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VOL. 122, PARTS 1 & 2 
29 MAY, 1998 


Transactions of the 
Royal Society of South 
Australia 


Incorporated 


Contents. 


Prideaux, G. J. & Wells, R. T. Sthenurus baileyi sp. nov., a new fossil ea 
from the Pleistocene of southern Australia - -  - 
Sheldon, F. & Puckridge, J. T. Macroinvertebrate assemblages of Goyder 
Lagoon, Diamantina River, South Australia - -  - 
Anstis, M., Alford, R. A. & Gillespie, G. R. Breeding biology of Litoria 
booroolongensis (Moore, 1961) and Litoria lesueuri 
(Duméril & Bibron, 1841) (Anura: Hylidae) and comments 
on population declines of L. booroolongensis - - - - 
Kolesik, P. A new genus and two new species of gall midge (Diptera: 
Cecidomyiidae) damaging young branches of Eucalyptus 
spp. in South Australia - - - - - - - - = - 
Steen, Z. & Schwarz, M. P. Within-nest behavior in a eusocial Australian 
allodapine bee Exoneura SERENE tridentata Houston 
(Apidae: Xylocopinae) - = - 
Field, S. A., Keller, M. A. & Austin, A. D. Field ectibiey aaa BehaOnE of the 
egg parasitoid Trissolcus basalis (Wollaston) Hymenoptera: 
Scelionidae) - - - - - - -~ = = - 
Beveridge, I. Woodwardostrongylus petrogale sp. nov. (Nematoda: 
Cloacinidae), from the stomachs of rock wallabies fetrogale 
spp.) from Amhem Land - - - - - - - = 
Nicholas, W. L. | Mesorhabditis kinchegensis sp. nov. (Nematoda: Rhabditidae) 
from arid soil in Kinchega National Park - - - - = - 


Brief Communications: 


McDonald, K. R. First Queensland record of the burrowing frog Cyclorana 
cryptotis Tyler & Martin, 1977 (Anura: Hylidae) - -— - 
O’Callaghan, M. G., Ockleshaw, E. & Allen, J. The prevalence and 
distribution of nematodes in the large intestines of sheep in 
South Australia - - - - - - = = = = - 


PUBLISHED AND SOLD AT THE SOCIETY’S ROOMS 
SOUTH AUSTRALIAN MUSEUM, NORTH TERRACE, ADELAIDE, S.A. 5000 


TRANSACTIONS OF THE 


ROYAL SOCIETY 
OF SOUTH AUSTRALIA 


INCORPORATED 


VOL. 122, PART 1 


TRANSACTIONS OF THE 
ROYAL SOCIETY OF SOUTH AUSTRALIA INC. 


CONTENTS, VOL. 122, 1998 


PARTS | & 2, 29 MAY, 1998 


Prideaux, G. J. & Wells, R. T. Sthenurus baileyi sp. nov., a new fossil Ranga 
from the Pleistocene of southern Australia- - - - - - 

Sheldon, F. & Puckridge, J. T. Macroinvertebrate assemblages of ‘Goyter Lagoon, 
Diamantina River, South Australia- -  - 

Anstis, M., Alford, R. A. & Gillespie, G. R. Breeding biology of Litoria 
booroolongensis (Moore, 1961) and Litoria lesueuri (Duméril & 
Bibron, 1841) (Anura: Hylidae) and comments on population 
declines of L. booroolongensis- - - - - - = - 

Kolesik, P. A new genus and two new species of gall midge (Diptera: 
Cecidomyiidae) damaging young branches of sik bane spP. in 
South Australia - - - 

Steen, Z. & Schwarz, M. P. Within-nest bebavtelie in a eisontia Australian 
allodapine bee Exoneura chide tridentata Houston 
(Apidae: Xylocopinae)- - - 

Field, S. A., Keller, M. A. & Austin, A. D. Field enslony aod ohesditerr of the egg 
parasitoid Trissolcus basalis (Wollaston) Hymenoptera: 


Scsifonidas) - = -—§ =~ 5S & » = # = & = 

Beveridge, I. Woodwardostrongylus  petrogale sp. nov. (Nematoda: 
Cloacinidae), from the stomachs of rock wallabies (Petrogale 
spp.) from AmhemLand- - - - - 


Nicholas, W.L. Mesorhabditis kinchegensis sp. nov. (Nematoda: Rhabditidae) 
from arid soil in Kinchega National Park - - - - - - 


Brief Communications: 


McDonald, K. R. First Queensland record of the burrowing frog Cyclonaien 
cryptotis Tyler & Martin, 1977 (Anura: Hylidae) - - - 

O’Callaghan, M. G., Ockleshaw, E. & Allen, J. The prevalence and distribution of 
nematodes in the large intestines of sheep in South Australia - - 


33 


45 


55 


65 


73 


79 


85 


87 


PARTS 3 & 4, 30 NOVEMBER, 1998 


Martin, H. A. Late Cretaceous-Cainozoic palynology of the Poonarunna No. | 
well, central Australia - - - - - - - - 

Kolesik, P. Rhopalomyia lawrenciae, a new gall midge species (Diptera: 
Cecidomyiidae) deforming leaves of Lawrencia squamata 
(Malvaceae) in South Australia- - - - - - - - 

Kolesik, P. Dasineura wahlenbergiae, a new species of gall midge (Diptera: 
Cecidomyiidae) damaging shoot tips of Wiphregrreipni stricta 
(Campanulaceae) in South Australia- - - 

Davies, M. & Watson, G. F. Developmental biology of Uperoleia talon Tyler, 
Davies & Martin, 1981 (Anura:Myobatrachidae)- - - 

Davies, M. & McDonald, K. R. A new species of frog (Anura: Mictany lice) fess 
Cape Melville, Queensland- - - - - - - - 

Davies, M. & McDonald, K. R. Developmental biology of Uperoleia altissima 
Davies, Watson, McDonald, ames & Werren, 1993 
(Anura: Myobatrachidae) - - + oe - 

Coleman, P.S. J. Changes in a Mangrove/Samphire Ciiiaiitey, North ae 
Creek, South Australia -  - - - - * + + = - 

Smales, L. R. New species of Seurechina (Nematoda : selsiiaion ‘sin in 
dasyurid marsupials from Australia- - - - 

Ferguson, M. A. & Smales, L. R. Spiroxys chelodinae Berry, 1985 (Nematoda: 
Spiruroidea) and Camallanus chelonius Baker, 1983 (Nematoda: 
Camallanoidea) from freshwater turtles ie Chelidae) in 
Queensland, Australia- - - - - ee 


Insert to Transactions of the Royal Society of South Australia, Vol, 122, parts 3 & 4,30 November, 1998 


89 


185 


STHENURUS BAILEYI SP. NOV., A NEW FOSSIL KANGAROO 
FROM THE PLEISTOCENE OF SOUTHERN AUSTRALIA 


By GAVIN J. PRIDEAUX* & RODERICK T. WELLS* 


Summary 


Prideaux, G. J. & Wells, R. T. (1998) Sthenurus baileyi sp. nov., a new fossil 
kangaroo from the Pleistocene of southern Australia. Trans. R. Soc. S. Aust. 122(1). 
1-15, 29 May, 1998. 

Sthenurus baileyi sp. nov., is described from Pleistocene deposits of Eyre Peninsula 
and the southeast of South Australia. The dentary is similar in size and morphology to 
S. occidentalis Glauert, 1910 but the cranium is much less inflated across the frontals 
and the rostrum less tapered anteriorly. Sthenurus baileyi is characterised by very low 
crowned molars, most similar to S. cegsai Pledge, 1992, S. brachyselenis Prideaux & 
Wells, 1997 and S. antiquus Bartholomai, 1963. Upper and lower premolars are 
similar to S. antiquus and §. brownei Merrilees, 1967. Overall, S. baileyi appears 
most closely related to S. antiquus and may represent the most plestomorphic member 
of the lineage containing the more brachycephalic sthenurine species. 

Key Words: Sthenurus baileyi sp. nov., Sthenurus antiquus, Sthenurus, 
Simosthenurus, sthenurine Kangaroo, Victoria Fossil Cave, Naracoorte, Brothers 
Islands, Eyre Peninsula, Pleistocene. 


Transactions ef the Royal Suciery of 8S. Aust, (1998), TI22( 1). 1-15, | 


STHENURUS BAILEY! SP. NOV., A NEW FOSSIL KANGAROO FROM THE 
PLEISTOCENE OF SOUTHERN AUSTRALIA 


by Gavin J, PRIDEAUX® & RODERICK T., WELLS* 


Summary 


PRipb AUN, G1 & Wrirs, RT. (1998) Srheniirus beilevi Sp. noy.. a new fossil kungaroo from the Pleistoeene 
of southern Australia. 7revey. R. Soe. 8. Auge, 122 (1), 1-15, 20 May. 1998. 

Sthenuras baileye sp. nov, is described from Pleistocene deposits of Eyre Peninsuli and the southeast of South 
Austrilia. The dentary is Similar insize and morphology to.8, eectenralis Glauert, 1910 but the cranium is much 
loss Inflated across the frontals and the rostrum less tapered anteriorly. Siemurus Dadlevé is characterised by very 
low crowned molars. most similar to8, ceesa/ Pledge, 1992. 8. brachyselenis Prideuux & Wells. 1997 and 4. 
untiquus Burtholomai, 1963, Upper sind Jower premolars are similar to $8. aarigqus and S. browne’ Merrilees. 
L067. Overall, §. badlevi appears most closely related to S. curignii and may represent the most plestomorphic 
iiember of the lineage vontuming the more brachycephilic sthenurine species. 


Kiy Worbds! Siediirus baileyi sp. nov., Sthenuras antiqiis, Sthenarus, Simeasthearus. sthenurine kangaroo, 
Victoria Fossil Cave, Naracoorte, Brotiers Islands, Lyre Peninsula, Pleistocene. 


Introduction 


Following its discovery in 1969, the extensive 
Pleistocene deposit within Victoria Fossil Cave iat 
Naracoorte. South Australia has yielded cremains 
from around one hundred vertebrate species, 
Included are slightly less than half of the known 
Pleistocene species of sthenurine kangaroos 
(subfannly Stheaurinuek Proceprodan rapha Owen, 
187d, Sthenuruy anderson’ Marcus. 1962.5. browne’ 
Merrilees, 1967, S. gilli Merrilees, 1965, 8. 
maddocki Wells & Murray, 1979, 8. cecidentalis 
Glauert, L910. 8. pales DeVis, 1895 (Wells ef eal. 
1984). and a oew sthenurine, & bailey sp.nov: The 
Species is also known from a single specimen 
collcered from an eroded cave on one of the Brothers 
Islands in Coffin Bay, Ryre Peninsula (Brown 1908: 
Fig. 1). Williams (1980) identified the cranium and 
ussociuted dentaries us Sifenurus cf moddock), bul tt 
is here designated us the holotype of 8. heilevi sp. 
nov, Deseription of the new species and a 
consideration of tts phylogenetic implications form 
the subject of this paper. 


Materials and Methods 


The material is housed in the South Australian 
Museum, Adelude (prefix SAMA) and Flinders 
University (prefix PU). Dental homology follows 
Flower (1867) and Luckett (1993). Dental 
nomenclature follows Tedford & Woodburne (1987), 
Ride (1993) or is stundard. Mensuration follows 


“Schou! of Biological! Scnnces, Phuders University of South 
Australia GMO Box 210) Adeluide S.Aust SOOT, Boil: 
wav ti prices @flinders cutiiut 


mADUha 
. 


= yeaycouRTe 


Pig. 7. Map of southeastern Australia showing location of 
deposits yielding Sthenurys bailey? sp. nev. 


Tedford (1966) and Wells & Murray (1979). Dental 
measurements (mm) are provided im Rible |. 


Systematics 
Order Diprotodontia Qwen, 1866 
Suborder Phialungerida Aplin & Archer, 1987 
Superfamily Macropoduidea (Gray. 1821) 
Fuimly Macropodidie Gray. 1821 
Subfamily Sthenurinve (Glauvert, 1926) 
Genus Sthenurus Owen, 1874 
Subgenus /’Simosthenurus Tedford, 1966 


Sthenurus (?Simosthenuruy) baileyi sp. noy, 
(FIGS 1-8) 


Holotype > SAMA P13670, partial cranium (wath Li 
3, dP2, dP3, M1-4. excivated P3; Fig, 2A,B, 3A, 4A, 


2 G. J, PRIDEAUX & R, T. WELLS 


TABLE 1. Cheek tooth dimensions of Sthenurus baileyi. 8. brachyselenis, S. brownei (eastern form), S. cegsat and S-antiquus: 
mein, standard deviation (parentheses), range (brackets). 

