Host-parasite associations of Eimeria spp. (Apicomplexa:Eimeriidae) in kangaroos and wallabies of...

InrernarionolJoournnlforPnras~rology Vol. 19, No. 3,pp. 241-263, 1989. OOZO-7519/89 $3.00 + 0.00 Printed in Grear Brirain. Pqamon Press plc

0 1989 Ausrralian Sociqfor Porasirology.

HOST-PARASITE ASSOCIATIONS OF EIMERIA SPP. (APICOMPLEXA:EIMERIIDAE) IN KANGAROOS AND WALLABIES OF

THE GENUS MACROPUS (MARSUPIALIAMACROPODIDAE)

IAN K. BARKER,*? MICHAEL G. O’CALLAGHANS and IAN BEVERIDGE~Z

*Department of Pathology, Ontario Veterinary College, University of Guelph, Guelph, Ontario, Canada NlG 2Wl *Central Veterinary Laboratories, Department of Agriculture, I.M.V.S., Frome Road, Adelaide, South Australia 5000,

Australia

(Received 21 June 1988; accepted 20 October 1988)

AbsfraCf-BARKER I. K., O’CALLAGHAN M. G. and BEVERIDGE I. 1989. Host-parasite associations of Eimeria spp. (Apicomplexa:Eimeriidae) in kangaroos and wallabies of the genus Macropus (Marsupi- alia:Macropodidae). International Journal for Parasitology 19: 241-263. Faecal samples from 514 kangaroos and wallabies representing 12 species of the genus Macropus were examined for oocysts of Eimeria spp. Six species of Eimeria were redescribed from their type hosts, and on the basis of finding homologous oocysts in the faeces of other Macropus spp., host ranges for these coccidia were extended. Eimeria hestermani Mykytowycz, 1964 is redescribed from M. giganteeus (eastern grey kangaroo) and is described from M. fuliginosus (western grey kangaroo), M. rufogriseus (red-necked wallaby), M. dorsalis (black-striped wallaby), and M. eugenii (tammar wallaby). E. toganmainensis Mykytowycz, 1964 is redescribed from M. rufus (red kangaroo) and the host range is extended to M. giganteus, M. fidiginosus, M. rufogriseus and M. eugenii. E. wilcanniensis Mykytowycz, 1964 is redescribed from M. rufus, and the host range is extended to M. giganteus, M. jidiginosus and M. robustus (euro or wallaroo). E. macropodis Wenyon & Scott, 1925 is redescribed from M. rufogriseus, and is described from M. giganteus, M. filiginosus, M. @us, M. irma (western brush wallaby), M. parryi (whip-tailed wallaby), M. dorsalis, M. eugenii, and M. parma (parma wallaby). E. fausti Yakimoff & Matschoulsky, 1936, E. cunnamullensis Mykytowycz, 1964 and E. purchasei Mykytowycz, 1964 are synonymized with E. macropodis. E. marsupialium Yakimoff & Matschoulsky, 1936 is redescribed from M. giganteus, and from M. fuliginosus. E. gungahlinensis Mykytowycz, 1964 is redescribed from M. fuliginosus, and from M. giganteus. Seven new species of Eimeria are described. E. jlindersi, new species, is described from M. eugenii, M. rufogriseus, and M. antilopinus (antilopine wallaroo). E. prionotemni, new species, is described from M. eugenii, M. panyi, M. rufogriseus, M. agilis (agile wallaby) and M. dorsalis. E. mykytowyczi, new species, is described from M. agilis, M. antilopinus, and M. parryi. E. panyi, new species, is described from M. parryi. E. yathongensis, new species, is described from M. fuliginosus and M. giganteus. E. parma, new species, is described from M. parma, and E. desmaresti, new species, is described from M. rufogriseus. E. kogoni Mykytowycz, 1964, and E. rufusi Prasad, 1960 are considered species inquirendae. The host- parasite associations of these coccidia, and of similar species of Eimeria in other genera of Macropodoid marsupials, are discussed in relation to the postulated phylogeny of the hosts.

INDEX KEY WORDS: Eimeria; coccidia; new species; Marsupialia; marsupial; Macropodidae; kangaroo; wallaby; Macropus; phylogeny.

INTRODUCTION

COCCIDIA (Eimeria spp.) have been described from only four of the 13 extant species of kangaroos and wallabies comprising the genus Macropus. Two species of Eimeria were described from the red- necked wallaby M. rufogriseus by Wenyon 8~ Scott (1925) and Zwart & Strik (1964). Three have been described in the red kangaroo M. r@s by Prasad (1960) and Mykytowycz (1964). Seven are reported from the eastern and western grey kangaroos, M. giganteus and M. fuliginosus, identified variously as M. gigunteus by Yakimoff & Matschoulsky (1936),

_FTo whom correspondence should be addressed.

or as M. cangunt by Mykytowycz (1964). These reports are qualified by the fact that they are, in many cases, based on one or two animals held in zoos (Wenyon & Scott, 1925; Yakimoff & Matschoulsky, 1936; Prasad, 1960; Zwart & Strik, 1964). Further- more, the population from which most of the coccidia of grey kangaroos were described (Mykytowycz, 1964) was subsequently demonstrated to be comprised of two sympatric, closely-related species (Kirsch & Poole, 1972). Hence the host-parasite associations of the coccidia of eastern and western grey kangaroos are uncertain. Only coccidia of the red kangaroo have been described from a large wild population of a well-defined species.

241

242 I. K. BARKER, M. G. O’CALLAGHAN and I. BEVERIDGE

Eimeria spp. are considered relatively host- specific, and seem morphologically conservative, host specificity evolving more rapidly than oocyst morphology (discussed in Barker, O’Callaghan, Beveridge 81 Close, 1988). As a result, the genus Eimeriu may prove a useful tool for investigating host phylogeny, since a ‘trail’ of homologous oocysts might serve as a marker for an evolutionary lineage of hosts (Vance & Duszynski, 1985; Wash, Duszynski & Yates, 1985; Duszynski & Moore, 1986, exemplify this approach).

On this premise, we have undertaken a study of the Eimeria spp. parasitizing kangaroos and wallabies, the marsupial superfamily Macropodoidea, seeking relationships among the host distribution of coccidial oocysts of various morphologic types, or ‘species’, and the phylogeny of their hosts (Barker, O’Callaghan, Beveridge & Close, 1988; Barker, O’Callaghan & Beveridge, 1988a, b). The Macropodoidea has radiated widely in Australia and New Guinea over the last 1.5 million years (Archer, 1984; Flannery, 1984). The phylogeny of the group is being clarified as tradi- tional taxonomic methods are supplemented by studies of cytogenetic, serologic and biochemical relationships (see Kirsch, 1977; Richardson & McDermid, 1978; Archer, 1984). However, altema- tive approaches to this problem may prove illuminat- ing or corroborative. Patterns of host-parasite association among coccidia of Australian marsupials also may be of interest to compare with coevolutionary relationships of helminths parasitizing the Marsu- pialia (Beveridge, 1986).

Significant homology was not evident among Eimeria oocysts from members of the Potoroidea (Barker, O’Callaghan & Beveridge, 1988a), and from some kangaroos and wallabies of the Maropodidae, other than Mucropus (Barker, O’Callaghan & Beveridge, 1988b), where comparisons could be made only at the level of host genus. However, homologies were present among the Eimeriu oocysts in nine species of rock wallabies (Petrogale spp.), such that a single Eimeria species (defined by similar oocysts) was present in up to six species of Petrogale. Furthermore, the patterns of distribution of distinctive species, or suites of species, of coccidia suggest that Eimeria may be tagging phylogenetic lines within that genus (Barker, O’Callaghan, Beveridge & Close, 1988).

Macropus, the largest genus in the Macropodidae, is comprised of three groups of species (Dawson & Flannery, 1985). The largest of these, the ‘brush wallabies’ of the subgenus Notamacropus, includes M. agilis, M. dorsalis, M. eugenii , M. irma, M. parma, M. panyi and M. rufogriseus. The subgenus Osphranter includes M. rufus, M. robustus, M. antilopinus and M. bernardus. M. giganteus and

M. fuliginosus comprise the subgenus Macropus. The reports of Eimeria spp. in the genus Macropus

cited above, based on one, or in the case of the grey kangaroos, probably two representatives of each subgenus, suggest that there is no homology in

coccidial oocysts among hosts of this genus. We undertook the study reported here to extend knowledge of the coccidian fauna parasitic in Macropus to most species in the genus, and to attempt to determine if apparent host-parasite associations might be at the level of host species, subgenus, or otherwise.

MATERIALS AND METHODS

Faeces were collected from animals of 12 species of Macropus (Table l), from a variety of localities (see results). Animals from which faeces were collected were shot, road- kills, captives, or free-living individuals identified on the basis of phenotype by competent zoologists during the course of other studies. Faeces were placed in 2.5% potas- sium dichromate, and oocysts were allowed to sporulate at room temperature; they were subsequently stored at 4°C. They were examined at invervals of a few days to 12 years after collection. Sporulation time was not determined, since specimens usually spent some time in transit to the laboratory. Oocysts were concentrated by flotation in sucrose solution (SG 1.30) and were examined under an oil immersion 100X Planachromat objective on an Olympus BH microscope with a Nomarski differential interference contrast system, or with a Planapochromat objective on a Reichert Zetopan Microscope. Measurements were made with an eyepiece graticule calibrated with an objective micrometer. Oocysts were recorded as they were encountered. They were assigned to species on the basis of oocyst morphology, on the conservative assumption that, in the absence of experimental information on cross-infectivity to prove otherwise, homologous oocysts from different species of host were the same species of Eimeria. The host in which a new species was most prevalent or common was desig- nated the type host, and localities from which oocysts were measured are indicated in the text by *. Where possible, a minimum of 20 oocysts of each species was measured from at least one individual of each host species infected. Measurements are recorded in the text, or in Tables 2- 11, as the mean + S.D. (range). Analyses of variance were per- formed on the dimensions of oocysts of species occurring in several different hosts. Least significant differences (lsd) were calculated at the 5% level, and are indicated in tables. Names of Australian states are abbreviated as follows: New South Wales-N.S.W.; Queensland-Qld.; South Australia- S.A.; Tasmania-Tas.; Victoria-Vie.; Northern Territory- N.T.; Western Australia-W.A. Station property names are included within quotation marks. Nomenclature of hosts follows Strahan (1983).

