――

3 1 11 3 -―

58 18 36 298 -―

7 2 1 1 1 27 -― - 1 -―

1 - - 1 - 3323-

- 2 -

- 16 876 3 7 -―

2 38 4 1,678 2,554 -―

- 4 462 22 3 -―

- 5 -- 65 41 -―

2 10 -- 4 2 -―

- 164 58 115 171 -―

4 - 7 -

1 1

415 421 869 926 3,740 1,284 332 496 88 111 6 25 740 818 6 40 21 21 31 34 12 12 10 10 6,286 7,334 6,290 8,582

8 3 8

-― - 14 13 17 1

5 1.249

13 3

iヨ I置

&,ll二

.‐

I:‖ : ; 1,:::七

:::士

:ま

‖ : 3

100 336100 336

Ю O■

References

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