Abbreviations: L= length, AW = width of anterior loph(id); PW = width of posterior loph(id); AH = crown height of anterior 
loph(id) on buccal side; PH = crown height of posterior loph(id) on buccal side; n = sample size. Note that crown heights 
are heavily dependent on degree of enamel wear, hence, frequently high standard deviations. 


Tooth — Species L AW PW All PH " 
UPPER DENTITION 
dP2 S. baileyi 10.5 75 10,0 62 6.1 | 
S. baileyi TYPE As above 
S. browne 10.9 (0.43) 8.7 (0.40) 10,8 (0,38) 7.0 (O51) 7.9 (0.00) 15 
(eastern form) [104-1 1.9] [8.1-9,5] {10.4-11.4] 16.0-8,0] [6.6-8.7] 
No veusat = = = rf - = 
S. antiqaus . - - - - - 
IPS S. batleyt 10.6 99 108 - - | 
S. baileyi TYPE As abave 
S. brawnei 11.3 (0.29) 10.7 (0.32) 11.0 (0,37) 5,7 (0.54) 5.9:(0,46) 15 
jeastern form) [10,6-1 1,7] [10,.2-11.3] [10,6-1 1.8] [18-64] [5.3-6.8| 
S. cegvat 93 8.6 92 63 63 | 
S. caentiyguns : 7 7 - 
PS S. baileyi 17.2 (0.14) 9.9 (0.28) 12.9 (0.21) 9.8 (0.28) 9.7 (1.06) 2 
[E71 -17,3] (9.7-L0.1] [12.7-13,0] [9.6-10,0] {8.9-10,4] 
S. baileyi TYPE {7.1 10.1 13.0 10.0 14 
S, browned 17.1 40,57) 10.9 (0.68) 13.7 (O.81) OLR (0.76) 9,7 (0.78) 21 
(eastern form) {16.2-18,0] [9.0-11,8] {12.1-15.0] [S.3-LL.1] [8.6-1 1.3] 
S. cezsai = - - . - 
S. antiques 14.4) 6.3 11.3 bt 11 | 
Ml S. hailevi 12.3 (0.14) 12.3 (0.21) 12.3 (0.2L) 6.3 (0.07) 6.6 (0.42) 2 
{12,.2-12.4| [121-124 {12.1-12.4] [6.2-6.3] [6.3-6.9] 
S. baileyi: TYPE 12.2 12 Pe | 6,2 63 
S. brawnei 12.9 (0.43) 12.4 (0.40) 12.3 (0,34) 6.1 (0,76) 6,5 (0.62) 28 
(eastern form) [12.2-13.6] {116-130 [ELB-13.1] [5.0-7.9| [S.3-8.1] 
S. cegyeail - : - - - 
S. antiques 12.4 12.0 12.) - - | 
M2 S. haileyi 13.8 (0.21) 13.2 (0.21) 13.1 (0.14) 6.9 (0.42) 7.2 (0.35) 2 
[13.6-13.9] [13.0-13,3 }13.0-13.2] [6.6-7.2| 6.9-7.4) 
S. baileyi ‘TYPE 13,6 13.3 13.0 V2 7A 
S. brownet 14,1 (0.37) 13.6 (0.46) 13.1 (0.43) 6.6 (0.76) 6.9 (0.58) 23 
(eastern form) [13.2-14.7[ {12.9-14.4 {12.5-14,3] [5.3-7.8| 5.8-8.0] 
S. cegsai - - - - 5.6 ! 
S. cautiquits 14,9 (0.07) 13.4 (0.14) 13.0 (0.21) 7.20113) 74 (1.34) 2 
[14.8-14.9] [133-135 {12.8-13.1] [6.4-8.0] 6.4-8.3] 
M3 S. baileyi 14.5 (0.00) 13.7 (0.21) 12.6 (0.35) TA (0.14) 6.8 (0.28) 2 
{14.5} [13.5-13.8 [12.8-13,3] [6.9-7,1] 6.6-7.0) 
S. baileyi TYPE 14.5 13.5 12.8 7A 6.6 
SL browne 14,5 (0,39) 14.9 (0,50) 12.9 (0.54) 6.7 (0.62) 7.0 (0.53) 19 
(eastern form) 13,7-15.5| [13.3-14,8] [12.3-14.5 [5.48.1] 5.9-8.1] 
NS cegseet 13.1 (0.07) 115 10.9 (O14) 5.4 (0.00) 5.5 (0.14) z 
13,0-13.1] [10.8- 11.0 [5.4] 5.4-5.6| 
S. antiquus 16.0 (0.35) 13.6 (0.49) 12.9 (0.85) $.3 (0.35) 4.3 (0,00) 2 
15,7-16.2] [13,2-13.9] [12.3-13.5 [8.0-8,5} [8.5] 
M+ S. hailevi 13.8 13,4 1L.3 6.7 6.1 | 
S_hailevé TYPE As above 
So hrawnet 14.0 (O43) 12.2 (1.48) 11.2 (0.40) &.1 (0.62) 7.4 (0.78) 16 
(eastern form) 13,.2-14.5] [11,3-13,0] [10.6-12.0 [7A-9.6} [6.5-8.8] 
S. ceesat 12.2 10 o5 55 a4 | 


S. cantiyuun 


TABLE 1. — Continued 


NEW PLEISTOCENE KANGAROO 


LOWER DENTITION 


up2 


ml 


m2 


ms 


m4 


S. baileyi 


S. buileyi TYPE 
S. brachyselenis 


S. browne 
(eastern form) 


S. cegsai 
S. aintiquus 
S, baileyi 


S. baileyi TYPE 
S. brachyselenis 


S, brawnet 
{custern form) 


SL cewyat 
S. antiqnus 
S. bailevi 


S, butileyi TYPE 
S. brachivselenis 


S. browne’ 
(eastern form) 


S. cegvat 
NS. cuvetigules 
S. baileyi 


S. baileyi TYPE 
S. brachyselenis 
S. brownet 
{eastern fort) 

S. cegsat 

S. antiquus 

S, baileyi 


S. baileyi TYPE 
S. brachyselenis 


SL brownet 
(eastern form) 


8. eersal 


SL cgentreptiues: 
S. baileyi 


So baileyi TYPE 
S. betchyselenis 
S, brawnel 
{eustern form) 


S. cevsal 
S. autiquus 


S. bailevi 


S. baile TYPE 
S. brachyselenis 
S, browned’ 
(eastern form) 
S.ceusal 

Si antiguas 


26 


uA 
8.1 


9.8 (0.45) 
[9.2-10.9] 


9.8 (0.14) 
[9.7-9.9] 


99 
10.2 


10.4 (0.36) 
[100-111] 


16.2 (0.78) 
[15.3-17.8 


15.3 
13,8 


16,2 (0.53) 
[15.2-17.0 


14.8 
17.6 


12.0 (41) 
[11.5-12,5 


12] 
13.9 


13.1 (00.64) 
|12.1-14,7] 


10,5 
13.8 


13.3 (0.46) 
{12.8-13.9| 


12.8 


14.7 (0.53) 
{13.8-16.1] 


12.7 
15.3 (0.85) 
[14.7-15.9] 


14.1 (0.64) 
[13.6-15.0] 


13.6 


14.9 (0.51) 
(13.7-15.7] 
12.8 
16.0 (0.99) 
115.3-16.7] 


13.9 (0.11) 
[13.7-14.0] 


13.8 


14.0 (0.43) 
[13,2-14.5] 


11.9 


15.4 (0.49) 
[15.0-15.7] 


a 
6.5 (0.34) 
[5.9-7,2] 


8.90.71) 
[8.4.9.4] 


Sab 
7.8 


9,2 (0,46) 
[8.4-10.0] 


8.0. (0.32) 
|7.7-8.4] 
7.9 
6.3 


8.5 (1.34) 
[R095] 


6.5 

8.2 
10.1 W.22) 
19.9-10.4] 

10.0 

909 
10.4 (0.49) 
[9.4-12.0] 


11.4 (0,27) 
[110-116] 


hs 


11.4 (0.39) 
{10.4-12.0] 


10.8 


11.9 (0.71) 
[A124] 
12.2 (0.37) 
[11.7-12.6] 
12.6 


12.0 (0.43) 
[112-129] 


Ho 


124 (042) 
[}2.1-12.7] 


12.1 (0.26) 
[11.8-12,3] 


2.2 


12.2 (0.48) 
| 11.3-13.0] 


10,3 


8.9 (0.99) 
[8.2-9.6] 
8.2 
6,1 


91 (46) 
[8.6-9.7] 


9.1 (0.28) 
[8.9-9,3| 


8.9 
8.2 


O93 (031) 
[8.7-9.8] 


9.7 (O44) 
9.1-10,3] 
95 
8.0 
10.3 (0.58) 
9.4-11.5] 
7.2 
10.1 


0.1 (0.26) 
9.8-10.4| 


10,2 
10.3 


10.6 (0.43) 
[96-115] 


1.0 


H.2(041) 
[10.7-11.7] 


7 


11,7 (0.33) 
[11.1-12.2 
10.3 


12,1 (0.78) 
[11.5-12.6] 


11,9 (0.42) 
[114-124] 


12.4 


12,2 (0,35) 
[11.4-12.9| 
10,2 
12.2 (0.49) 
[1 1.8-12.5] 


11.1 (0.42) 
[10.5-1 1.5] 


11.2 


11.2 (0,40) 
{10.6-12,0] 


8.6 
11.0 


73 


73 
5.7 
7.5 (082) 


9.5 (1.04) 
17.9-1 1,0} 
89 
73 
9.9 (0.85) 
[8.9-11.4] 
8.3 


7.1 (1.01) 
[6.3-8.5| 


64 
9.3 


8.6 (1.04) 
16.4- 10.0} 


5.5 


83 (0.49) 
[7.8-8.9] 


8.0 


9.5 (0.94) 
[8.2108] 
59 
lhe 


8.2 (4.93) 
[7.0-8.9] 


78 


9.3 (0,61) 
(8.4-10.3] 


TA) 
10,5 


74 (043) 
[7.1-8,0) 


71 


8.1 (0,621 
\7.4-9.6) 


6.2.71) 
(5.7-6.7| 


5.7 
5.5 


7TALU7 
[6.40-8.2] 


6.0 


70 


7.2 (0.63) 
[6.0-8.2] 


9. (1.03 
[7.3-10,2] 
91) 
6.7 
9.9 (0.83) 
[8.9-1 1.3] 
71 
4 


7.0 (0.76) 
[6.3-8.0] 


6S 
94 


8.7 (1.07) 
[6,5-10.2] 


85 


8.4 (0.63) 
[7,8-9.2] 
8.0 


9,5 (0.80) 
[8.0-11.2] 
59 
10.4 


TITY 
[6.9-8.4| 


TS 


9.1 (0.73) 
17.5104] 


64 
10,3 


7.2: (0.35) 
[6.9-7.8] 


7A 


TA(OTS) 
|6.5-8.8] 


45.3 


7.7 (0.35) 
[7.4-7.9] 


te 


i 


17 


‘ G.I. PRIDEAUX & RT. WELLS 


SA) left and ngnt dentanes (with il, dp2. dps. nil - 
4. excivated pa: Fig. 3B, 4B, 5C,D), apparently: 
collected from a bone breccia in-an eroded cave on 
the western end of west Brothers. Island (34 35' 8, 
135° 20' EE), Coltin Bay. Eyre Peninsala South 
Austulin (Brown 1908: Williams L980: Fie. 1). 
Other mammals frona the deposit include Macrapus 
rufagrisens, Potoraus plaivops, Pyeudocteirus sp. 
Rarus fuscipes and Neopheco cinerea. A huge bird 
femur previously atiibuted to Gereverinty preween 
(Rich 1979) belongs toy Drwnailis rovaehollandice 
(J, MeNatnara pers, con, 1996), Age of type 
locality ty considered Pleistocene because all taxa 
identified to species are only known from the 
Quuteruiry. Sumhirly, the genus Sihenieuy appeurs 
nol lo have existed anywhere beyond the lite 
Picistovene, Details of collection are uncertain bul 
probably retrieved by DLR. George around 1902 (1 
MeNcuirrunt pers, eaimm. 1996), 