RESULTS

The prevalence of infection with each species of Eimeria in the various hosts is recorded in Table 1. All species of Eimeria recognized are described or redescribed below; new species are erected, and where appropriate, synonymies are established. Diagnostic features allowing identification of the coccidia described in Macropus spp. are summarized in Table 12.

Eimeria hestermani Mykytowycz, 1964 Figs. 1, 16-19

Description. Oocyst ovoid; 26 measured from two animals from two localities, Table 2; double oocyst wall; outer layer rough, brown, partly opaque, 1.6-

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- -

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- -

- -

- -

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M. e

ugen

ii 71

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7/20

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20/2

0 -

- -

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f st

udy.

244 I. K. BARKER, M. G. O’CALLAGHAN and I. BEVERIDCE

FIGS. 1-5. Drawings of oocysts of Eimcriu spp. from kangaroos and wallabies of the genus Mmropus. Scale = IO pm. all figures to same scale.

FIG. I. E. hesrermuni from M. giganreus.

FIG. 2. E. iogunmainensis from M. rujrs.

FIGS. 3-5. E. flindersi: FIG. 3, from M. eugenii; FIG. 4, from M. rufogriseus; FIG. 5, from M. anlilopinus.

3.2 pm thick, readily fractured and separated from inner layer; outer layer slightly thicker around micropyle 4.0-6.0 pm in diameter present at the more pointed end; inner clear, colourless, 0.8 pm thick; polar granule variably present; no oocyst residuum; four ellipsoidal sporocysts, more pointed at end with Stieda body and prominent lenticulate substieda body; two sporozoites with finely granular cytoplasm filling each sporocyst; refractile globule 5.6-6.8 pm in diameter, up to 9.2 iurn long; sporocyst residuum a tight aggregate of granules, up to 8.0 pm in diameter,

or loose aggregate of granules scattered about equator of sporocyst.

Diagnosis. This species is readily recognized by its large size, and distinctive oocyst wall.

Type host. Macropus giganteus, eastern grey kangaroo. Localities: Yathong Nature Reserve* near Mount Hope, N.S.W.; Bondo*, N.S.W.; Yan Yean, Vie.

Other hosts. M. fuliginous, western grey kangaroo. Eighty-five measured from four animals in three localities, Table 2. Localities: Second Valley*, !%A.;

Coccidia of Macropus spp. 245

14 15

FIGS. 6-15. Drawings of oocysts of Eimerb spp. from kangaroos and wallabies of the genus Macropus. Scale = 10 ,um, all

figures to same scale.

FIG. 6. E. prionotemni from E. eugenii.

FIG. 7. E. wilcunniensis from M. rufus. FIG. 8. E. mykyfowyczi from M. ugilis.

FIG. 9. E. puryi from M. paryi.

FIG. 10. E. macropodis from M. rufogriseus.

FIG. 1 I. E. mursupialium from M. giganteus.

FIG. 12. E. yathongensis from M. fuliginosus.

FIG. 13. E. parma from M. parma.

FIG. 14. E. gunguhlinensis from A!. fuliginosus.

FIG. 15. E. desmaresti in M. rufogriseus.

Para Wirra Conservation Park*, S.A.; ‘Pine Plains’*, sured from one animal, Table 2. Locality: ‘Harvest via Patchewollock, Vie.; Kangaroo Island, S.A. Home’*, via Charters Towers, Qld.

M. rufogriseus, red-necked (Bennett’s) wallaby. M. eugenii, tammar wallaby. Twenty measured Twenty-three measured from four animals, Table 2. from pool of faeces, Table 2. Localities: captive Localities: Connerville*, Bridgenorth, and Launces- animals, Flinders University, Adelaide*, S.A.; ton, Tas. Kangaroo Island, S.A.

M. dorsalis, black-striped wallaby. Twenty mea- Comments. Mykytowycz (1964) did not describe

246

16

I. K. BARKER, M. G. ~‘CALLAGHAN and I. BEVERIDGE

18

- 10pm

FIGS. 16-19.


Stieda and substieda bodies, but they are visible in his illustration. Pellerdy (1974) recognized the likely identity of Globidium mucosum (Zwart & Strik, 1964) with E. hestermani Mykytowycz, 1964, and synonymized the former with the latter on the grounds of priority. Since the population of grey kangaroos (as M. canguru) from which Mykytowycz (1964) described the species was comprised of both M. giganteus and M. filiginosus (see Kirsch & Poole, 1972), the former is nominated here as the type host for the redescribed species. We confirm that this species also occurs in M.fuliginosus, though oocysts were not found in material examined by us from 92 M. jidiginosus at Yathong Nature Reserve at Mount Hope, N.S.W., the type locality for the species. We extend the host range to h4. dorsalis and M. eugenii. Endoge- nous development of gametocytes takes place in the lamina propria of the small intestine (Barker, Harri- gan & Dempster, 1972), and we have observed developing oocysts characteristic of this species in tissue sections from the intestines of M. agilis (Barker, unpublished), from which oocysts have yet to be described. Though there is considerable variation in size among the oocysts from the several host species, in the absence of experimental information on host specificity, we have included all in E. hestermani.

Eimeriaflindersi new species Figs. 3-5,20-22

Description. Oocyst ovoid to nearly ellipsoidal; 20 measured from one animal, Table 3; double oocyst wall; outer layer finely mamillate at more pointed end of oocyst, surrounding micropyle, otherwise smooth, clear, dark tan, 1.4 pm thick; inner layer clear, 0.4 ,U m thick; micropyle up to 5.6 pm in diameter; polar granule present; oocyst residuum absent; four ellipsoidal to slightly ovoid sporocysts, typically distributed in basal portion of oocyst, away from micropyle; inconspicuous Stieda body, and lenticulate substieda body present; two sporozoites with finely granular cytoplasm, nearly filling sporocyst; refractile globule 4.0 pm in diameter; sporocyst residuum granular aggregate up to 8.0 pm in diameter, at equator of sporocyst.

Diagnosis. The combination of larger oocyst and sporocyst size, and tan hue, distinguish the oocyst from E. toganmainensis, which it resembles. E. lagorchestes in the spectacled hare-wallaby (Barker, O’Callaghan & Beveridge, 1988b) overlaps the upper end of the oocyst size range, but differs in having a thicker, diffusely mamillate wall.

Type host. Macropus eugenii, Fig. 2. Locality: Kangaroo Island*, S.A.

Other hosts. M. rufogriseus, Fig. 3. Twenty-one measured from two animals, Table 3. Locality: Connerville*, Tas.

M. antilopinus, antilopine wallaroo, Fig. 4. Eigh- teen measured from one animal, Table 3. Locality: captive, Pallarenda, Townsville*, Qld.

Comments. E. flindersi in M. eugenii and M. rufogriseus is clearly separate from E. toganmainensis, which also occurs in those hosts, on the basis of size and colour. The specimens of E. flindersi in M. antilopinus, from a single animal, lack mamillation and tan colouration of the oocyst wall and are inter- mediate in oocyst and sporocyst dimensions between E.flindersi in M. eugenii and M. rufogriseus, and E. toganmainensis. On the grounds that the sporocysts are relatively wider than those of E. toganmainensis, the oocysts from M. antilopinus are included, with reservation, in E. flindersi. The species is named for Matthew Flinders, who charted the waters of much of the Australian coast, including Kangaroo Island and Tasmania. Oocysts in faeces; endogenous stages undescribed.

Eimeria toganmainensis Mykytowycz, 1964 Figs. 2,23-26

Description. Oocyst bluntly ellipsoidal to ovoid, occasionally slightly flattened on one side; 65 measured from three animals in two localities, Table 4; double oocyst wall; outer layer smooth, clear, slight purple-pink cast, 1.2 ,~rn thick; outer wall thins at more pointed end, forming micropyle 3.2-4.0 pm diameter; inner wall clear, colourless, 0.4 ,um thick; polar granule variably present; no oocyst residuum; four sporocysts, usually distributed in basal portion of oocyst, away from micropyle; sporocysts approximately ellipsoidal, slightly pointed toward end with inconspicuous Stieda body and lenticulate substieda body; two sporozoites with finely granular cytoplasm, nearly filling sporocyst; refractile globule up to about 5.6 pm; sporocyst residuum granular aggregate up to 6.4 pm diameter at equator of sporocyst.

Diagnosis. This species is separated from E.flindersi on the basis of smaller oocyst and sporocyst size, and colour. It differs from E. prionotemni, new species, in disposition of sporocysts within the oocyst, in having a micropyle, and by consistently lacking an oocyst residuum. It is larger than E. wilcanniensis and E. mykytowyczi and differs from them in having a Stieda/substieda body. E. lagorchestes in the spec-

FIGS. 16-19. Photographs of Eimeria hestermani oocysts from various spp. of kangaroos and wallabies of the genus Macropus. Scale = 10 pm, all figures to same scale.

FIG. 16. E. hestermani in M. giganteus, the type host.

FIG. 17. E. heslermani in M. fuliginosus. FIG. 18. E. hestermani in M. rufogriseus.

FIG. 19. E. hestermani in M. dorsalis.