Divaeiveasts 

Cranium similar iy srze to Stenuray ecenlentalin 
bul fronts less expanded. rostrum shorner and 
brouder with wider misals and larger minal aperture, 
PA similar to So deevwied but wath relatively marrow 
shallow longitudinal basin and mwa accessory 
cuspales unieror jo promiment poslerchuceut 
uceessary cusp. Upper molurs very low crowned. 
With shart preemmuluen weak pustprotoerista and 
very Well developed postparaerista. Dentary sivrlur 
ine sivve and morphology to S. vecidesitalis and 4S, 
cniieitys — Barthologiai, (463 but with mare 
posteriorly inflated pleryeoid fossu thiunm in any 
Siieniias species, Posteroventral batder uf 
neassclerie Fossa expanded laterally imu wide shelf, 
similar to SL cess, So aif and S. widddowki, iL 
intermediate belweens, occidentalix anil S. browees 
in genera! shape and degree of prociumbeney. pF 
must similar in morphology to 8. entiguds but lower 
erowned, with straighter lingual crest. Lower molars 
very low crawned. with anteropesterionly short 
trivonid, well-developed premetactistid. aud very 
reduced onstidd obhgua and parucnstid producure i 
morphialugy closest lo Neves, § uniques and S, 
hrachvselenis Prideaus & Wells, 199) hut wader 
relative to leagth, 


Description af holawpe 

Vertical portion of prenaxila thived dorsally 
prowidibg -clonwate contac)! with uasals. Diusterma 
shorn, unterinr V/s comprising premasxilla and 
posterior i minxitla. Tneisive foramina long. narrow, 
untecior border level with posterior extreme of 13 
alveolus (Pig. 3A). Rostrum short, lapered anteriorly 
(Fig. 2A). Buceinitor fossa on maxilla rather shallow 
ameriolly, deeper postetiorly gthterion to zygomatic 
wich. Massetenic process well-formed, railier narrow, 


eroded off ventrally on leit and nent sides. Nasals 
very broad posteriorly dnd, although broken 
antenorly, clearly short. Nasotrontal suture gently 
sinusoidal (Fig. 2A). Prontals moderately inflated 
anteriorly: supriorbital crests only slightly 
developed (Fig, 2A). Temporal crests: moderately 
developed, not tully conversent upon sagittal suture, 
Large infraorbitul foramen positioned anteroventral 
und tnestal to Lichrymial foramen, just below orbital 
centre. Palatal vacuities extend anteriorly to anterior 
extreme of MI (Pig, 4A). Right lateral extremly of 
broken postpalatine bar devel wilh M4 jnterloph 
yullev. 

1] crown ruther long. moderutely wide. wath 
vertical occlnsal surface faving posteriorly, 12 very 
small splint-like, Vasize of LL, 13 high crowned, but 
qyuile short iunteropostertarly (Pig. 2B). Small 
anterolingual lobe evidenton 3. 

dP2 reminiscent of P3, rognded in general outline. 
especially lingually, bur much shorter relative be 
width (Fig, 4A). Buceul and lingual crests siraisht, 
excepl for buccal curvature of lingual crest a 
posterior extremity, Anterior bisio sinh quite deer 
und separated from longuddimal basin by tow 
transverse ridgelet. Posterior basin appears to juve 
bee relabvely Targe, approx, hall size ol 
longiludinal basin. 

JP3 completely molarifurnm und similar in general 
outline t ML but differs by tiiving Lophs orientated 
obliqgely (not perpendiculird ta buceal and Trial 


sides of tooth (Fig. TA). In addition, precigulum 


very sha@bt terminating before reaching lingual 
extreme of tooth. Premetaerista appears well- 
developed. Profoloph appeurs to have been vary 
curved. convex anteriarly. No enamel crentlarans 
present i interloph valleys yery low. barely 
delectible postprotocristy positioned just Wagual of 
dP3 midline (Piz. 44). Postmetacwnulecrisia curves 
dorsobuecally frome metuconule to mect verticul 
postinetlcrishi. Saud) accessory crest positioned 
mesial ta postnetacristia, slight poslink centrally 
positioned on pasterior meraloph tuce, 

PS rounded jn oulline and tipered antenorly (Mig 
SA). Longitudinal basin shallow, Buceal erest barely 
excerding lingoul crest m height. Anteriarly, lingwil 
erest begins. tO cin purillel fa buceal crest then 
posterior 4/4 of crest curves oul Tingually. Small 
umertor busin present and separated from 
longitudinal basin by drunsverse ridge descendiny: 
from anterior buceal cusp terminating adjacent to 
anterior Tinguad casp. Posterior basin short. well- 
formed and separated [rom longitudinal basin by 
low tringverse ridge originating From posterior 
lingual cusp and onentated obliquely (slightly 
unterobueaily) to meet low down on buccal crest, 
Main posterobuccal aevessury cusp well-formed, nul 
quite as high as posterior parcot buceal erest. Three 


NEW PLEISTOCENE KANGAROO 5 


= 


aw / 


—) , 2a * mM 


mp 


Vig. 2. Sitenaras ballew! sp. oy, ccaniuin, A. Holotype 1 P1AG70) dorsal view. BL Holotype lateral view, Seale bars = 10min 
Abbrevs; d = ditistema, [= front, i= niaxilla. mp = tasscterte process. W= basalonls = qesolrontal sature, p= parietal 
pire Prem. s = supragrbital erest. 


“ G, J, PRIDEAUX & R.'T, WELLS 


Hig. 4. Schein baileyi sp. nov. cranium and dentaries. 


Stereopai of holotype dentary cectusal view. Scale bars 


A. Stereopair of holotype cranium (PI3670) palatal view. B 
= 10mm. Abbrevs: a = anterior rool oF uscending tinus. b = 


buccinlor fossa an Alisa © = tandibular condyle, i = incisive foramina, pb = posteroyenteal border Of masseteric 


fossa, pv = palatal vacuities, 


small poorly separated accessory cuspules 
positioned anterior to mui accessory cusp (Pig. SA). 

Upper molars very low crowned, with protoloph 
equal in width lo metaloph in MI-2, but wider in 
M3-4 (Fig. 4A). Precingulum short. bueeal extreme 
terminating at distinct cuspule. representing cither 
stylir cusp A or B. Slight crest (probable puracrista) 
connects cuspule posteriorly ta puracone. Two to 
four slight vertical crenulations centrally located on 
precinguliim, with most togual probably temant 
preprotocrista (forelink). Postprotocrista weak, low, 
ascending buccally across fice of protoloph into 


interloph valley, uniting with vertical crenulation 
directed posteriorly from mid-pomr on protoloph, 
Postparacrista strongly developed, forming buccal 
border Of interloph valley, meeting slight 
premetierista on anterior face of metiloph (Pig. 4A) 
Interloph valley with few very fine to no enamel 
erenulations, Postmetaconulecrisit sweeps ucross 
posterior luce of metuloph terminating just posterior 
toend of postmetacrista, Two to three small distinet 
crenulations enclosed by postmetuconulecristy an 
metaloph posterior Mee, 

Pentary moderately proportioned, except for 


tt 


NEW PLEISTOCENE KANGAROO 


Fig. 4. Sthenurus baileyi sp, noy, cheek tooth rows, A, Stereopair of holotype (P13670) left upper cheek tooth row occlusal 
view, B, Stereopair of holotype (P13670) right lower cheek looth row occlusal yiew. Scale bars = 5 mm. Abbreys: co = 
crisid obliqua, pe = preciagulum, pd = paracrisud, ppe = postparacrista, ppre = postprotocrista, f = trigonid, 


Fig 5. Sthenarus baileyi sp. noy. premolars, A, Stereopair of holotype (P13670) left P3 close-up occlusal view. B, Stereapair 
of paratype (FUOQTG67) left P3 ceclusal view. C. Stereopuir of holotype left p3 close-up occlusal view, D. Stereopair of 
holotype Jell p3 occlusal view. Seale hars = 5 mm. Abbrevs: ac = uccessory cuspules. be = buccal crest, lb = longitudinal 
basin. le = lingual crest, me = main crest, my = median valley, phue = posterobuccal accessory cusp. 


x fF 1 PRIDEAUN ART WHEELS 


posteriorly intlited pterygoid fossa and lateral 
Axpinston of posteroventral border of mussereric 
fossee into wide shelf. Ramus moderately deep for 
wilh, partivularly in region of symphysis, 
Sympbysis gently tapered anteriorly and posteniorhy 
only extended short way beneath genial pit. below 
anterior root of dps. Digastrie eminence present hut 
Hol patticularly prominenr Digastric sulcus 
eatending from below anterior extreme ob pterygoid 
fossa to helow m2 hypolophid. Diustenta short, with 
fedian dorsal groave deep. relaively wide. Very 
shullow buceindter sulcus arises near posterior 
extreme of diastema, dorsal to large anterior mental 
forsmen. Bueeinalor sulcus deepens slightly 
posteriorly, terminates below ml hypolophic 
Posteriar mentil foramen positioned below m2 
hypolophid. hallway between dorsal and ventral 
borders of mans, 

Anlenor ront of ascending ramus begins adyucent 
to m3 hypolophid (Fig. 3B), extending posteriorly to 
form huceal border of postuveolur fussa, Preryyor 
fossa inflated postertarly, projechine well beyond 
border Of Mmasseteric fossa when viewed Jaterally 
Masseterie toss deep. due largely ty laterally 
expanded posteroyentral border (Pig. 38), Ventral 
border of nasseterte fossa ut sume honeatal level as 
posterior region Of buceinator sulets. Musseteric 
Jofamen moderately latee. vertical in opeptation, 
Inferior mandibular foramen rither small, At 
untenior exueme of pterveoid fossa. anteromedial to 
iitenior mundibular foramen, dorseventally wade 
mylohyoid grouve present. This appears to hive 
heen partially overhung by shagp adterodorsally- 
directed process al anteremedial border of pterygoul 
fossa, and similurly-shaped  posteroventrally - 
direcied process positioned helow postenoe extreme 
ot postaiveotur fossa. Mandibular condyle 
moderately laree (hig. $B) Angular process well- 
developed. rising dorsally to acute point. 

il rather shor. slender upturned. with oeclusal 
surface aba horicantal level just above Base of check 
well crowns. dp2 an both sides ef holotype toa wart 
or fragmentary to interpret. Likewise, dpa very 
worn, allhough clearly molinform, possessing, low 
bur well-defined parm premeta-. prehypo- unit 
preentu-cristids. 

p3 considerably longer than any molur, with main 
(Lingual) erest cxtending trom posterofingual vormer 
ly failing oi toeth anteriorly (Pig. SC.D). ‘Three 
cUspules form anterior part of main crest. with each 
bearing pair of literal ridgelets, one descending 
buccally. one fingtally. Buceal ridgelets lernimpate at 
fiw shelf formed by three conlluent cuspules, 
loved immediately anterior to buccal crest, Buccal 
crest straight. shor. equal in length to and mirroring 
shape af pestetior part of main crest, Median valley 
rather narrow. modenitely deep, Toward ts posterity 


(A, median valley waversed by coarse ridgelet (Fig. 
SCD). 

Lower molars very low erowned, with protolophid 

und hypolophid ecclusal suttaces finear and close to 
parallel. Trigonid very short, with paracristid low 
and composed of two moieties. Degree of separation 
of anterior and posterior muiglies increases from m1 
to m4. Posterior part of paracristid sweeps smoothly 
dnterolingually across protolephid face, terniinates 
on bueeal side of anterior part Tn more posterior 
Molars, aMenor conjponeat of paraeristid shifted 
more lingually bul posterior extreme remains within 
buceal V/s of anterior protolophid face. originaliny 
well below lophid apex. A few fine enamel 
crenulations umse low down on antenor fuce ol 
prololophid and descend (ito tigontd basi, Lingual 
side of Ingonid bordered by welledeyeloped 
premetacristid. which termigates at puraconid 
Previngulid small and positioned anterobuceal ws 
paracrisiid, extending lingually as very thin 
pebinsija at antefior extreme of moku. Cristid 
obliqust (prehypnenistid)y low, sumihirly developed 
and aligned ia similar position on hypolophid: as 
pursertsod on prutolephid. Preentoctisuid’ very low 
and barely detectable. Asie from these weakly 
developed crests, shallow interlophid valley bears no 
enumel crenulations, Pastener face ol bypotoplre 
with low. shallow inflation: 
Paranpes: Prom Victoria Fossil Cave, Naracoorte 
South Australia (37 00'S. 140° 48) Ba, PUOOOA Jeit 
dnd nghe adult dentures: PUOLG7, let) PA, M1 (PS om 
Fis, SB), FUOL6S, right pa. FUOI94, partivl right 
denlary; SAMA PI6S31/P 16558. lett and right autuls 
denturies (left dentary in Fig. GA): P28287_ right 
juvenile derntary: P28659. right M2. lett M4, 
PUOON4, PUOH67 and SAMA P28659 may belang to 
sane I)dividual based of proximity ty deposit 
degree of enamel wear und occlusal hb Specimens 
collected’ by Prideauy. Wells and others. Age of 
deposit $s medial to lite Pleistocene (Wells er ul. 
1O84: Ayliffe er al 10 yrress). 