TABLE

~-DIMENSIONS

(urn

) OF OOCYSTS

AND

SPOROCYSTS

OF

Eim

eria

he

ster

man

i IN M

acro

pusg

igan

teus

, M

.fuli

gino

sw,

M.

rufo

gris

eus,

M

. do

rsal

is A

ND

M.

euge

nii

Hos

t

Ooc

yst

Spor

ocys

t

Len

gth

Wid

th

Len

gth

Wid

th

n*

xi

S.D.

Ran

ge

it S.D.

Ran

ge

it S.D.

Ran

ge

,Ct

S.D.

Ran

ge

M. g

igan

teus

26

64

.7

2.10

61

.6-6

8.6

45.3

1.

95

41.6

-48.

3 24

.9

0.96

23

.0-2

6.6

13.4

0.

83

12.0

-15.

4

(typ

e)

M. f

ulig

inos

us

85

71.1

3.

32

63.2

-77.

0 50

.3

3.23

43

.2-5

6.0

24.6

1.

60

19.2

-29.

4 14

.2

0.67

12

.8-1

5.8

‘M.

cang

uru’

50

66

.1

- 63

.2-6

9.0

46.6

-

45.0

-49.

1 24

.5

- 22

.1-2

6.6

13.6

-

13.0

-14.

6 M

ykyt

owyc

z (1

964)

M

. ru

fogr

isew

23

64

.9

2.89

60

.8-7

2.0

40.5

2.

10

37.6

-44.

8 25

.2

2.07

21

.6-3

0.6

13.1

1.

01

12.0

-15.

3 Z

war

t an

d St

rik

6 62

.8

- 59

.0-6

5.0

43.8

-

41.0

-47.

0 23

.7

- 20

.0-2

6.0

11.7

-

11.0

-12.

0 (1

964)

M

. do

rsal

is

20

63.5

1.

61

60.0

-67.

2 44

.2

1.13

41

.6-4

5.6

25.9

1.

05

24.8

-28.

0 13

.3

0.53

12

.0-1

4.4

M.

euge

nii

20

59.8

1.

86

56.8

-62.

4 39

.2

1.36

36

.0-4

0.8

24.6

1.

27

22.4

-27.

2 13

.3

0.54

12

.0-1

4.4

Pool

ed

174

67.1

4.

95

56.8

-77.

0 46

.3

5.02

36

.0-5

6.0

24.9

1.

54

19.2

-30.

6 13

.7

0.84

12

.0-1

5.8

(ori

gina

l)

Isd

(5%

) 4.

3 4.

1 0.

6 0.

4

*Num

ber

of o

ocys

ts

mea

sure

d.

iMea

n di

men

sion

.

TABLE

~-DIMENSIONS

(,um

)o~

OOCYSTSAND

SPOROCYSTSOF

Eim

eria

flin

ders

i NEW

SPECIES,IN

Mac

ropu

seug

enii

,M

rufo

gris

eus

AND

M.

anti

lopi

nus

Ooc

yst

Spor

ocys

t

Len

gth

Wid

th

Len

gth

Wid

th

Hos

t n*

xt

S.D.

Ran

ge

J3

S.D.

Ran

ge

Xi

S.D.

Ran

ge

Xi

S.D.

Ran

ge

M.

euge

nii

20

44.2

1.

55

42.4

-48.

8 29

.2

1.19

26

.8-3

2.0

18.1

0.

73

16.8

-19.

4 10

.6

0.60

9.

6-11

.6

(typ

e)

M.

rufo

gris

eus

21

46.8

1.

80

42.0

-49.

0 26

.0

0.63

25

.5-2

7.5

17.6

2.

00

14.3

-22.

4 11

.1

0.60

10

.0-1

3.0

M.

anti

lopi

nus

18

41.2

2.

34

37.6

-47.

2 24

.3

1.76

20

.8-2

8.0

15.0

0.

76

13.6

-16.

0 11

.1

0.54

10

.4-1

2.0

Pool

ed

59

44.2

2.

95

37.6

-49.

0 26

.6

2.34

20

.8-3

2.0

17.0

1.

88

13.6

-22.

4 10

.9

0.63

9.

6-13

.0

Isd

(5%

) 4.

5 4.

0 2.

7 0.

5

*Num

ber

of o

ocys

ts

mea

sure

d.

tMea

n di

men

sion

.

249

FIGS. 20-28. Photographs of oocysts of various Eimeria spp. from kangaroos and wallabies of the genus Macropus. Scale = 10 ,um, all figures to same scale.

FIGS. 20-22. E.Jlindersi: FIG. 20, from M. eugenii, type host; FIG. 21, from M. eugenii, showing mamillations on outer surface of oocyst wall at pole around micropyle; FIG. 22, from M. rufogriseus.

FIGS. 23-26. E. toganmainensis: FIG. 23, from M. rufus, type host; FIG. 24, from M. giganteus; FIG. 25, from M. rufogriseus; FIG. 26, from M. eugenii.

FIG. 27. E. prionotemni from M. eugenii. Note oocyst residuum.

FIG. 28. E. parryi from M. parryi.

TA

BL

E

~-D

IME

NS

ION

S

(pm

) O

F O

OC

YS

TS

A

ND

S

PO

RO

CY

ST

S

OF

Eim

eria

to

ganm

aine

nsis

IN

Mac

ropu

s ru

fus,

M

. gi

gant

eus,

M

. fui

igin

osus

, M

. ru

fogr

iseu

s A

ND

M.

euge

nii

Hos

t

Ooc

yst

Spor

ocys

t

Len

gth

Wid

th

Len

gth

Wid

th

n*

xi

S.D

. R

ange

xi

S

.D.

Ran

ge

Yt

S.D

. R

ange

*z

i S.

D.

Ran

ge

M.

rufu

s

(typ

e)

Myk

ytow

ycz

(196

4)

M.

giga

nteu

s M

. ful

igin

osus

M

. ru

fogr

beus

M

. eu

geni

i

Pool

ed

(ori

gina

l)

lsd

(5%

)

65

41.0

2.

27

35.2

-44.

8 26

.2

2.18

21

.6-3

0.0

15.5

0.

73

14.4

-17.

0 9.

8 0.

63

8.8-

I 1.

5

50

38.7

-

34.1

-44.

3 24

.9

- 23

.2-2

7.3

15.8

-

13.0

-17.

7 9.

7 -

8.2-

10.9

20

35.2

1.

74

32.0

-37.

6 19

.9

0.91

18

.4-2

1.6

14.6

0.

75

13.6

-16.

0 9.

0 0.

48

8.0-

9.6

35

37.2

1.

67

33.6

-40.

0 21

.4

1.22

19

.2-2

4.0

16.1

0.

86

14.0

-17.

6 9.

0 0.

60

8.0-

10.4

24

36

.4

2.63

32

.8-3

6.2

21.2

1.

14

19.2

-24.

0 14

.2

1.03

12

.0-1

6.0

8.5

0.50

7.

8-9.

6 20

35

.5

1.77

32

.8-3

6.2

21.3

0.

95

19.2

-22.

4 14

.1

0.67

13

.3-1

6.0

8.7

0.38

8.

0-9.

6

164

38.1

3.

15

32.0

-44.

8 23

.1

3.01

18

.4-3

0.4

15.1

1.

09

12.0

-17.

6 9.

2 0.

76

7.8-

11.5

2.6

2.8

1.0

0.6

*Num

ber

of o

ocys

ts

mea

sure

d.

iMea

n di

men

sion

.

TA

BL

E ~

-DIM

EN

SIO

NS

(,

um)o

~ O

OC

YS

TS

A

ND

S

PO

RO

CY

ST

S O

F E

imer

iapr

iono

tem

ni

IN M

acro

pus

euge

nii,

M.

part

yi,

M.

rufo

gris

eus,

M

. ag

ilis

AN

D M

. do

rsal

is

Ooc

yst

Spor

ocys

t

Hos

t

M.

euge

nii

(typ

e)

M. p

arty

i M

. ru

fogr

iseu

s M

. ag

ilis

M.

dors

alis

Pool

ed

Isd

(5%

)

Len

gth

Wid

th

n*

xi

S.D

. R

ange

xt

S

.D.

Ran

ge

20

37.1

2.

11

32.0

-40.

8 22

.0

0.99

20

.0-2

3.2

45

35.2

2.

37

30.4

-41.

2 21

.2

0.90

20

.0-2

3.7

20

33.7

1.

24

32.0

-36.

0 21

.4

0.72

19

.6-2

2.4

31

35.1

1.

79

31.2

-38.

6 21

.3

1.05

18

.4-2

2.6

2 32

.4

- 32

.0-3

2.8

22.6

-

22.4

-22.

8

118

35.2

2.

25

30.4

-41.

2 21

.4

1.00

18

.4-2

3.7

1.3

0.4

Len

gth

Wid

th

xt

S.D

. R

ange

X

i S.

D.

Ran

ge

15.7

0.

50

14.4

-16.

0 10

.1

0.57

9.

6-11

.2

14.3

0.

85

12.8

-16.

5 8.

6 0.

60

7.2-

9.8

13.4

0.

70

12.0

-14.

4 8.

6 0.

41

8.0-

9.2

14.4

1.

27

12.4

-16.

5 8.

3 0.

34

7.8-

9.2

14.4

-

13.6

-15.

2 8.

4 -

8.0-

8.8

14.4

1.

13

12.0

-16.

5 8.

8 0.

79

7.2-

11.2

0.6

0.5

*Num

ber

of o

ocys

ts

mea

sure

d.

tMea

n di

men

sion

.


tacled hare-wallaby (Barker, O’Callaghan & Bever- idge, 1988b), differs in being larger, with a thicker oocyst wall.

Type host. Macropus rufus, red kangaroo. Locali- ties: Yathong Nature Reserve*, near Mount Hope, N.S.W.; ‘Avenel’, via Broken Hill, N.S.W.; captive, Cleland Conservation Park, Adelaide*, S.A.

Other hosts. M. giganteus. Twenty measured from two animals, Table 4. Localities: Bondo*, N.S.W.; Mitta Mitta*, Vie.

M. fuliginosus. Thirty-five measured from two animals, Table 4. Localities: Second Valley*, S.A.; Para Wirra Conservation Park*, near Kersbrook, S.A.; Bangham, S.A.; Kangaroo Island, S.A.

M. rufogriseus. Twenty-four measured from three animals, Table 4. Localities: Connerville*, Bridge- north*, and Launceston*, Tas.

M. eugenii. Twenty measured from one animal, Table 4. Locality: Kangaroo Island*, S.A.

Comments. Mykytowycz (1964) did not refer to the presence of a Stieda/substieda body in E. togun- mainensis, and his illustration is inadequate in this regard. He described this species from M. rufus, but not from M. canguru (= M. giganteus and M. fuliginosus). This may be the result of geographic variation in prevalence, since we also failed to find E. toganmainensis in samples of 92 M.fuliginosus and 82 M. giganteus from Mount Hope, a site studied by Mykytowycz (1964), though it was present in smaller samples of both species from other localities. Oocysts from M. rufus were somewhat larger than those from other hosts, but this was not considered sufficient to justify a new species. Oocysts in faeces; endogenous stages undescribed.

Eimeriaprionolemni new species Figs. 6,27

Description. Oocyst bluntly ellipsoidal or slightly ovoid; 20 measured from two animals, Table 5; double oocyst wall; outer layer 1.2- 1.6 ,u m, thinning at one pole of oocyst, but no micropyle; light tan hue, clear; inner layer 0.4-0.8 pm, clear; polar granule present; oocyst residuum, comprised of a cluster of three-eight granules, usually present; four ellipsoidal to ovoid sporocysts, with inconspicuous Stieda body, and lenticulate substieda body; two sporozoites with finely granular cytoplasm; refractile globule up to 5.6 X 4.0 pm; sporocyst residuum finely granular clump up to 5.6 pm diameter at equator of sporocyst.

Diagnosis. This species is separated from E. toganmainensis in disposition of sporocysts within the oocyst, in lacking a micropyle, and by usually possessing an oocyst residuum. It is separated from E. wilcanniensis and E. mykytowyczi, new species, in lacking a micropyle, but possessing a Stieda/substieda body. E. thylogafe from the Tasmanian pademelon (Barker, O’Callaghan & Beveridge, 1988b) has a thicker, irregular oocyst wall. E. aepyprymni, from the rufous bettong (Barker, O’Callaghan & Beveridge, 1988a), and E. inornata from rock wallabies (Barker,

O’Callaghan, Beveridge & Close, 1988), are similar, but are considered distinct on the basis of phylogenetic separation of their hosts from the Macropus spp. infected with E. prionotemni.

Type host. Macropus eugenii. Localities: captive, Flinders University, Adelaide*, S.A.; Kangaroo Island, S.A.; captives, Adelaide Zoo, and Gilles Plains Animal Resource Centre, Adelaide, S.A.

Other hosts. M. panyi, whip-tailed wallaby. Measurements from one animal, Table 5. Localities: captive, Adelaide Zoo, Adelaide*, S.A.; 9 1 km north of Rockhampton, Qld.; “Inkerman”, via Home Hill, Qld.

M. rufogriseus. Measurements from one animal, Table 5. Locality: Launceston*, Tas.

M. agilis, agile wallaby. Measurements from one animal, Table 5. Localities: captive, Adelaide Zoo, Adelaide*, S.A.

M. dorsalis. Measurements from one animal, Table 5. Locality: captive, Pallarenda, Townsville*, Qld.

Comment. Though most specimens contained an oocyst residuum, two/five infected M. eugenii from different localities, and the single M. agilis infected had no oocyst residuum. E. prionotemni and E. toganmainensis occurred together in only three M. eugenii, where they were clearly separate. The species epithet is based on the host distribution, which includes most of the ‘brush wallaby’ species formerly included in the putative subgenus Prionotemnus (Bartholomai, 1975). Oocysts in faeces; endogenous stages undescribed.

Eimeria wi1cannien.G Mykytowycz, 1964 Figs. 7.29, 30

Description. Oocyst approximately ellipsoidal, slightly pointed at one end, or slightly ovoid; 25 measured from two animals, Table 6; double oocyst wall; outer smooth, clear, colourless, 0.8-l .O ,u m thick; inner clear, 0.4 pm thick; thinning of outer oocyst wall, and protrusion of inner wall at more pointed end of oocyst, but micropyle often equivocal; no polar granule or oocyst residuum; four ellipsoidal sporocysts, slightly pointed and thickened toward one end, but no clear-cut Stieda body; no substieda body; two sporozoites nearly filling sporocyst; refractile globule 3.2-4.0 pm diameter; sporocyst residuum loose aggregate of small granules, usually scattered at equator of sporocyst.

Diagnosis. E. wilcanniensis is separated from oocysts with overlapping size range, other than E. mykytowyczi, new species, on the basis of absence of a Stieda/substieda body. It is differentiated from E. mykytowyczi on the basis of oocyst shape and sporocyst size.

Type host. Macropus rufis. Localities: Yathong Nature Reserve*, near Mount Hope, N.S.W.; ‘Avenel’, via Broken Hill, N.S.W.

Other hosts. M. giganteus. Twenty measured from two animals, Table 6. Locality: Yathong Nature Reserve*, near Mount Hope, N.S.W.


- IOym

FIGS. 29-39. Photographs of oocysts of various Eimeria spp. from kangaroos and wallabies of the genus Macropus. Scale = 10 pm, all figures to same scale.

FIGS. 29,30. E. wilcanniensis: FIG. 29, from M. rufus, type host; FIG. 30, from M. robustus.

FIG. 31. E. mykytowyczi from M. agilis.

FIGS. 32-34. E. macropodis: FIG. 32, from M. rufogriseus, type host; FIG. 33, from M. parma; FIG. 34, from M. fuiiginosus.

FIG. 35. E. marsupialium from M. giganteus.

FIG. 36. E. yathongensis from M. fuliginosus.

FIG. 37. E. gungahlinensis from M. fuliginosus.

FIG. 38. E. parma from M. parma.

FIG. 39. E. desmaresti from M. rufogriseus.

M. fuliginosus. Forty measured from two animals measured from one animal, Table 6. Locality: from two localities, Table 6. Localities: Second “Nonning”*, via Port Augusta, S.A. Valley*, S.A.; captive, Adelaide Zoo, Adelaide*, S.A.; Comments. This species was originally described Para Wirra Conservation Park, near Kersbrook, S.A.; by Mykytowycz (1964) from the red kangaroo Kangaroo Island, S.A. M. rufus. The original description fails to refer to the

M. robustus, common wallaroo, or euro. Twenty presence or absence of a Stieda/substieda body, and

TABLE ~-DIMENSIONS

(urn

) OF OOCYSTS AND SPOROCYSTS OF E

imer

ia w

iica

nnie

nsis

IN M

ucro

pus

rufi

s,

M. f

digi

nosu

s,

M.

giga

nteu

s AND

M.

robu

stus

Len

gth

Ooc

yst

Wid

th

Len

gth

Spor

ocys

t

Wid

th

Hos

t

M.

rufu

s

(typ

e)

Myk

ytow

ycz

(196

4)

M. f

ulig

iflo

sis

M. g

igan

teus

M

. ro

bust

us

Pool

ed

(ori

gina

l)

Isd

(5%

)

n*

xt

S.D.

Range

,?t

S.D.

Range

Xi

S.D.

Range

it

S.D.

Range

25

32.2

1.

45

30.0

-34.

0 20

.3

0.95

18

.0-2

1.6

13.1

0.

60

12.0

-14.

0 9.

8 0.

47

9.0-

11.0

100

31.8

-

28.0

-37.

8 20

.3

- 18

.4-2

3.6

11.9

-

9.5-

14.3

9.

3 -

8.2-

10.9

40

34.8

1.

71

31.2

-38.

4 20

.4

0.57

19

.2-2

1.6

14.3

0.

75

12.8

-15.

2 9.

3 0.

41

8.8-

10.4

20

35

.4

1.83

31

.2-3

8.0

21.0

0.

96

19.2

-23.

0 12

.9

0.75

11

.2-1

4.4

9.7

0.44

8.

8-10

.4

20

32.8

1.

42

31.2

-36.

0 21

.4

0.45

20

.8-2

2.4

11.4

0.

54

10.4

-12.

0 9.

0 0.

57

7.6-

10.4

105

33.9

2.

04

30.0

-38.

4 20

.7

0.85

18

.0-2

3.0

13.2

1.

25

10.4

-15.

2 9.

4 0.

54

7.6-

11.0

2.0

0.7

1.6

0.5

*Num

ber

of o

ocys

ts m

easu

red.

SM

ean

dim

ensi

on.

TABLE ~-DIMENSIONS (fim)oF OOCYSTSAND SPOROCYSTSOF E

imer

iam

ykyt

owyc

izi

IN M

acro

pusa

ntii

opin

us,M

.agi

lis

AND h% p

arty

i

Ooc

yst

Spor

ocys

t

Hos

t

Len

gth

Wid

th

n*

ft S.D.

Range

X?

S.D.

Range

M. a

gili

s

(typ

e)

M.

anti

lopi

nus

M. p

arry

i

Pool

ed

Isd

15%

)

21

31.3

1.

17

29.6

-33.

6 17

.5

0.46

16

.8-1

8.4

20

28 .

6 1.

86

25.6

-33.

6 15

.9

0.26

1X

-16.

0 29

32

.7

2.01

28

.0-3

6.5

18.6

0.

89

16.0

-20.

0

70

31.1

2.

43

25.6

-36.