Features out preserved adequately nm holotype cue 
deseribed from paratype SAMA P2822. 

Up? -cquil in length to dp3, yery similar iy 
morphology to p3 bul wider relative fo length. As in 
po. tinee cuspules dominute untenior half of main 
crest. cuch with tunsverse ndgelet on bieeal sule 
Ridgelets likely to have terpiinated 16 Gity Cuspules 
like p3, but due to considerable weur Sustained have 
become conflucnt with biiceal crest, vonveying an 
iHpression of More clopgale crest. 

Completely molaritorm. dps bears prololophn! 
tapered more tlaward Jophicl apex than hypolophid, 
As with dp2. wear has rernoved several leutures 
However, casdd oblique uppears more stmongly 
developed than in niolars and curved directly frou 
hypoconid apex into interlophid valley. terminate 


NEW PLEISTOCENE KANGAROO 9 


Es 


Fig. 6. Sthenurus baileyi sp. nov, left dentary, A. Paratype (P16558) lateral view. B. Paratype (P16558) mesial yiew. Scale 
bars = 70 mm. Abbrevs: amf = anterior mental foramen, bs = buccinator fossa, d = diastema, de = digastric eminence. ds 
= digastric sulcus, imf = inferior mandibular foramen, mf = masseteric fossa, pmf = posterior mental foramen, ptt = 
pterygoid fossa, s = symphysis, 


Ww fi J PRINBAUX & &.T WELLS 


cen(tally on posterior protolophid faee. Very weak 
precarocristid also present, curving from entocenicd 
jolo interlophid yalley. lenmmating Tingual to eristiel 
obliqua. Enamel crenulations, similar to those on 
molars, gppear to have been present on anterior 
lophid faves. Slight, rounded posteingulid on 
posterior face oF hypolophid appears confluent with 
slight postentourstiut. 


Etyvinedtoey 

Named in honour of Mr Edwin Ed” Bailey whose 
efforts aver Une last 25 years have contibuted so 
much to fhe success oF pulaeontological work in the 
Naracoorte Caves, 


Veridian 

Unfortunately, puly one cranium is khown af, 
bakeyi sp now and variation within the upper 
denution ean unly be assessed by comparison of PS 
ind MI-3, which are each represented by two 
specimens, P3 is very similar in the holotype and 
PUOI67, with the slight occlusal wear in PUOTG7 
responsible for most of the superficial differences 
between the specimens. In the holotype. PA is 
slightly wider anteriorly, both across the whole tooth 
and tie longitudinal basin. The lingual surlace of the 
holotype P3 is slightly more convex and rounded 
than FUOL67, The three cuspules anterior tothe main 
posterobuccul weessory cusp ure More sepuruted in 
FUOIGT. 

Only ote slight difference is deteerable on 
comparison of MI-3 of PI3670, FUOQT67 (MI) and 
P28059 (M2-3). The postparacrista 1s larger in the 
holotype. While greater weur sustiined hy FU A7 
und P2S86S9 could decount for these differences, 
consideration of the mater mm which teeth oeelude 
suggests that they are more likely la reflect 
morphological variarion. 

Conplete or purtiu] denuiries are known for five 
maiwviduals, with three charucters clearly variable 
Depth and exten of the digastric sulcus is the most 
varidble character, Although deep and extending 
frony (he anterior extreme of the pterygoid fossa uy 
below the m2 hypolephicd in PIS670 and 
PLOS3 1/PL6S358. the suleus i much shallower und 
only extends to below the m4 protalophid: in 
FLOOO4. In P28282. the divastric suleus is even 
shallower thus pepating the diagnostic unlity of this 
chamacter, The degree lo which the plerygond (oss is 
inflaied posteriorly alsa yaries belween specimens, 
Inflation is greatest in PLO53 WP TASS8, slightly less 
mo the bGlatype (P13670). PUOOO4 and P28282. and 
ledst in PUO204, However. ibis sulficient in the latter 
(Oo mink fl US @ distinctive feature of So hetilevi. 
Dentary deprh relative te width js greater in 
PLOS31/P1 6558 (depth to width ratio below (2-3 = 
179) compared fo FUOQ04 (1.65) and P2822 


(1.61). The ratio is lowest in the hololype (1.46). 
Intraspecifie variation in dentary depth rehitive ta 
width ms cormonty observed in sthenurine species 
known Irom even small sumple sizes, 

Variauion in pa size is common in all sthenurines, 
Mivluding S$. ballew. While most of the paratypes are 
very similar in size. PIGS30/PI6551 and the 
holotype are noticeably shorter and narrower 
Morphology ovaries only  shehily between 
individaals, primarily in the form of the buccal erest 
and minor variation in width of the median valley- 
The anterior half of the buccal crest in P28282 os 
slightly higher than the postertor half and curves 
posterolingually, becoming conlluent with a 
transverse ridvelet which crosses the median yalley 
This buceul crest roorpholoagy is not observed in any 
of the other specimens, although a very similar 
wansverse Pidgelet traverses the median valley in 
PI3670, Apart from this. feature, only the relative 
jofladon of the anterior region of the p3- varies 
slightly. A p3 referruble ro $8. baileyi as also known 
from Lindsay Hall Cave, near Madura on the 
Nullarbor Plain, Western Austra bat this speemmel 
remuins in the private collection of L. atehern Perth, 
This specimen is inseparable in sive and morphology 
from the South Australian Specimens. 

There iy little varfation in both sive and 
morphology of the lower molars, although the 
premetucrisud. paracrnishd and cristid obhqua of the 
paratypes are slightly more weakly developed than 
the holotype and the anterior lophid fees bear more 
fine eoumel crenulaGons, In addition. the 
postemeuhd os more shelf-like im each of the 
paratypes than in the holotype. except i) FUO2Od 
where there is a larger inflation of the ventrobuccal 
region of the hypalaphid posterior face. 


Comparison with other terxe 

Cranium. Although P3 was unerupled ine the 
holotype of S. baileyi sp. nov. the presence ol M4 in 
seclusion indicates that P3 eruption was imminent, 
An examination of other species for which a wuod 
age series ts known, reveals that lithe change in 
morphology or size in OSL uspects OF the eruniunt 
and dentary occurs From (his ontogenetic stage toile 
stage where P3 is erupted. This means that direct 
comparisons with older represemtuitives of ather tavu 
ave lenable. Leis worth noting that the two saniples al 
§. browne! and S. cecidenntlis with whieh 8. Auileys 
Is compared come from Naracoorte and ure 
considered fo represent the castent tons of both 
species, Although very similar in overall 
mormbology. they can he distinguished from the 
Loporypic Western Australiuy samples by their larger 
overul sive and Shelly smaller dentition relative to 
Ji Ane. 


NEW PLEISTOCENE KANGAROO Vl 


‘The cranium of 5. barfleviis very similiar in size and 
brachycephaly to 8. eceidenralis, The premaxillac 
are also similar in relative size and morphology. 
Although rostral length of the two species is similar, 
the bucginator fossa on the side of the maxilla is 
deeper in 8. eceidentalix. This is coupled with a 
mesially concave aspect to the edge of the diastema, 
in contrast to the less distinet edge and shallow 
buceinator fossa in §. hadley’, This condition is more 
reminiscent of S. gilli and 8. andervant. 


The rostrum ofS. baileyi docs not taper to the same 
degree anteriorly as 8. vecidenialis, both becuuse (he 
frontals are less expanded and its qarial aperture is 
proporGonaly kuger Among the Siientiris species 
for which the splinehnocrunitun is known, lateral 
inflation Of the frontal region (particularly anteriorly) 
nd formation of supraorbital crests is greatest in 8, 
madidocki, S. occidentalis, S. stirling? Wells & 
Tedford, 1995 and §. Arawret The frontal region is 
relatively narrow in §. gilli, S. détdersant and S, 
Hindale? Tedford, 1966, The proportions displayed in 
S, baileyi are intermediate between these two groups, 
particularly bemween §. browne? and S. erlli. 
However. the nusals of S. beflev are very wide and 
constitute a greater proportion of the dorsal aspect of 
the fosteum than any other Sthemris species. except 
S. maddocki. Overall, the short and broad nature of 
the rostrum ts characterisue of S. balleve. 

The anterior extent of the palatal vacuities in 9 
haley is akin te a number at other species, 
feyminatings close wo the GPS metaloph, or whit 
would be close to the posterior extreme of the P3 if 
it were in Geclusion, The masseteyic process appears 
iu have been well-developed, allowing for the 
damave in the holotype, and is intermediate between 
S. mudedocks and §. browner in size. 


Upper Dentition. In S\ baileyi sp. noy., the crown of 
V1 is slightly Jonger and broader than §, browne? wad 
is most stimulir to §. occidentalis, Weis not us high 
crowned us that of S. gi// and not as broad as i 8. 
anderson, 8. atlas (Owen, 1838). S. trrdalei or S. 
pales. The small, cvlindrieal 12 js iitermediate 1 
size bewween 4, browne? and 8. vecidentaliy, 13 ts 
Most SHNIRW H size all general morphology tos. 
frrovene? but the buceul surfave iysmoolh and That, not 
hearing any vertically-orientated undulations. [n this 
respeel, S. beilevé is similar to &. eccidernteliy ands, 
will. 

though slightly shorter aad less inflated huerally 
thin 104, browne, dP2 OF S. batlev? sp. ouy. is 
closest in overull morphology to that species. 
Onentalion of fhe buccal and lieu! crests is alse 
similir but the posterior basin appears to have been 
larger nS. Aeiewin PA of S. hatlevi is amtust 
reminiscent af S. hawnel and S. eartiqutis in 
morpholowy, partivilarly iy the shape and aricntiwon 


of the buccal and Jingual crests and the anterior basin 
(Fig. 7). However, 8 bailey? possesses a shallower 
and narrower longitudinal basin and a prominent 
posterobaceal accessory cusp with ‘two cuspules 
amlertor to it (Pig. 7). The posterior basin is smaller 
than in either S. aariquus or S$. browne. Height of the 
linghal crest in §, aitiquay is considerably lower 
relative io [he buccal crest than iy either S. browne! 
or §. bailey’. In addition, the §. ctiquus P3 is also 
smuller relative to the size of the molars. The amount 
of wear that dP3 has undergone has obliterated 
several characters useful for compurison. However, 
the tooth appears to have heen generally similar to 
that oF S. browner but with a smaller precingulum, 
larger premetucrista and many fewer and finer 
enamel erenulations on the loph faces and interloph 
valley: 

The very low crowned nature of (he S. heilevi 
upper molars ts only upproached among Sthemuerta, 
hy S. ceased, So antiguas and § prddocki. Similar to 
S. cegsad and §. aatiqats, there are few erenulations 
on the Joph faces and the interloph yalley but the 
postprotocrista is more Weakly developed in S. 


hailey, The postparactista 1s) more strongly 
i ry 
j es 
4 f . = - 
. —_ ea é 
5 = —_ - 
i : 
a] } 1. 
, a it 4 - 
— HT 
I 
a 
am, 


Pig, 7, Comparntive sketches oh len PS inpechisal view. A 


Tivcdronemas  pucktider Bo Sikes juirtipins 
(posterolinuiiil corer reconsiticted — ineaniplete 1 
awtal speenmen) CN. hoilevt sp. naw DOS Ayo 
Sele burs= 3 Wii, Abbrevs! d= unteriar t= lineal 


12 G. J. PRIDEAUX & R. 1. WELLS 


developed in S$. baileyi than any other species. 
including 8. hrawnei, The slight nature of the 
precingulum is similar to that of S. antiguas. 


Dentary, Th general morphology, the dentary of 8. 
baileyi sp. nov. is most similar to 8. occidentalis, 8. 
anriguus and 8. gilli, and to the former two in size. 
The ramus dilfers from §. eveidenralis in the 
following features: it is slightly narrower lor its 
depth, the symphysis extends only just beneath the 
genial pit. the il and symphysis ure slightly more 
procumbenot, the diastema is slightly longer, the 
divastric sulcus ts shallower and less extensive. the 
posteroventral border of the massetcri¢ fossa is more 
fared laterally and the pterygoid fossa is more 
inflated posteriorly. Sthenuruy baileyi differs froin §, 
antiguas uw its longer cheek tooth row relative to 
ramus depth. Morphology of the §. baileyi symphysis 
Most resembles that of S$. maddevki, where the 
symphysis tapers gently anteriorly and only extends 
slightly below the genial pit, However, unlike 4. 
meaddocki, the orientation of i} closely approximates 
that of the anteroventral border of the symphysis and 
in this respeer is similar to S, vceidentelis and S. 
browne’. Morphology of the iL crown and its degree 
of procumbency are intermediate between 3, 
vecidentalis aus. browne. Relative to the length of 
the ramus, the diustema of 3S. baileyi is 
proportionally longer than that of S. vecidentalls, 5. 
browned and S, ail/i, ets mast similar in length tos. 
maddacki bul is nol convex. dorsally as in this 
species. Depth and extent of the digastric suleus are 
similir to, but slightly more pronounced than in 8. 
meddock?, The degree of intraspecifie variation in 
depth and extent of the digastric sulcus also seen 
similar between (he lwo species. Literal expansion 
of the pusteravential border of the masseturic Tossa 
into wide Shell as similar tod. ceeseh So aly and §, 
maddocki, The pterygoid (ossa is more inflated 
posteriorly than aay other Siremumy species anu att 
his respeer, Y feadeve resembles Proceptodon, 

Size dnd morphology Gh dp2 most resembles that 
ot S. fie ied DUT Ss net us narrow unlerionky rohitive 
fo the posterior part of the tooth, The median valley 
moatso narrower Superficially, the dp2 tceal ens 
uppears Similar te fengthe no thalalS. Avo tal this 
Hipresvint es created because The wou sustaiedd babs 
resiicd Mm the erest becoming eontuent wilh the 
Sn cuspbles teins aiferiog dp3 is alse seine om 
size tnd ierplolaey fo that ots, Arewited bir the 
Erstit) ODT ia Ts Con Giet Yell Oe biypoconiied 
upes ine there cin Teweronamed crenulatioms on the 
tophid fees. In these ehanniiers he 8. bain ps 
more Closely resembles dat at \y aeeviteraeaten 


In morphology, stae relative to the olor. ame 
onenwiiion oF the tain und buccal crests. the 8 


hetles!d oF is Similan te Hat lS. geting Be TY 


Fig, Ko Comparative sketches of right pa in oeclusal view. A, 
Sthenurus brachyselenis, BOS. uanguus, CS. baileyi sp. 
noy. D. 8) browne/, Scale bars = 5 im. Abhreva: a = 
anterior, |= lingual, 


differs by being smaller, lower crowred, slightly 
nove inflated anteriorly and having a slaighter main 
crest orientated from the posterolingual to the 
anterobuiccal dormer of the tooth. bn S. adit, the 
posterior part or (he main erest wends. anterohuceally 
(hen straightens anteriorly along the tooth’s midline, 
The S. baflevé p32 shures with S. brachyselenix the 
major features OF the tum erest aod a tidgeled 
traversing the median valley bub is easily 
Jistinguished by abs Larger sive, shiglitly grearer widrh 
relative to length (ratio Q.55 conipaned with 0,52) ane 
longer. straighter buecal crest (Pig. &). da size wud 
general outline, the S. betlevi ps is also sinitiar to. 
browne! hut is lower erowned aid hears o 
cobsidenibly shorter buecul crest. 

li sive and crown height. the lower migtars ahs. 
baflev ewe somewhabs imino those ofS. sees aad 
S cviguis, Sut ave Most stint ta these at § 
Ivacliyveleinis They differ from the later ru ther 
wrealey WIL polative to length (roti O54 compara 
wilh (72) ah the aulerioy! cinguhin oer hens 
symmetrically tapered auleriorly wou the shell-ike 
inflioem ci jhe posterior fare of the hyputophued 
heli much fess pronounced, In general db. duilew 3. 
hrachvseloma, 8S. ceases and 3. iets huve 
relutively smooth lophidh Mees. with only a lew line 
cnamel enenulations 


NEW PLEISTOCENE KANGAROO A 


Diseussion 


Sthenuras baileyi sp. neve retains a sure of 
cramoadental characters that suggest a fiirly 
plesiomorphic position within the genus, Although 
the deposits trom whieh the species ts known are 
Pleistocene in ape S. Aeileve is most closely 
comparable with the Phioeene S$, airiqgilix from 
Chinchilla in southeastern Queensland and 4, 
bravhyselenisx, a species of uncertain age from 
Wellington Caves, eastern New South Wales. p3 is 
very similar fo So @atiyius, but considerably nvore 
denved than S$. hrachyselenis, given its greater 
robustness relative to tbe molars ang longer. 
straighter buccal erest. In &, brachyvelenis, pd is 
quite arrow and has a short, erescentic buceal crest 
resincted to its posterobyecal corner, features. which 
are considered plesiomorphic for the genus 
(Podeaux & Welly 1897). The lower molirs are 
Intenmediaie herween S. hrachyseleniy ands. 
auuiquus i gener) morphology, bat. unfortunately, 
no appee moles lor Wie Former are known, However, 
the upper molars af S. baileyi are very sumilur to 
those of S. aeriguis. Based on a comparison of single 
fipper Molurs. these two species would be diffieule to 
separate. However, P3 ts notably more derived in 3. 
hiulevi, the lingual cizulum having become raised 
into a crest subequal in height with the buccal crest. 
lu 3. cartigaus itis markedly lower. 

Although the only known cranium is incomplete, 
S. baileyi can be clearly distinguished from all 
species Of Sphemens for which the cranium is 
Known. While exhibiting a similar degree ot 
brachycephaly (o 8. occidentaliy, §. bailey? possesses 
a shorter. broader rostrum and a less inflated frontal 
reoron (hun any of the other brachyeephalic 
Pleistocene species, Ineredsed inflation of the 
Irontils appenrs to have co-evolycd with jneredsed 
check tooth coniplexity in the fineaye (ar passibly 
linvages) lowding to the nore briehyveephalie 
(shorter-faced) Species. eg. $. brane &. 
eecidentatis wl 8, nidddbekh The modest degree at 
fronrel inflation. relatively stmple Jow crowed 
inolars and short bucwal crest not joming (he main 
vhest aiteriorly on pd provide a conceryable 
auWeCdent merpholowy to phese other specres, 

Uuformunaely. only one ramus and one munilha 
fraginent of So civigers see Known bul given: Lhe 
Jentil somihiritics between this species and A 
boiler, Nhe Vikelthood amay be thut these reflect 
overall cranial srmihidies. Althoweh the dentary ol 
Soaiiques ds incomplete. ole inypartant differends 
ja the craniuon of this speeres and §, barlert taay be 
nuiened by the Jonger cheek tooth row relitiye to 
denny depth observed: in aetiquas (nd 242 
compared with 18S lor S. fades). Ths sugpesrsy a 
relatively longer dewlany und: Uerelon oa foie 


clongate cranium than fea S. baeeyy. This feature, tn 
conjunction with the shghily higher crowned molars, 
und more distinel ctisiid obliqua and parucristid. may 
make S. antiquis a possible structural precursor to 
the Jineaye that led to the more dolichovephaliv 
(longer-faced) Pleistocene species. This contention is 
supported by the fact that the ingualscrest of the §. 
antiguus P3 is notably lower than the bueciul crest. 4 
featute shared hy the more dolichucephalic species. 
In (he more brachycephalic species the erests. tend to 
be subequal in height. Since musing oF the lingual 
cinguluny into a erest is a synapomorphy for all 
sthenurines excluding the plesiamorphic late 
Miocene Hadnmomas puckridel Woodburne, 1967 
(Fig, 9), a lower crest muy be Tegarded is a more 
plesiomorphic condition 


Despite the celidnce on relatively limited Pliueene 
maternal, the suyilarities. between 8. bailev and Ss. 
antiquus imply a close relationship. They are more 
derived than Ss. cegved and §&. brachyselegis but more 
plesinamorphic than ony deserthed Pleistocene 
species. Features not shared with each other are 
either those shured with the more dolichocephilic 
species in the case of §, antiques, or with the more 
bruchycephalic species in the case of §. baileyi. We 
Tedford’s (1966) subgeneric (generie sensi Flannery 
1983) definitions hold (ie. Simayieniray = 
brachycephalic, low-erowped cheek teeth with low 
links and many coarse enantel erenulations, 
Sdicnurns sense stricto = dolichocephalic, high 
erowned cheek teeth with Strong links and few fine 
enamel! crenulations), then 9. andfguns may represent 
ihe least demyed species in the subgenus Sihenusts, 
while S bullevi may fulfil a similar position in 
Simosthenurny (Fig. 9). Beeause 8. nolabitis 
Bartholomai, 1963, au apparently derived 
Johchocephalic species co-Gccars. with So antégines 
in the Pliocene Chinchilla deposit, the diverzence of 
the shorter- and longer-faced sthenaurine groups must 
huve oeeurred much eurher im the Pliocene 
Similarly, very derived species co-seeur with Ss. 
fuileyi an the Pleistocene, but all that thie 
demonstrates is thal 8) aetiquits anc S. bailey? anc 
striiciural precursors to the doli¢chocephalic une 
brachycephalic lineages. rather than part ol Wret 
direct ancestry 

So given ther verisimiltude, are ihe dillerneces 
between §. udev? and So amtignas sufhetent 1 
Warrant phivement ji dillerent subgenera’! While 
they dee net possess muny of ie extreme character 
states Todfoed (MOGs uscd to deline the subeeneri, 
the questiau ts plywogenenedihy irrelevant so lors us 
Sritesuuriens ail Sehenireiy SA ae monophyletic 
The validity @f these tusa ty curreniy under 
Trvestiwanou Dy on of ts (GSP) and requires sine 
revision, sive (he Humber at descr beat erhenurine 


14 G. J, PRIDEAUX & R. T. WELLS 


Srenurus 
brachyseleniy 


Sthemuris 
eegsal 


Hadronomas 
puckridgi 


longer faced = Syhenuruy — Sthenuris — shorter faced 
Sthenurus spp. antiqnis baileyi Sthenurus spp 


Fig. 9, Possible phylogram of basal relations in the Sthenurinae, based on the following synapomorphies. |, Cranium 
relatively large, neurocranium flexed dorsally relative to rostrum: occiput close to vertical. broad and deep with well- 
developed lambdoid crest; large palatal yacuities, narrow post-palatine bars; deep jugal expansion torming ectoglenoid 
process; Jaterally expanded supraorbital crests; ectotympanic thick, wide, cancellous and yentrally-kveled: ascending 
ramus relatively vertical. with pterygoid fossa elevated and deep: digastric sulcus / eminence well-developed: 12 very 
small and splint-like; 13 dominated by buccal crest. lingual crest restricted to anterolingual corner: upper meisors form V- 
shape when viewed yentrally, C1] absent: p3 bears posterobuccal cingulum: molars fairly short relative width and squarish 
in occlysal view: molar lophs relatively straight and close to parallel; lower molars with posterior face of hypolophid 
inflated ventrally. 2. Rostrum broad and deep; zygomatic process of squamosal relatively deep; dentaries unkylosed at 
symphysis; mandibular ramus deep and wide, with depth at symphysis barely shallower than beneath molars: P3 with 
lingual cingulum raised into crest, separated from buccal or main crest by longitudinal basin traversed by ridgelets: p3 
with buccal cingulurn raised into crest, 3. p3 with curved buccal crest separated from main crest by wide median valley: 
p3 widened posteriorly: molars with more fine enamel crenulations. 4. p3 wider overall relative to length, with longer 
buceal crest: lower molars with crisud obliqua and paracristid shifted more lingually, 5, Cheek tooth row long relative to 
ramus depth, higher crowned molars, more prominent cristid obliqua and paracristid. 6. Cheek tooth row short relative to 
rumus depth, brachycephalic: retained lower crowned molars, low cristid obligua and paracristid. 


species has roughly doubled since Tedford’s (1966) 
review. Almost certainly, §. cegsai and 8. 
brachyselenis have va place within the two 
subgenera because they lack muny of the delimiting 
character states and appear to be the earliest 
derivations from the sthenurine lineage, post- 
Hadronomas piuckridgi (Fig. 9). We await the 
discovery of further Pliocene species to confirm 
exactly where S. cegyeai and 8. Arachyselenis fit 
within the sthenurine radiation. As more taxa 
become available more light will inevitably be 
thrown on this paramount phase in sthenurine 
diversification, 


Acknowledgments 


We thank N. Pledge and J. McNamara (South 
Australian Museum). R. Molnar (Queensland 
Museum) and R. Jones (Australian Museum) for the 
loan of specimens. L. Hatcher loaned the Western 
Australian specimen for comparison. J. McNamura is 
sincerely thanked for proyiding data on the type 
locality, preparing the holotype and his characteristic 
perspicacity when commenting on an early draft of 
this paper. We are alsa grateful to J. Long and M. 
Atcher for their constructive criticism of the 
manuscript. 