5 17

.5

1.29

15

.2-2

0.0

3.3

2.2

*Num

ber

of o

ocys

ts m

easu

red.

tM

ean

dim

ensi

on.

Len

gth

Wid

th

.ut

S.D

. R

ange

it

S.D.

Ran

ge

10.5

0.

57

9.6-

12.0

7.

8 0.

43

7.2-

8.8

10.5

0.

50

9.6-

11.2

7.

7 0.

36

6.8-

8.0

11.3

0.

69

10.3

-12.

4 8.

2 0.

43

7.2-

9.2

10.9

0.

72

9.6-

12.4

7.

9 0.

47

6.8-

9.2

0.8

0.5

,. .

_-_

.__.

, _,_

.i., _I

., ,.-

_- --

-_ _

__- --

r.

_I__

“(”

-....

~ .

,___

=,_

. __

___.

_g

_,__

,,,_

,_ ._

.. __

_ I_

,. __

_ .._

. __ . . .

,,.


the illustrations are inadequate in this regard. We have redescribed the species based on material from the type host. Mykytowycz (1964) did not recognize E. wilcanniensis in grey kangaroos, but we found it rarely (two/82) in M. giganteus at Mount Hope, N.S.W., though not in 92 M.filiginosus from that locality. However, it was present in seven/43 M. fuliginosus from Kangaroo Island and mainland S.A., where it also occurred in M. robustus.

Eimeria myky#owyczi new species Figs. 8,3 1

Description. Oocyst irregular, elongate ellipsoid/ ovoid, often flattened on one or both sides in profile; 2 1 measured from one animal, Table 7; double oocyst wall; outer layer smooth, clear, colourless, 0.8- 1.0 pm thick; inner clear, 0.4 ,~rn thick; thinning of outer wall and protrusion of inner wall at more pointed end of oocyst, but micropyle equivocal; polar granule variably present; no oocyst residuum; four ellipsoidal sporocysts, slightly pointed and thickened at one end, but no clear-cut Stieda body; no substieda body; two sporozoites nearly filling sporocyst; refractile globule 3.2-4.0 pm diameter; sporocyst residuum loose aggregate of granules usually scattered about equator of sporocyst.

Diagnosis. E. mykytowyczi is separated from E. wilcanniensis on the basis of consistently irregular oocyst shape, and a combination of smaller oocyst and sporocyst dimensions, as well as presumed host specificity. It is separated from other coccidia of Mucropus spp. with overlapping oocyst size range on the basis of lacking a Stieda/substieda body. E. obendorfi of the Tasmanian pademelon (Barker, O’Callaghan & Beveridge, 1988b) is smaller, and lacks a micropyle. E. godmani of rock wallabies (Barker, O’Callaghan, Beveridge & Close, 1988) is similar, but has a thicker, tan oocyst wall.

Type host. Macropus agilis. Locality: ‘Tipperary’*, N.T.

Other hosts. M. antilopinus. Twenty measured from one animal, Table 7. Locality: captive, Pallar- enda, Townsville*, Qld.

M. parryi. Twenty-nine measured from one animal, Table 7. Locality: captive, Adelaide Zoo, Adelaide*, S.A.

Comments. This species closely resembles E. wilcunniensis, from which it is differentiated on morphometric grounds, in the absence of information on host specificity. Named in honour of Dr Roman Mykytowycz, formerly of the C.S.I.R.O. Division of Wildlife Research, who first studied coccidia in kangaroos systematically. Oocysts in faeces; endogenous stages undescribed.

Eimeria parryi new species Figs. 9, 28

Description. Oocyst elongate ellipsoid, 2 1 measured from one animal, 28.7 + 1.20 (25.6- 30.4) X 13.1 +- 0.54 (12.0-14.4)pm; double oocyst

wall; outer smooth, clear, colourless, 1.0 pm thick; no micropyle; polar granule present; four elongate ellipsoidal sporocysts, 12.3 k 0.85 (11.2- 14.4) X 6.0 k 0.39 (5.6-6.4) pm; tapering slightly to inconspicuous Stieda body; no substieda body; two sporozoites, nearly filling sporocyst; refractile globule 3.2 X 2.4 pm; sporocyst residuum a few granules up to 1.6 pm diameter scattered in sporocyst.

Diagnosis. This species is separated from others known in Macropus spp., except E. parma, new species, in M. parma, by possession of a sporocyst with a length : width ratio of about 2 : 1. The oocyst of E. parryi is larger than that of E. parma. The oocyst is narrower, and mean sporocyst length is shorter than E. sharmani in rock wallabies (Barker, O’Callaghan, Beveridge & Close, 1988) which, coupled with the generic difference in hosts is considered justification for separation of the species.

Type host. Macropus parryi. Localities: captive, Adelaide Zoo, Adelaide*, S.A.; 78 km north of Rockhampton, Qld.

Comments. Named for the host. Oocysts in faeces; endogenous stages unknown.

Eimeria macropodb Wenyon & Scott, 1925 Synonyms E. fausti Yakimoff & Matschoulsky. 1936. New

synonymy. E. cunnamullensis Mykytowycz, 1964. New synonymy.

E. purchasei Mykytowycz, 1964. New synonymy. Figs. 10,32-34

Description. Oocyst irregular, ovoid, ellipsoid, or cylindroid, often slightly asymmetric or flattened on one side, usually somewhat pointed at one end; 30 measured from four animals from three localities, Table 8; double oocyst wall; outer clear, colourless, smooth, 0.8 y m thick; inner clear, 0.4 pm thick; outer wall thinning at pointed end of oocyst, often with slight concave inward distortion of inner wall, but no clear- cut micropyle; polar granule present; no oocyst residuum; four roughly ellipsoidal sporocysts, slightly pointed at end with inconspicuous Stieda body; no substieda body; two sporozoites filling sporocyst, each with refractile globule 1.6-3.2 pm diameter; finely granular residuum scattered about equator of sporocyst.

Diagnosis. Shorter, narrower oocyst than E. wilcanniensis, E. mykytouyczi and the other larger species in Macropus spp.; no micropyle. Shorter sporocyst than E. parryi. More elongate, irregular shape than E. marsupialium. Generally larger, less regular shape than E. yathongensis, new species, which is consistently ovoid. Larger oocyst, and sporocyst than E. parma; larger oocyst, elongate shape compared to E. gungahfinensis. Larger than E. desmaresti. Small forms similar to E. petrogale in rock wallabies (Barker, O’Callaghan, Beveridge & Close, 1988); E. bicofor in the swamp wallaby; and E. Zumholtzi in Lumholtz’ tree kangaroo (Barker, O’Callaghan & Beveridge, 1988b), but considered separate species on basis of generic difference of

TA

BL

E ~

-DIM

EN

SIO

NS

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n) O

F O

OC

YST

S AN

D SPOROCYSTS OF E

imer

ia

mac

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pusg

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, M

. ful

igin

osus

, M

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ma,

M

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M.

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M

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, M

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2 M

. ru

fogr

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s

(typ

e)

M. g

igan

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M

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igin

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M

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fus

M.

irm

a M

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yi

M.

dors

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M

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ed

300

25.0

2.

52

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(5%

) 1.

6

50

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34

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62

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20

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9.3

*Num

ber

of o

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TABLE ~-DIMENSIONS

(urn

) O

F O

OC

YST

S AN

D SPOROCYSTS OF E

imer

ia

mar

supi

aliu

m

IN M

acro

pus

giga

nteu

s A

ND

M. f

ulig

inos

us

Hos

t

Ooc

yst

Spor

ocys

t

Len

gth

Wid

th

Len

gth

Wid

th

n*

Xi

S.D

. R

ange

xt

S.

D.

Ran

ge

it S.

D.

Ran

ge

it S.

D.

Ran

ge

M. g

igan

teus

(typ

e)

M. f

ulig

inos

us

M.

giga

nteu

r Y

akim

off

& M

atsc

houl

sky

1936

44

22.5

1.

93

19.2

-25.

9 16

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1.08

14

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8.2

10.8

0.

62

9.6-

11.9

6.

9 0.

55

5.6-

7.7

35

22.3

1.

54

19.2

-24.

8 16

.6

0.79

15

.2-1

8.4

11.0

0.

53

10.4

-12.

0 6.

8 0.

49

5.6-

7.4

77

22.2

-

20-2

6 18

.0

- 16

-20

- -

8-12

-

- 6-

8

Pool

ed

(ori

gina

l)

79

22.4

1.

76

19.2

-25.

9 16

.5

0.96

14

.4-1

8.4

10.9

0.

58

9.6-

12.0

6.

9 0.

53

5.6-

1.7

*Num

ber

of o

ocys

ts

mea

sure

d.

iMea

n di

men

sion

.


hosts. Large forms separated from E. obendo@ in Tasmanian pademelon (Barker, O’Callaghan 81 Beveridge, 1988b) by size and shape of sporocyst, and possession of Stieda body; from E. ringaroomaensis in the same host by smaller sporocyst size, and lack of substieda body.

Type host. Macropus rufogriseus. Localities: Connerville*, Launceston* and Bridgenorth, Tas.; Bondo*, N.S.W.

Other hosts. M. giganteus. Fifty measured from two animals, Table 8. Localities: Yathong Nature Reserve*, near Mount Hope, N.S.W.; Bondo, N.S.W.; Mitta Mitta and Yan Yean, Vie.; captives, Cleland Conservation Park, Adelaide, S.A.

M. fuliginosus. Thirty measured from four animals, Table 8. Localities: Yathong Nature Reserve*, N.S.W.; Second Valley*, Para Wirra Conservation Park*, Kangaroo Island, Bangham, S.A.; ‘Pine Plains’, via Patchewollock, Vie.

M. rufis. Forty measured from two animals, Table 8. Locality: Yathong Nature Reserve*, near Mount Hope, N.S.W.

M. irma, western brush wallaby. Twenty-five measured from three animals, Table 8. Locality: captives, Murdoch University, Murdoch*, W.A.

M. parryi. Thirty-three measured from three animals, Table 8. Localities: 78 km north of Rock- hampton*, Qld.; captive, Adelaide Zoo, Adelaide*, S.A.; 91 km north of Rockhampton, Qld.; ‘Inkerman’, via Home Hill, Qld.

M. dorsalis. Forty-two measured from 11 animals, Table 8. Localities: (all Qld.) ‘Harvest Home’*, via Charters Towers; Milman*; ‘Apis Creek’*, via Marlborough; Clements Creek*, north of ‘Apis Creek’; captive, Pallarenda, Townsville*.

M. eugenii. Twenty measured from five animals, Table 8. Localities: (all S.A.) Kangaroo Island*; captive, Flinders University and Adelaide Zoo, Adelaide*; captive, Kingston (Southeast).

M. parma, parma wallaby. Thirty measured from three animals, Table 8. Localities: captive, Macquarie University, Sydney*, N.S.W.; Adelaide Zoo, Adelaide*, S.A.

Comments. There was often wide variation in shape and size of oocysts having the characters described, within a particular host species and individual, and a broader range in oocyst size among host species (Table 8, Fig. 40). However, we were unable to confidently differentiate more than one species, on the basis of oocyst size, within any host. We conclude that the strong homology among the oocysts present in the various hosts of the genus Macropus, despite differences in mean oocyst length and other para- meters (Table 8) justifies lumping them as a single species, in the absence of experimental information on host specificity. Wenyon & Scott (1925) described, and Triffitt (1926) reiterated the description and illustrated, an asymmetric oocyst in M. rufogriseus with a wide range of dimensions, and other character- istics (‘weak’ micropyle, ‘slight’ Stieda body), similar

to those we found in the same host. We conclude, therefore, that E. macropodis Wenyon & Scott, 1925 is valid. E.fausti Yakimoff & Matschoulsky, 1936, described from only a ‘few’ oocysts in M. giganteus, falls within the size and shape range for this species. No micropyle or Stieda body were described. We conclude that E. fausti is compatible with the species we redescribe, and assuming no host specificity, we synonymize E. fausti Yakimof & Matschoulsky, 1936, with E. macropodis Wenyon & Scott, 1925. Mykytowycz (1964) described two species, E. cunnamullensis and E. purchasei from grey kangaroos ‘M. canguru’, apparently separating them on the basis of (overlapping) overall oocyst size and shape. Work- ing with material from his type locality for E. purchasei, and from grey kangaroos of known species, we do not feel confident in separating the two species of coccidia. Micropyles were described in E. purchasei and E. CunnamuIlensis; though the outer oocyst wall thins at the apex, there does not appear to be a clear- cut micropyle in our material. Mykytowycz’s (1964) illustrations are small, but we do not recognize a micropyle in them, though it is easy to understand the interpretation that one is present. On the basis that the morphologic characters and size of oocysts resembling E. cunnamullensis Mykytowycz, 1964, and E. purchasei, Mykytowycz, 1964, are similar to E. macropodis Wenyon & Scott, 1925, and assuming no host specificity, we synonymize the former two species with E. macropodis. E. macropodis is the most prevalent coccidian within each host species, and has the broadest host range among Mucropus of any of the coccidia encountered. The only hosts in which it was not found (Table 1) inhabit xeric habitats and had a low overall prevalence of infection with coccidia (M. robustus), or were only available in small numbers (M. antilopinus, M. agilis). Mykytowycz’s (1964) failure to observe the species in red kangaroos may relate to this, since we found it only at low prevalence in that host (Table 1). Further investigation may demonstrate that E. macropodis has a universal host range among Macropus. Oocysts in faeces; endogenous stages undescribed.

Eimeria marsupialium Yakimoff & Matschoulsky, 1936 Figs. 11.35

Description. Oocyst subspherical to ellipsoidal or ovoid; 44 measured from seven animals from four localities, Table 9; double oocyst wall; outer smooth, clear, colourless, 1.0 pm thick; inner clear, 0.4 ,D m thick; no micropyle; no polar granule or oocyst residuum; four elongate ellipsoidal-ovoid sporocysts; inconspicuous Stieda body present; no substieda body; two sporozoites filling sporocyst; refractile globule 2.4-3.2 ,~rn diameter; sporocyst residuum irregular granular aggregate scattered throughout sporocyst.

Diagnosis. Separated from E. macropodis on the basis of shorter oocyst, higher oocyst width:length ratio, greater sporocyst length, relative symmetry of


17 - .a rufogrisaus

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OOCVST LENGTH (pm)

FIG. 40. Limits of oocyst dimensions of Eimeria macropodis measured from M. rufogriseus, the type host, and from M. parma, M. dorsalis, M. parryi and M. irma. The most extreme dimensions of oocysts of this species measured from any host (see Table 8) are illustrated in this figure. Large symbols mark the mean dimensions of oocysts measured from each host

species.

oocyst, and lack of any suggestion of micropyle. E. parryi and E. parma have sporocyst length:width ratios of about 2 : 1. It is longer, has larger sporocysts and lacks the surface mamillation of E. gungahlinensis.

Type host. Macropus giganteus. Localities: Bondo*, and Yathong Nature Reserve, near Mount Hope, N.S.W.; Mitta Mitta*, and Yan Yean*, Vie.; captive, Cleland Conservation Park, Adelaide*, S.A.

Other host. M. fuliginosus. Thirty-five measured from two animals, Table 9. Localities: Yathong Nature Reserve*, near Mount Hope, N.S.W.; Bangham*, Second Valley, Para Wirra Conservation Park, near Kersbrook, and Kangaroo Island, S.A.; ‘Pine Plains’, via Patchewollock, Vie.

Comments. Yakimoff & Matschoulsky (1936) described this species from M. giganteus, in a zoo, and illustrated ellipsoidal-ovoid oocysts, without a micro- Pyle, of approximately the same size as those we report here. They offer no mean sporocyst dimensions, with lower minimum sporocyst length and width limits than we observed, and indicated absence of a Stieda (Schneider) body. The Stieda body we observed is not prominent, and despite the incon- sistencies noted, we conclude that we are dealing with E. marsupialium Yakimoff & Matschoulsky, 1936.

Eimeria yathongensis new species Figs. 12,36

Description. Oocyst consistently ovoid; 30 measured from two animals from two localities, Table 10; double oocyst wall; outer smooth, clear, colourless, 0.6 ,~rn thick; inner clear, colourless, 0.3 pm; no micropyle; no polar granule; no oocyst residuum; four approximately ellipsoidal sporocysts, slightly pointed toward end with inconspicuous Stieda body; no substieda body; two sporozoites nearly filling sporocyst; refractile globule 3.2 pm diameter; sporocyst residuum a loose aggregate of small granules scattered throughout sporocyst.

Diagnosis. Differentiated on the basis of combination of oocyst size and shape, and greater sporocyst length, from E. macropodis in grey kangaroos. Thinner, less refractile oocyst wall, lower width:length ratio and more consistent shape than E. marsupialium. Larger oocyst and sporocyst size, different shape, smooth surface in comparison with E. gungahlinensis. Different oocyst and sporocyst shape from E. parma.

Type host. Macropus fuliginosus. Localities: Yathong Nature Reserve*, near Mount Hope, N.S.W.; Bangham*, Second Valley, Kangaroo Island, Para Wirra Conservation Park, S.A.

TA

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(typ

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30

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6.6

20

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99

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10.6

0.

36

10.4

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2 5.

9 0.

42

5.6-

6.6

50

21.3

1.

02

19.0

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59

10.0

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*Num

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TA

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s

Len

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gth

Spor

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t

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t

M. f

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us

(typ

e)

M. g

igan

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‘ M. ca

ngur

u’

Myk

ytow

ycz

(196

4)

Pool

ed

(ori

gina

l)

n*

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S.D

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70

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7.0-

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7.7

50

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50

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8.4

8.8

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5

120

18.4

1.

93

13.0

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12

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5.

0-7.

7

*Num

ber

of o

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d.

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Other host. M. giganteus. Twenty oocysts measured from two animals, Table 10. Locality: Yathong Nature Reserve*, near Mount Hope, N.S.W.

Comments. This species was always present in relatively small numbers, in animals infected with one or the other, or both, of E. macropodis or E. marsupial&m, never alone. It was much more prevalent in M. fuliginosus than in M. giganteus, at Mount Hope. The specific name is based on the locality where it was present in both species of grey kangaroos. Oocysts in faeces; endogenous stages unknown.

Eimeria parma new species Figs. 13,38

Description. Oocyst irregular, blunt ellipsoid, occasionally more pointed at one end, or flattened on one side; 12 measured from one animal, 20.7 & 0.75 (20.0-21.6) X 11.6 kO.34 (11.2-12.0)pm; double oocyst wall; outer smooth, clear, colourless, 0.8 pm thick; no micropyle; inner clear, 0.4 ,um thick; no polar granule; no oocyst residuum; four sporocysts, 11.7kO.64 (11.2-12.4) X 5.1 kO.31 (4.8-5.6)pmm; inconspicuous Stieda body; no substieda body; two sporozoites, not filling sporocyst; refractile globule 2.4 pm diameter; sporocyst residuum an aggregate of small granules at equator of sporocyst.

Diagnosis. This species resembles E. panyi in having a sporocyst with a 1ength:width ratio of about 2: 1; however the oocyst is significantly shorter. It is similar to E. volckertzooni in the quokka (Barker, O’Callaghan & Beveridge, 1988b), but the generic and geographic/ecologic separation of the hosts is considered to justify separate species.

Type host. Macropus parma. Locality: captive, Adelaide Zoo, Adelaide*, S.A.

Comments. Oocyst in faeces; endogenous stages unknown.