NEW PLEISTOCENE KANGAROO | 


a 


References 


Avtibhn Lo K., MARIANELI. PLC. MeCuntocn, M, T. 
Mortimer. G. E., HELLSTROM, J. C., Moriarty, kK. C. & 
WrLts. R.T. (in press) S00 ka preeipitabon record from 
southeastern Australia: evidence for interglacial relative 


undily. Geclowy. 


Brows, H. Y,L. (1908) Bone breccia and rock phosphate at 
Brothers Islands p. 343 de “Records of the Mines of South 
Australia, 4th edn” (Government Printer, Adelaide), 


FLANNERY, To FE 


(1983) Revision in The macropodid 


sublamily Sthenurinae (Marsupiala: Macropodoidea) 
wad the relationships of the species of Tropasodon and 
Lagastrophus. Aust. Marimet. @ 15-28. 


(1984) Kangaroos: 15 million years of Australian 


hounders pp. 817-835 Ja Archer, M. & Clayton, G. (keds) 
“Vertebrate Zoogeography and Evolution in Austria” 


(Hesperian. Perth). 


Fiownr, W. H. (1867) On the development and succession 
af teeth in the Marsuptala, Piri. Trans, R. Sec. Lond. B 


157, 631-641. 


Lucnhnti, W. PF (1993) An ontogenetic as 


ment of dental 


homelogies in therian mammals pp. 182-204 fa Szalay. 
KS, Novacek, M_ J. & MeKenna. M. C: (Eds) “Mammal 
Phylogeny® (Springer-Verlag. New York), 

PRIpb AUX, God. & Wares. R. T. (1997) New Sthenarus 
species (Macropodidae, Diprotodontia) from Wellington 
Caves and Bingara, New South Wales. Prac, Linn Sec. 


NSW 1E7, ISL-L96. 


Rich, PV. (1979) The Dromornithidae, an extinet family of 
large ground birds endemic to Australia, Bie Min. Res. 
Bull, 184, 1-196, 


Rink, W. D. L. (1993) Jackmehoneva pen, nov, and the 
genesis of the macropodiform molir, Mem. Ass, 
Australas, Palaeontely 15, 441-459, 


TEprokD, RM. (1966) A review of the niaecrapodid genus 
Sthenurus. Univ, Calif, Publ. Geel, Sei. 87, 1-72. 


— & Woopsurknrt, M, O, (1987) The Uartidae, a 
new family of vombatiform marsupials from Miocene 
strata of South Australian and ain evaluation of the 
homology of molar cusps in the Diprotodontia pp. 401- 
418 da Archer, M. (Ed.) “Possums and Opossums: 
Studies in Evolution” (Surrey Beatty & Sons, Sydney). 

Wetts, R, T. & Murray, PF (L979) A new sthenurine 
kangiroo (Marsupiala, Macropodidae) fram south- 
eastern South Australia. Trey. R. Soo. 8. Ast. 103, 213- 
219. 


. Mortarty, KL & Witiiams. D. L. G. (1984) 

The fossil yertebrate deposits of Victoria Fossil Cave 

Naracoorte: an introduction to the geology and fauna, 
Ausi, Zool, 21, 305-333. 

Wiitams, Do L. G. (1980) Catalogue of Pleistocene 


vertebrate fossils and sites in South Australia. Trans. R- 
Soc, S. Aust. 104, (01-115. 


Appendix 


Material used for comparison with S. baileyi. See “Introduction” for abbreviations, except AM = Australian Museum, QM 


= Queenslund Museum. 


Species 


Sthenurus antiqtys 
S. brachyselenis 

S. browned 
(eastern form) 


S. cegsat 


S. gilli 


So mnaddocki 


S. acetdentalis 
(custern form) 


S. orecn 


S. pales 


Registration Number 


QM F293]. F2973 
AM F31026 


SAMA P2043, 
FU 0202, FU 0271 


SAMA P31800 
(tiolatype) 


SAMA P16528, 

P16629, P2()797. 
FU 0246 

SAMA P16627, 

P16643. P16G73 


SAMA P20798, P27799 


QM F2923 (holotype) 
SAMA P27797 


Locality 


Chinchilla, Darling Downs, Qld 
Wellington Caves, NSW 


Victoria Fossil Cave, Naracoorte, SA 
Corra Lynn Cave, Curramulkia, SA 


Victoria Fossil Cave, Naracoorte, SA 


Victoria Fossil Cave, Nuracoorte. SA 
Victoria Fossil Cave. Naracoorte, SA 


Darling Downs, Qld 
Victoriit Possil Cave, Naracoorte, SA 


MACROINVERTEBRATE ASSEMBLAGES OF 
GOYDER LAGOON, DIAMANTINA RIVER, SOUTH 
AUSTRALIA 


By FRAN SHELDON* & JIM T. PUCKRIDGE* 


Summary 


Sheldon, F. & Puckridge, J. T. (1998) Macroinvertebrate assemblages of Goyder 
Lagoon, Diamantina River, South Australia. Trans. R. Soc. S. Aust. 122(1), 17-31, 29 
May, 1998. 

The wetlands in the arid zone of Australia have considerable significance as drought 
refuges. Despite this their biology is poorly documented. Goyder Lagoon is an arid 
freshwater wetland in the driest region of Australia, the central Lake Eyre Basin, 
South Australia. 

The abundance and richness of macroinvertebrates was examined at 11 sites within 
Goyder Lagoon. Insects, comprising 76% of taxa and 63% of individuals, dominated 
the assemblage. The prosobranch gastropod Thiara balonnensis (Smith) was the most 
abundant taxon, the prawn Macrobrachium australiense (Ortmann) comprised the 
greatest biomass. 

Key Words: Macroinvertebrates, semi-arid river, functional feeding groups, Goyder 
Lagoon, variability. 


Treinsaetions af ile Reveal Savierny ofS. Aas (199%) $2207), 02-31. 


MACROINVERTEBRATE ASSEMBLAGES OF GOYDER LAGOON, 
DIAMANTINA RIVER, SOUTH AUSTRALIA 


- HF py af 
by FRAN SHELDON & JIM T. PUCKRIDGH 
Sunimary 


Stiecuoy, B& Prccaipoe, 21) ( 1994) Mavcoinyertebrate assemblages of Goyder Liaoon, Didukintiia River, 
South Australian Tas, RoSed, S. Anse 12201), 17-3129 May. 199%. 

The wetlunds in the wie zone of Austialia have considerdble significance a4 drought refuges. Desptic this (heir 
biology is poorly documented. Goyder Lagoon rs an arid freshwater wetlind in in the driest region Ol Ausialia 
the central Lake Eyre Basin, South Australia. 

The abundance and richness of macroinvertebrates was examined at TT sles within Goyder Lagoon, Insects. 
vomprismye 760% of taxa and 63¢¢ of madiyiduals. dominated the assemblage. The prosobranch gasteupod Titan 
batonnensis (Siwith) was the mostubundant een, the prawe Membrchinncaustralieive (Orimang) comprised 
the grenlest biomass, 

Collectors were (he dominant fecding group ueross all habitats with tbe ridos of different Functional Feeding 
Ciroups (FRCS) suggesting that Goyder Lagoon is heterotrophic, dependent an allochthonous organie matter ys 
iLcurhon source, 

Multivariate analyses separated temponiry fron) perinanent habitats, The prawn Af ausirafiense, ihe yabbie 
Cheray destructor (Clark), We ephemeropterin Tagmaneceeniy arecata AtThu-Tercedor S& Suter anc the 
Wichppreran Bones. sp, dominuled at frequently inundated sites whereas infrequently inuridated Sites. Were 
dominated hy the nomnectid Aviiharey sp,, and the curinids Micrmiecte spp- 

There were suiking diNerences in assemblages trom different sites, even those From (he samme mesohabiiat type 
channel, Witerhole or billabong), ‘This may reflect diferent successional (rajectories wilhin the assemblage at 
cul site afler hydrological isolution. This supports the hypothesis that the variable flows characteristic of 
Gover Lagoon ace inporkint for mamtaining macvolnvertebrate diversily, 


Key Worps Macrainvertebrates, semi-arid river. funetional feeding groups. Goyder Lagoon, variability 


Introduction 


The establishment sind maintenance of specific 
types of wetlands amd wetland processes are 
controlled, it least in part, by the prevailing 
hydrological revime (Mitsch & Gosselink 1986). 
Although it sounds contradictory, wethuids do occur 
in and landscapes. Dryland rivers and their 
assocrated wetlands have a number of unique 
hydrotoical features (Molles & Dahm 1990; Walker 
etal, 995; Puckridge er al 1998). The dominant 
hvdrolovieal rhythms of drylkimd myers are not 
seasonal or annual but vefleet. in part, large weather 
phenomena such as the El Nifio Southern Oscillation 
(NSO) (Walker.e/ af. 1997). In the Australian arid 
fone, significant freshwater Wetlands. are mainly 
associated with nver channels wre floodplains and 
intermittent river Flow ensures that these wetlands do 
not accumulate sult from year to year und beeurne 
silt lakes (MeComb & Luke 1988). 

Wethids are amongst the world's most productive 
ceysysiems. Arid zone wetlands have consiteruble 
rewional significance for migratory birds (Breen 
{9Ul> Kingsford 1995; ANCA 1996), Despite this, 


Cooperitive Rescurch Centre Tur Prestivuter Ecology 
Departed of Zandogve Pie Unteeyerey ob Adelayde Aust. SUG 


their biology is poorly documented, in generul 
(Breen J991) und almost nothing ts Known of 
Australia's acid freshwater wetlands (Puckridge in 
press). 

This study describes the macroinvertebrate 
assemblage composition ul eleyen sites. within 
Goyder Lagoon. an arid freshwater wetland on the 
lower reaches of the Diamantina River. Lake Eyre 
Basin, South Australia (Pig. |), The Diamantina 
River system is one of the major wetland systems mn 
Austrutia thar remains substantially unmodified by 

auerresouree development and. as such. is likely to 
harbour w relatively pristine brota (ANCA 1996), 
Little, however, is known of the pracroinvertebrate 
fauna of any of the rivers in the central Lake Eyre 
Basin, Puckridge & Drewien (J98S%) and Rei & 
Puckridge (1990) list same of the taxa found in the 
Coongie Lakes system but there are no published 
data on the macroivertebrates of Goyder Lagoon, 
The highly variable Mows in Goyder Lagoon are 
likely to produce a diversity oF habitat types with a 
rutize of inundation frequencies and hence a 
spectrum oe different habiluts for aquatic 
invertebrates over time (Boulton in press), 

Multivariate analyses were used to identity the 
environmental yvariibles structuring the assemblages 
ai euch site and for different habiuat types. ‘The 
Functional Feeding Group (FFG) composition 
(Cummins & Klug 1979) of aysembliwes at cach site 


18 F. SHELDON & J.T. PUCKRIDGE 


Diamantina River, Birdsville 


one G6 


Eyre Creek — Diamantina River 
pshiplor Clifton Hills Outstation 
Pelican Lagoormh, + /ADHS Gr EGS 


PAIS 


Koonchera Dune 


G4&G5 
’ 
p CHHS 
Pootha Pootha 
Yelpawaralinna Ss G10 & G11 
G9 
We 
0 50 100 


km 


Fig. 1. Position of sampling sites (cf. Table 1) in Goyder Lagoon and on the Diamantina River. The heavy line marks the 
boundary of Goyder Lagoon. ADHS = Alton Downs Homestead; CHHS = Clifton Hills Homestead. 