Eimeriagungahlinensis Mykytowycz, 1964 Figs. 14,37

Description. Oocyst spherical-subspherical, 70 measured from two animals from two localities, Table 11; double oocyst wall; outer 0.8 pm, slightly mamillate, clear, colourless; no micropyle; inner 0.4 pm, clear, colourless; no oocyst residuum; polar granule 1.6 pm diameter; four ovoid-ellipsoidal sporocysts; Stieda body present; no substieda body; two sporozoites filling sporocyst; refractile globule 1.6-2.4 pm diameter; sporocyst residuum a granular aggregate up to 4.8 pm diameter.

Diagnosis. This species is distinguished from other coccidia in Macropus by its small size and spherical- subspherical shape, in combination with its mamillate oocyst surface. The sporocyst is smaller than that of E. marsupialium, with which species oocyst size may overlap.

Type host. Macropus fuliginosus. Localities: Yathong Nature Reserve*, near Mount Hope, N.S.W.; ‘Pine Plains’*, via Patchewollock, Vie.; Kangaroo Island, Second Valley, Para Wirra, near Kersbrook, S.A.

Other host. M. giganteus. Fifty measured from one animal, Table 11. Locality: Yan Yean*, Vie.

Comments. This species is redescribed from M. fuliginosus at Mykytowycz’s (1964) type locality, Mount Hope, N.S.W., and from northwestern Victoria. Though it is relatively common in M. filiginosus, it is rare in M. giganteus (Table 1). Oocysts in faeces; endogenous stages unknown.

Eimeria desmaresti new species Figs. 15,39

Description. Oocyst irregular, blunt ellipsoid, often flattened on one side; 22 measured from one animal, 16.8 + 1.05 (14.6-18.4) X 10.3 + 0.55 (9.7- 1 1.4 pm; double oocyst wall; outer clear, colourless, smooth, 0.8 pm thick; inner clear, 0.4 pm thick; no micropyle; polar granule present; no oocyst residuum; four approximately ellipsoidal sporocysts, 7.5 * 0.73 (6.8-9.7) X 4.5 + 0.46 (3.9-4.9) pm; more pointed at end with obvious Stieda body; no substieda body; two sporozoites, filling sporocyst; refractile globule 1.6 iurn diameter; sporocyst residuum a few granules scattered at equator of sporocyst.

Diagnosis. Differentiated on basis of smaller size, shape, and more prominent Stieda body from E. macropodis and E. marsupialium . Differentiated from E. gungahlinensis on the basis of shape and smooth oocyst surface, and from E. parma on the basis of small oocyst and sporocyst size. It overlaps the oocyst length of E. quokka in the quokka (Barker, O’Callaghan & Beveridge, 1988b), though with a smaller mean dimension; the two species are considered justified based on the generic and geographic/ ecologic separation of the hosts.

Type host. Macropus rufogriseus. Localities: Comrerville*, Bridgenorth, and Launceston, Tas.

Comments. Named in honour of Anselme Gaetan Desmarest, French zoologist and veterinarian who described several species of wallabies and kangaroos, including the type host of E. desmaresti.

DISCUSSION

We recognized all but two of the Eimeria spp. previously described from kangaroos and wallabies, though, for reasons presented above, we synonymized E. fausti Yakimoff & Matschoulsky, 1936; E. cunnamullensis Mykytowycz, 1964 and E. purchasei Mykytowycz, 1964 with E. macropodis Wenyon & Scott, 1925.

We were unable to recognize oocysts correspond- ing with E. kogoni Mykytowycz, 1964, despite exam- ining hundreds of samples from grey kangaroos from the type locality (Mount Hope, N.S.W.), and else- where. In the early stages of this work, prior to the recognition of the species distribution of E. wilcanniensis, we mistakenly diagnosed oocysts of that species, in grey kangaroos, as E. kogoni. We found that E. wilcanniensis in the grey kangaroos is larger than the same species, as described by Mykytowycz (1964) in the red kangaroo (Table 6). Without

TABLE 12-DIAGNOSTICFEATURESOFOOCYSTS

OF E

imer

ia

SP

P.IN

KANGAROOSAND

WALLABIES OFTHE GENUS

Mac

ropu

s,

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knowledge of the criteria for his separation of E. kogoni and E. wilcanniensis, and whether Stieda and substieda bodies are present or not, which Mykytowycz (1964) did not describe or illustrate adequately, we do not know if E. kogoni is a synonym of E. wilcanniensis. We regard it as a species inquirenda.

Similarly, we did not find oocysts in M. rufi(s, or in any other species, which corresponded with E. rufusi Prasad, 1960. The description resembles E. marsupialium most closely, but the sporocyst size is smaller than in that species, which, as well, we did not encounter in red kangaroos. Hence, we also regard E. rufusi Prasad, 1960 as a species inquirenda.

Though we attempted to examine large numbers of individuals within each host species, we were not always successful (Table 1). Undoubtedly, the suites of coccidia infecting hosts with small sample sizes in our study will be expanded by further investigation. Species sampled almost exclusively in localities with xeric environments (M. r&s, M. robustus) had a low prevalence of coccidial infection, which may have influenced the probability of encountering the full range of coccidial species parasitizing those hosts. Within the hosts with the greatest coccidial species diversity (M. giganteus and M. fuliginosus) populations from drier areas (e.g. Mount Hope) had lower overall prevalence of infection and fewer species/ infected animal.

Without experimental attempts to cross-infect hosts with oocysts derived from other host species, the biologic validity of the species of Eimeria described from more than one host in the genus Macropus is unknown. We have elected to lump homologous oocysts from congeneric hosts as single species of Eimeria, a conservative practice which reflects their similarity, but is amenable to revision as more information becomes available. It also serves to emphasize patterns of host-parasite association.

The earlier impression, that little homology exists among the Eimeria of various members of the genus Macropus, does not hold. E. macropodis infects nine species, and E. toganmainensis infects five species, in both cases encompassing all three subgenera in Macropus (Table 1). E. jlindersi and E. mykytowyczi each infect three species, encompassing the subgenera Osphranter and Notamacropus; E. wilcanniensis infects four species, encompassing Osphranter and Macropus (Table 1). E. hestermani infects five species, in Macropus and Notamacropus (Table 1). E. marsupialium, E. yathongensis and E. gungahlinensis are all found in, and limited to, species of the subgenus Macropus (Table 1). In addition, E. prionotemni is limited to, but widespread among, species within the subgenus Notamacropus.

On the basis of these patterns of host-parasite association, we suggest that E. macropodis may have a long-established relationship with Macropus, per- sisting in most, if not all, species as they radiated. The presence of Eimeria spp. resembling small forms of

E. macropodis in Wallabia bicolor (E. bicolor, Barker, O’Callaghan & Beveridge, 1988b), in Petro- gale spp. (E. petrogale , Barker, O’Callaghan, Beveridge & Close, 1988) and in Dendrolagus lumholtzi (E. lumholtzi, Barker, O’Callaghan & Beveridge, 1988b) further may suggest a long association of a species group of Eimeria with macropodid marsupials. This may also explain the presence of E. macropodis in all three subgenera of Macropus, if, as Richardson & McDermid (1978) propose, the genus is paraphyletic, M. giganteus and M. fuliginosus, allied with Wallabia, standing apart from the other Macropus spp.

E. desmaresti is similar to E. quokka in Setonix brachyurus (Barker, O’Callaghan & Beveridge, 1988b), and E. prionotemni closely resembles E. inornata in Petrogale spp. (Barker, O’Callaghan, Beveridge & Close, 1988) and E. aepyptymni in Aepyprymnus rufescens (Barker, O’Callaghan & Beveridge, 1988a). The latter homology is the only one recognized between coccidia in members of the Potoroidea and Macropodidae. E. panyi and E. parma share the characteristic of having a sporocyst possessing a high ( 22: 1) 1ength:width ratio with E. sharmani in Petrogale spp., and with E. volckertzooni in Setonix.

Though adding some weight to the notion that it may be possible to recognize lineages of homologous Eimeria spp. among the Macropodoidea which may be coevolving with their hosts, these observations contribute little to clarifying our understanding of the many-branched phylogenetic ‘tree’ of genera reflect- ing the recent radiation of the kangaroos and wallabies (Richardson & McDermid, 1978).

Within Petrogale, distinctive Eimeria species, or suites of species, seemed to mark species groups of hosts, as defined by other taxonomic approaches (Barker, O’Callaghan, Beveridge & Close, 1988). The same may occur to some extent within Macropus, in that E.prionotemni appears limited to wallabies in the subgenus Notamacropus, while a suite of three Eimeria species (E. marsupialium, E. yathongensis and E. gungahlinensis) is limited to the two closely- related species of kangaroos comprising the subgenus Macropus. The remaining host-parasite associations were not compatible with any particular hypothesis regarding phylogenetic relationships within the genus Macropus (see Richardson & McDermid, 1978; Archer, 1984).

Hence, while examination for homology of Eimeria spp. may have very limited use as an approach to understanding the phylogenetic relationships among the Macropodidae at the generic level, it may have some application to studies at the species level, especially within complex genera. Since many of the genera in the Macropododoidea are monospecific, and most others, with the exception of Petrogale and Macropus, are relatively small, opportunities to test this hypothesis further within the Macropodoidea are limited. Our observation of host specificity, for at least

262 I. K. BARKER, M. G. O’CALLAGHAN and I. BEVERIDG~

some species of coccidia, at the generic level, or speculatively (though we have not followed this possibility in assigning species names), at a higher taxonomic level in the Macropodoidea, is consistent with a pattern observed by Wash et al. (1985) in other mammalian hosts with high levels of karyotypic variation and polymorphism.