MACROIN VERTEBRATES OF GOYDER LAGOONS 14 


and in different habitats is explored with ratios of 
different FPGs Used to mdieate ecosystem altrihdtes. 
Miributes such as the rekuive jmportance of 
fictevotrophy or aulolrophy (Suggested by 3 
domimunee of cither dearivarous eollectors or 
herhivafous: scrapers) und the relative amounts of 
Course Particulate Organic Matter (CPOM) and Fine 
Particulate Organic Matter (FPOM) in transport or ia 
slonige (suggested by the presence of filtering 
collegtors: Gr gathering collectors) (see Merrit & 
Cummins (996) were examined. Goyder Lagoon 
may he similar to (he Chow iki wethind of the lawer 
River Murray, South Austatat where colleetors 
domimuted The assemblige suggesting ollovhihaneus 
ordavie Cirbon was Ldeniinint Hood source (Boulton 
A& Tayi 997}, 


Materials and Methaeds 


Sry aired 


Goyder Lagoon is a 3030 kt inernddenr ternvil 
Comin & Williniis 1994) floodplain wethind sit ihe 
rerminus OF the Diaiantini/Georg ii river systen 
whieh has uo cachmenr aren of 365,000 ki (Pies. 1), 
Phe Diamantina is ane ol the most hydrologiowlly 
variable sven systems ih the world | Pochrupe er af, 
}9U8) Govder Lagoan beains 80 km below thie kiwi 
of Birds wille where fhe Diuinidna River splits ite 
an easter and a western unabraneh. These barnches 
then radiate into a redieuhite system of fine channels 
(o form the castern und western) intechal deltas. The 
custern internal deli ts larger and more frequently 
Hooden, Water thal makes ils way (hrough the maze 
of Cioyder Lagoon draiims to Lake Eyre vin Warburton 
Creek. The lagoon is bounded hy the dunefelds of 
the Sampson Desert to the west und northwest ane by 
the gibber plains ol Stiets Stony Desert to the cast. 

The survey wus conducted during November 1993 
when wuler levels in the laoon were low. the fast 
pion flodding having oevurred in nid L991. AU bot 
the deepest waterholes Were dry dnd sampling was 
(herefore Confined lo these waterbodies. In the period 
JURS-1993 [lows occured samewhere in the lagoon 
Y oul ol LO yours (bandsat dati; South Agstealian 
Department for Eovironment. Hertlaye and 
Aboriginal Affairs). Flouding Crequency ane Cloned 
posicdence (imes wppeien Tram the Landsat record. 
diminish in the ligeon downstroam, and ins the 
northern fagoow tron) east la west. The inundation 
frequency oF cach site within the lagoon was ranked 
using an estimate from the overall gradien( in 
inundation frequency evident in the Landsat record 
(1986-1993) und cheeked with loeal Jandowners’ 
knowledize. 

Sites were chosen 


represent abo range of 


hydrological conditions and a spatial distribution 

aluog the lagoons axis (Fig. 1), In order of 

inundation frequency. snes meluded: 

* the permanent channel of the Diamantina Rivet 
upprosimately | Km dowastreat from the town of 
Birdsville (GG), 

* the pernunent channel of the main (northeasiern) 
branch of the Dianmntina in Goyder Lagoou at 
Chitton Hills Outstation (G7) and w closely 
associated backwater (GS), 

‘un isolated waterhole at Andrewilla Station on it 
semi-permunent segment of the northwestern 
branch (G2), closely ussocnited backwater (Ci). 
and a onearby isokued billabong, Pelicun Lagoon 
(G3). 

‘4a deepened but impermanent portion of the 
reticulate channel system ol the eastern dele 
formed by the iitrusion of Koonchera Dune (G4. 
GS), 

eo segment of the less frequently tlowded 
Warburton chanel io ahe southern dagounm, Poolttia 
Pootha Waterhole iGl0). und ai associutead 
billabong (G11), 

+ un isoliled waterhole on a tess Hequently Mlooded 
wabraneb of the Warburton downstraant of GT, 
Yelpawuralinna Waterhole (G9). 


Aquitic’ systems such as Goyder Laguon can be 
desided Tote maero-, mesa. tind miero-habihuts 
(Walker ef af }9US). Lisuie fis hiertirehical system. 
Goyder Lagoon is a distinet mycrohabitat. Within 
Unis maerohubitut exist mesohabitats such as 
channels, waterholes aid billaboogs and within hese 
ae Microlabitits such ws emergent and submerged 
vewelation, submerzcd wood aad other substrate, 

In this study four mesohabitit types, channel. 
witerhole, backwater and billabong were sampled 
(Table 1). Overall microhubitats tneluded emerent 
vegetation such as sedges (Cyerky spp.) and semi- 
aquatic grasses (eg. Cyredon spp.) Torming 
sometimes dense stuns along the ed@es ot tre 
waterbodies, aquatic macrophyles (ee, Colvecuias 
sp. Miriepivilon spp. submerged woody debris 
(snags') represented by the roots or Lullen lambs of 
trees Sue os river redguntm® (Avelyn 
cimeldileasis Delinh. var ebtuse  Blakelys 
Coolihah (Eyvealpai colihah Blitkely & Jacobs 
ssp arid (Blukely) b, Johnson & K. Hill) and River 
voobah (dca ia verephyla Cun ex Benth), course 
particulate organie matter such as packs of lea! litter 
und (wigs From riparian vegenition, and unvegedited 
littor) arews free of veeetition, woody detritus oF 
other cover. The latter were further divided into silt 
and clay suBstaitum, Sandy substratum: or rok 
substratum, Ninely (wo sumiples were collected from 
microhabitals present within the TT sites (Pig t, 
‘able 1). 


20 PF. SHELDON & J, T. PUCKRIDGE 


Pabur |. Lise of sites (ane their abbreviations) wrauped by niesohabitar. 
rhe number of samples collected from each microhabitat at each site ts alsa given. 


Mesohabitat Site Site Name Microhabitats No. 

Number Samples 

Backwater Gl Backwater of Andrewilla Waterhole Leaf litter 4 

Silt 4 

GH Backwater of Clifton Hills Waterhole Polygonum 3 

Lignum 4 

Silt 3 

Waterhole (2 Andrewilly Wilerhole Rock 4 

Sill 8 

G4 Koonchera Dune Walerhole Sand 4+ 

(caster portion) Sill 4 

GS Koonehery Dune Waterhole Lignum 4 

(westem portion) Sand 4 

Sill + 

G7 Clifton Hills Waterhole Polygonum 4 

Lignum 4 

Silt 5 

Shag 4 

ci9 Yelpawaralinnit Watterhole Emerg. vey 4 

O10 Pootha Pootha Waterhole Emerg. veg 3 

Silt 3 

Channel Go Main channel of Diamantina River Polygonum 3 

(Birdsville) Grass 3 

Rock ‘f 

Silt 4 

Billabong G3 Pelican Lagoon Billabong Silt 2 

Gil Small Billabong Silt I 


Sample collection und processing 


Each site Was considered a distinct mesohabitat 
und was stratified into the microhabitats defined 
above (Table 1). The most prevalent of these were 
then sampled randomly (Boulton & Lloyd 1991), 
Macroinvertebrates were collected in replicate 
samples from each microhabitat by sweeping a SOO 
um mesh pond net ever 5 m-* for 20 sec, Samples 
were preserved in 70% ethanol and later washed 
through nested sieves (4000, 2000, 1000, 250 pm) 
and the organisms hand-soerted, enumerated, and 
identified as far as practicable, Unidentified 
specimens Were recorded as separate taxa (e.g, “tiny 
Zygoptera’”), Each taxon was assigned to one of four 
broad functional feeding groups (FFGs): collectors, 
predators und serapers (Cummins & Klug 1979), 
with collectors incorporating, filterers and gatherers. 
The assigned FPPGs, particularly “collectors”, ure 
tentative as the diets of most taxa are unknown; 
assumptions uboul diet were therefore based on 


MBollros, AW. (1988) Composition and Dynamics ot 
Macromyerebote Communities in “Pwo Intermittent Streatys. 
PhD thesis, Monash Uniwersity CGunpub.y 

2 Seen A (}994) Lattetal Eeology ofa Repuluted Dryland River 
(River Murray, South Austratia) with Reference to the Gastyapoua 
PHD thesis, Unversity of Adelaide (unipub ). 


taxonomic affinities. Classification of taxa int 
functional feeding groups followed Cummins & Kluy 
(1979), Boulton (1988)', Boulton & Lloyd (1991) 
und Sheldon (1994)") At each sampling site. spot 
measurements were recorded of depth, Séevhi 
lranspurency, temperature, dissolved oxygen CYS] 
oxygen probe) and conductivity (Hanna HE 8733 
conductivily meter): sulinity was computed trom 
temperature corrected conductivity values using the 
equations in Williams (1960). 


Data analysis 
Hierarchical analysis of variance (Underwood 
IO81) was used fo explore patterns in fichness and 
abundance between mesohabital und microhabitat 
scules. As the sampling design was unbalanced, 
subsets of the total dataset were used, Subsets were 
chosen so there were equal replicutes within each 
level making the design balanced. The lollowing 
scpurate unalyses were performed: 
i. Microhabitats (emergent vegetation, sill) 
bested within sites (G6, G7, G8, GIO), 
i. Microhabitats (ignum, silt) nested within 
sites (G5, G7, G&S). 
All analyses were performed in SYSTAT vy, 5.03 
(Wilkinson 1990), Data were rendered normal by 
(ranslorming using log ),(X + 1). 


MACROINVERTEBRATES OF GOYDER LAGOON Zz 


Hierarchical patlerns in the dita were also explored 
using inultivariate statistics from the PATN software 
puckuge (Belbin 1993). Multivariate analysts allows 
patleriis. between samples to be explored. ‘lwo 
multivariate approaches were used: classifieation (oy 
derivation of discrete proupings of samples) and 
ordination (arringement of samples ina space ot a 
few dimensions). Both techniques urrange samples 
on jhe basis of their species conjposition, with those 
samples that cluster or group, together more likely to 
be similar in species composition (ler Braak 1947). 
Data were log,,(% + 1) ininsformed before analysis 
and then range standardised as suggested by Belbin 
(1993); the Bray-Curtiy coefficient Was Used as. the 
measure of dissimilarity between samples. AIL taxa 
occurring in one OC Mure Samples were retained in 
the analyses. Mlexible-UPGMA was used to cluster 
the samples, with the ANOSEIM procedure (Clurke 
1993) used to test for significunt clusters of samples 
ata meso- and microhabilal scale, The SIMPER 
procedure Trom the PRIMER software package 
(Clarke & Warwick 1994) was used to identify the 
percentage contributions of different taxa to the 
simple groups. Semi-Strong Hybrid multidimensional 
scaling (SSH) ordinations were also computed trom 
the Brey-Curtis similarity matrix generated [rom the 
transformed and standardised data: setuugs of 30 
itenutions and J ratio-ordinal cut value of 0.8 over 
100) random starts was used, Solutions were 
calculated in two, three and four dimensions. The 
solution presented here had a stress of 0.18, The 
Stress ineasure gives an indication of the “goodness- 
of-lit” for the ordination (Kruskal & Wish 1978); an 
ordination with a good fil fur sumples within the 
ordination space basa stress value of less than 0,2, 
Sample groups from omesohubitats. sites and 
Mmicrohabitais were mapped on to the ordination 
plots. 