The origin of coccidia infecting Australian marsupials is cause for speculation, if the entire fauna is derived from a common ancestor which immigrated to Australia from South America, via Antarctica, as Gondwanaland broke up (Archer, 1984). Coccidia have yet to be described in the Dasyuridae, though gametocytes, which may be those of Eimeria, have been recognized in sections of the intestine of Ante- chinus stuartii (Barker, Beveridge, Bradley & Lee, 1978). They are found in the Perameloidea (our unpublished observations), the Vombatidae (Barker, Munday & Presidente, 1979), many members of the Phalangeroidea (Presidente, 1984; Speare, Haffenden, Daniels, Thomas 81 Seawright, 1984; our unpublished observations), as well as the Macro- podoidea. Assuming that they radiated from the common ancestor, it seems probable that some members of the Dasyuridae will be found to be infected by Eimeria, unless this coccidian parasite was ‘captured’ by an ancestral marsupial from another vertebrate class, following an early divergence by the Dasyuridae. Certainly, Eimeria has successfully para- sitized the echidna (Barker, Beveridge & Munday, 1985), and the platypus (Barker, Whittington & McCall, unpublished), members of the Monotremata, which appears to have evolved entirely within Australia (Augee, 1984), and which presumably adopted a coccidian fauna by capture.

Acknowledgements-We thank Dr David Spratt, Dr Richard Speare, Dr Dale McCullogh, Dr Rob Close, Mr Keith Dempster, Dr Barry Munday, Dr David Obendorf, Dr Jon Dunsmore, Mr Peter Johnson and many others for their assistance in obtaining specimens. Animals were collected under permits from appropriate fauna authorities. Liz Moore and Jeanette Clarke provided laboratory support. Mark Fitzgerald of the IMVS, Adelaide, developed and printed photographs. Particular thanks are due to Dr Jack Arundel, Department of Veterinary Paraclinical Sciences, University of Melbourne, who fostered the interest of I.K.B. and LB. in studies on Australian wildlife, and provided support for the early phase of this work, which has been carried on intermittentlv since 197 1. Specimen collection was facilitated by grants io LB. from the Australian Research Grants Scheme. This work was supported in part by an Inter- national Collaborative Research Grant from the Natural Sciences and Engineering Research Council of Canada. awarded to I.K.B.

REFERENCES

ARCHER M. 1984. The Australian marsupial evolution. In: Vertebrate Zoogeography and Evolution in Australasia (Animals in Space and Time) (Edited by ARCHER M. & CLAYTON G.), pp. 633-808. Hesperian Press, Carlisle, Western Australia.

AUCEE M. 1984. Quills and bills: the curious problem of monotreme zoogeography. In: Vertebrate Zoogeography and Evolution in Australasia (Animals in Space and Time) (Edited by ARCHER M. & CLAYTON G.), pp. 567- 570. Hesperian Press, Carlisle, Western Australia.

BARKER I. K., HARRIGAN K. E. & DEMPSTER J. K. 1972. Coccidiosis in wild grey kangaroos. International Journal for Parasitology 2: 187-l 92.

BARKER I. K., BEVERIDGE I., BRADLEY A. J. & LEE A. K. 1978. Observations on spontaneous stress-related mortality among males of the dasyurid marsupial Ante- chinus stuartii Macleay. Australian Journal of Zoology 26: 435-447.

BARKER I. K., MUNDAY B. M. & PRESIDENTE P. J. A. 1979. Coccidia of wombats: correction of host-parasite relationships. Eimeria wombati (Gilruth and Bull, 1912) comb.nov. and Eimeria ursini Supperer, 1957 from the hairy-nosed wombat and Eimeria arundeli sp. n. from the common wombat. Journal of Parasitology 65: 45 1-456.

BARKER I. K., BEVERIDGE I. & MUNDAY B. L. 1985. Coccidia (Eimeria tachyglossi n. sp., E. echidnae n. sp., and Octosporella hystrix n. sp.) in the echidna, Tachyglossus aculeatus (Monotremata:Tachyglossidae). Journal ofProtozoology 32: 523-525.

BARKER I. K., O’CALLAGHAN M. G. & BEVERIDGE I. 1988a. Eimeria spp. (Apicomplexa:Eimeriidae) parasitic in the rat-kangaroos Hypsiprymnodon moschatus, Potorous tridactylus, Aepyprymnus rufescens and Beitongia gai- mardi (Marsupialia:Potoroidea). International Journal for Parasitology 18: 947-953.

BARKER I. K., O’CALLAGHAN M. G. & BEVERIDGE I. 1988b. Eimeria spp. (Apicomplexa:Eimeriidae) parasitic in wallabies and kangaroos of the genera Setonix, Thylo.pale, Wallabia, Lagorchestes and- Dendrolagus (Mar&pialia:Macropodidae), International Journal for Parasito- logy 18: 955-962.

BARKER I. K., O’CALLAGHAN M. G., BEVERIDGE I. &CLOSE R. L. 1988. Host-parasite associations of Eimeria spp. (Apicomplexa:Eimeriidae) in rock wallabies, Pefrogale spp. (MarsupiaIia:Macropodidae). International Journal forParasitology 18: 353-363.

BARTHOLOMAI A. 1975. The genus Macropus Shaw (Marsupialia:Macropodidae) in the upper Cainozoic deposits of Queensland. Memoirs of the Queensland Museum 17: 195-235.

BEVERIDCE 1. 1986. Coevolutionary relationships of the helminth parasites of Australian marsupials. In: Coevolu- tion and Systematics (Edited by STONE A. R. & HAWKSWORTH D. L.), pp. 93-l 17. Clarendon Press, Oxford.

DAWSON L. & FLANNERY T. 1985. Taxonomic and phylogenetic status of living and fossil kangaroos and wallabies of the genus Macropus Shaw (Macropodidae: Marsupi- alia), with a new subgeneric name for the larger wallabies. Australian Journal ofZoology 33: 473-498.

DUSZYNSKI D. W. & MOORE D. W. 1986. Coccidian parasites (Apicomplexa:Eimeriidae) from insectivores- II. Six new species from Japanese shrew moles (Talpidae). JournalofProtozoology 33: 276-281.

FLANNERY T. 1984. Kangaroos: 15 million years of Australian bounders. In: Vertebrate Zoogeography and Evolution in Australasia (Animals in Space and Time) (Edited by ARCHER M. & CLAYTON G.), pp. 817-835. Hesperian Press, Carlisle, Western Australia.

KIRSCH J. A. W. & POOLE W. E. 1972. Taxonomy and distribution of the grey kangaroos, Macropus giganteus Shaw and Macropus fuliginosus (Desmarest), and


their subsnecies (Marsunialia:Macropodidae). Australian Gliders (Edited by SMITH A. P. & HUME I. D.), pp. 283- Journal 0fZoolo‘gy 20: 3 1 S-339. _ ’ 302. Australian Mammal Society, Sydney. ’ _ _

KIRSCH J. A. W. 1977. The comoarative serologv of Marsu- STRAHAN R. 1983. The Cornvlete Book of Australian pialia, and a classification of ‘the marsupials: Australian Mammals, pp. 207-223. An&s and Robertson, Sydney. Journal of Zoology, Supplementary Series No. 152: l- TRIFFITT M. J. 1926. Some sporozoan parasites found in the 152. intestinal wall of Bennett’s Wallaby (Macropus benneni).

MYKYTOWYCZ R. 1964. Coccidia in wild populations of the Protozoology 2: 31-46. red kangaroo Megaleia rufa (Desmarest) and the grey VANCE T. L. &DUSZYNSKI D. W. 1985. Coccidianparasites kangaroo Macropus canguru (Miiller). Parasitology 54: (Apicomplexa:Eimeriidae) of Microrus spp. (Roden- 105-115. tia:Arvicolidae) from the United States, Mexico, and

PELLERDY L. P. 1974. Coccidia and Coccidiosis, 2nd Edn, Japan, with descriptions of five new species. Journal of pp. 378-381. Paul Parey, Berlin. Parasitology 71: 302-311.

PRASAD H. 1960. A new species of coccidia from the red WASH C.D., DIJSZYNSKI D. W. & YATES T. L. 1985. kangaroo Macropus rufus Mamm. Zeitschrif jYir Parasit- Eimerians from different karyotypes of the Japanese enkunde 20: 385-389. wood mouse (Apodemus spp.), with descriptions of two

PRESIDENTE P. J. A. 1984. Parasites and diseases of brush- new species and a redescription of Eimeria montgomeryae tail possums (Trichosurus spp.): occurrence and signific- Lewis and Ball, 1983. Journal of Parasitology 71: SOS- ante: In: Possums and Gliders (Edited by SMITH A. P. & 814. HUME 1. D.), pp. 171-190. Australian Mammal Society, WENYON C. M. & SCOTT H. H. 1925. Exhibitions of Sydney. sections of the small intestine of a Bennett’s wallaby

RICHARDSON B. J. & MCDERMID E. M. 1978.Acomparison (Macropus bennetti) containing various parasites. Trans- of relationships within the Macropodidae as determined actions of the Royal Sociev for Tropical Medicine and by allozyme, cytological and immunological data. Ausfral- Hygiene 19: 7-8. ian Mammalogy 2: 43-51. YAKIMOFF W. L. & MATSCHOULSKY S. N. 1936. Coccidiosis

SPEARE R., HAFFENDEN A. T., DANIELS P. W., THOMAS in the kangaroo. Journal of Parasitoloav 22: 5 14-5 15. A. D. & SEAWRIGHT C. D. 1984. Diseases of the Herbert ZWART P. &ISTIIK W. J. 1464. Globid&is in a Bennett’s River ringtail, Pseudocheirus herbertensis, and other wallaby. TijLschrift voor Diergeneeskunde 89 (Suppl. 1): North Queensland rainforest possums. In: Possums and 138-143.

Host-parasite associations of Eimeria spp. (Apicomplexa:Eimeriidae) in kangaroos and wallabies of...

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