Rehidionships und 


between environmental 


community data are usually many, conrplex snd 
nonlinear (Gauch 1982). One aun of condueting 
multivarigte analyses is to detect major differences im 
Species composition for sample groups which are 
potentially related to environmental differences 
(Boullon 1948)', Spearman Rank cornlavons (Zar 
1984) between the physicochemical variibles 
measured in the Held and the ordination scores on the 
SSH axes were calculated 10 examine relationships 
between environmental factors and macroinvertebrate 
ussemblige composition, 


Results 


Environmental conditions 


The environmental conditions for all sites are given 
in Table 2. The Ditumuntina River earries a high load 
of yery fine suspended sediment with purticle 
diameters of less than | pm. This contributed to 
Secchi depth measurements well below 20 em at all 
sites, Salinites were highest at those sites in the 
centre of the Lagoon which were the most 
chsconnected from any recent flow of water, sites on 
Andrewilla Station (G1 and G2),at Koonehera Dune 
(G4 und G4) and Pelican Lagoon Billabong (G3). 
Pootha Pootha Waterhole (GIO) at the southern end 
of the Lagoon had a lower salinity, probably the 
result of recent rain_ Sites on the main channel of the 
Diamantina (G6) and Clifton Hills Outstation (G7 
and G8) also had relatively low salinities due to a 
recent small flow through the system. Al sites were 
well oxygenated (ncur saturation) with imstantineous 
Avaler temperatures manging from 20.34° C. 


Parterns of richness dnd abundance 

A total ef 7.363 individuals from 54 taxa was 
collected in. 92 sumples (Appendix), Insects were the 
dorminant group comprising 76% of (axa andl 63% of 
individuals (Fig, 20). OF the Insecta, the Diptera 


TabLE 2. Environmental conditions measwad in the livaral zene of each site in Gowler Lagoon, Diamantina River South 


Atuyinlia ta Noventber 199), 


Datawere not uvailable tof sites Go and G11 - n/a = nol avnilables 


Site Sample Depth Salinity 

(em) (moh't 
Gl Fd 309 
(2 Oi 26) 
Ga aM) 8265 
Ga 57 (299 
Gn 4a 2133 
Gin Wa wa 
G7 62 172 
Gs 74 }2> 
Gy 63 769 
G10) as 142 
ml al no 


Seecht Temp Oxyyven 
(em) (°C) (Se Saluralion) 
7 Wa ou 0) 
6.0 230 ony 
20.0 34.0 V0 
40 227 ORD 
40 24,7 WAY 
Wil ws ta 
a0 26,0 KA 
ay 25,4 THs 
40 Td) Sho 
My 716 nl 


nia oil mi 


22 PF. SHELDON & J.T. PUCKRIDGE 


comprised 39% of taxa and 41% of individuals (Fig. 
2b), The prosobranch gastropod Thiara balonnensis 
(Smith) was the most abundant taxon, with the 
freshwater prawn Macrobrachinm australiense 
(Ortmann), the corixids Micronecta spp.. the 
predatory chironomid larvae Coelopynia sp. and the 
predatory caddisfly larvae Oeverty sp. also common. 
The prawo M, ausrraliense comprised the greatest 
biomass. Five taxa occurred only once. Observed at 
Koonchera Dune Waterhole, but not collected, was 
the freshwater crab Holthuisiena (Austroielphiuse) 
transverse (Martens), 

Both the number of taxa (richness) and the number 
ol individuals (abundance) differed between the 
inesohabitars (Fig. 3). Backwaters and waterholes 
had more individuals than the channel or billabongs. 
However, richness was similar for all mesohabitats, 


When the mesohabitats were split into siles there 
were differences between sites from the same 
mesohabitat. Of the backwaters, GI had many more 
individuals and a greater richness than G&, Similar 
differences occurred between the billabongs G3 and 
G11 and there was also variation in both richness and 
abundance for different waterhole sites (Fig. 3), 
Numbers of taxa und individuals also differed 
between microhabitats depending upon the site, 
Large numbers oF individuals were found in leat 
litter packs in GI and silt microhabitat trom beth C4 
and G5, fewest individuals were found in snag and 
silt microhabitats from G7 and the rock microhabitat 
from G6. Emergent vegetation and submerged aress 
of hgnum contained the greatest number of taxa 
while the fewest number of taxa were from snug 
microhabitats. 


Bivalvia 
(CD Gastiopeda 
O Oligochaeta 


WB Hirudinea 
Ej Crustacea 
[J Insecta 


(_]Ephemeroptera 
| Odonata 
3) Hemiptera 


ff] Coleoptera 
Diptera 
(]Trichoptera 


Fig. 2 fi Percent representation of the major invertebrate groups in samples from all sites within Goyder Lagoon, 


na? 


mber 1993, in terms of the number of (i) taxa in each group and (if) abundance of individuals. (b). Percent 


representation of the major insect Orders in samples from all sites within Goyder Lagoon. November 1993. in terms ol 
(iil) the number of taxa in cach Order and (iv) the abundance of individuals, 


MACROINVERTEBRATES QF GOYDER LAGOON 


Mean numbers of individuals (FP, , 6.246, 
p>0.05) did not differ between sites G6, G7, G8 and 
GIO bot mean numbers of taxa (Fy, = 12.178, 
p<O0.01) did with GIO having more taxa than the 
other sites (Fig. 3). Mean numbers of taxa (Fy), = 
Q.586, p>0,05) and individuals (Fy), = 1.607, 
p>0.05) did not differ for emergent vegetation und 
Sut microhabitals nested within these sites. There 
were also differences in mean number of taxa (FP, = 
148.08. p<0.001) and individuals (FP), = 103.47, 
p<0.00T) between the sites G5, G7 and GS, with GS 
having more taxa and mdividuals than the other sites 
(Fig. 3). However, again there were no differences 
for either taxa (Vy), = 0.192, p20.05) or individuals 
(Fy, = 0.603, 750.05) between lignum and. silt 
microhubiliaty nested within the sites. 


Multivatiite dadlysis of samples 


UPGOMA classtlieation of the sample data 
indicated Wo main groups, The majority of samples 
foie the more temporary habitats (Cel, G2, G3, G4, 
G5, GY. G10) lonmed one group (A-D) while those 
from more permanent habitats (G6, G7 G8) formed 


8 


Abundance 
8 


250 


g 


(b) 


Wa 


rs) 


Sit 
Lignum 
Silt 
Silt 


Leal Liter 


Polygonum 


(c) 


Nig. 3 


23 


the other (E-H) (Table 3, Fig. 4). The first group (A- 
D) further split into Group A-B, containing samples 
from the western anabranch of the lagoon (G1, G2) 
which is the more frequently inundated of the 
temporary habitats, one sample from G6 and one 
from G7, and Group C-D containing samples from 
the extreme temporary habitats on the lower section 
of the eastern anabrinch and the southern end of the 
lagoon (G4, G5, G9, GIO). The second group 
separated into Group E-G containing a majority ot 
sumples from Clifton Hills Outstation on the upper 
section of the eastern anabranch (GI and G&) and 
Group H containing most of the samples from the 
permanent channel of the Diamantina River al 
Birdsville. The single sample [rom the infrequently 
inundated billabong associated with Pootha Pootha 
Waterhole, GIL, was an outlier clustering with the 
nore permanent sites in groups A-D, 
SIMPER analysis showed thal the prawn 
Macrobrachium australiense, the yabbie Cheray 
destructor (Clack), the ephemeropteran nymph 
Tasmanoeoenis. arcuata Alba-Tercedor & Suter and 
the trichbopreran larvae Leveitiis sp, dominated the 
asseinbluge in the more frequently inundated sites, 


Ute | 
eee peas Channel Billabong eee 
= Mile = Y 


| Whoa! 


G3 oe G5 GIO Gif 
Silt Emerg. 
Veg. 

5 80 Bo mK 

unui a | 

EgE= oO 2 = uz = 

§88a 3 5? & § i ge% 

8 § 5 a 

& 


. Mean (486) of richness ind abundance for each of the three sampling scales. (a). Mesohahitats. Uh}, Sites. within 


Ealulontuat (c). Microhabitats within sites, collected from Goyder Lagoon, Novernber }993, Points indicure richness 


und bars abundance, 


24 


F, SHELDON & J, T. PUCKRIDGE 


TABLE 3, Flexible-UPGMA groups by site, mesohabitat and microhahitat. 


Those taxa contributing more than 5% to the similarity of the sample groups are also indicated. 


UPGMA No. 
Group Samples 
A 21 
B I 
Cc 23 
D 1 
E 8 
F 22 
G 2 
H 5 
z 
S 
a 
> 06 5 


00@°e@,. 
@r-eee @): 


® @ 
o 7m 00 > 


No, Samples 


from Sites 


7Gl 
12 G2 
| G6 
1 G8 


1 G7 
2G3 
1G7 
12.G5 
8 G4 
4G9 
6 G10 


Snag 


0.8940 


22 


22 


No. Samples 
from 
Mesohabitats 
8 Backwater 
12 Waterhole 

1 Channel 


1 Waterhole 


2 Billabong 
21 Waterhole 


10 Waterhole 


3 Backwater 
4 Channel 
| Waterhole 


4 Channnel 
11 Waterhole 
6 Backwater 
1 Billabong 


2 Waterhole 


5 Channel 


1.0132 


No, Samples 
from 
Microhabitats 


4 Organic 


Matter 
13 Silt 
4 Rock 


| Polygonum 


1 Silt 
8 Sand 


4 Lignum 


4 Gruss 
3 Sedge 
3 Silt 


3 Silt 
| Grass 
| Snag 


3 Polygonum 


1 Rock 
2 Snag 


7 Lignum 
4 Palygonium 


6 Silt 
2 Grass 
2 Silt 
3 Rock 
1 Silt 


| Polygonum 


1.1324 


1.2516 
' 


Contribution of taxa to sample 


group (SIMPER) 


Thiara balonnensis 
Coelopynia sp. 
Micronecta spp. 
Macrobrachium 
australiense 


no taxa 
Oevcetis sp. 
Coelopynia sp. 
Bezzia sp. 


Enithares sp. 

M. australiense 
Micronecta spp. 
Coelapynia sp. 

Anisops spp. 

M. australiense 
Austrogomphus australis 
Cherax destructor 


M. australiense 
Tasmanocoenis urcuata 


no laxa 


Ecnamus sp. 
M. australiense 


1.3708 1.4900 


57.6 
40.6 


Fig. 4. Flexible-UPGMA dendogram, using Bray-Curtis dissimilarity, of the 92 samples collected from Goyder Lagoon in 
November 1993. Entities in sample groups A-H are listed in Table 3. A pictorial summary of the microhabitat 
representation of samples is also given. Large circles depict greater than 70% of the samples from the microhabitat occur 
in the sample group, medium cireles 30-70% and small circles 1-30%, 


MACROINVERTEBRATES OF GOYDER LAGOON oF 


G6, G7, G8 (Table 3, Fig. 5). The prosobranch 
vastropod Thiara balommensis, predatory tanypod 
lurvae Coelopynia sp. and corixids Micronecia spp. 
dominated samples from sites at Andrewilla (G1, 
G2) whereas the predatory tuichopteran larvae 
Oecetiy sp. and Coelopynaia sp. dominated samples 
from sites at Koonchera Dune (G4, GS) as well as the 
temporary billabong G3, The infrequently inundated 
sites GO and G10 had an assemblage dominated by 
the highly mobile predatory hemipteran Enitheres 
sp.. the prawn M. australiense and corixids 
Micronecta spp. (Table 3, Fig, 5). 

Ordination highlizhted the differences between 
Thesuhabitats and between sites. When grouped by 
mesohabitat, those samples from backwaters. were 
dispersed across Axes | and 3 and grouped low on 
Axis 2 (Pig. 6). Samples trom waterholes were 
dispersed across all three axes whereas samples from 
channels grouped centrally on Axes | and 2 and low 
on Axis 3. Billabong samples were central on Axes | 
und 3 aind grouped high on Axis 2. When samples 
were grouped according to sampling site, their 
disthbution along Axis 2 refleeted the inundation 
frequency of the site (Fig. 7). Samples from the more 
permanent sites Go, G7 and G8 grouped low on Axis 
2, those from {he less frequently inundated G1 and 
G2 prouped centrally on Axis 2 while those from the 
more extreme temporary sites G4, G5, G9 and G10 
grouped high On Axis 2, Samples showed no distiner 
groupings according to microhabitat on any of the 
three axes (Pig. 8). 

The ANOSIM procedure suggested diflerences 
belween groups of simples at cach level: significant 
differences were located between sumple groups ata 
mesohabitat (R=L 118, p<0,001), site (R=1.362, 
pe0.0O01) and microhabitat (R= 1.079, p<0,001) level, 


Inman Frequent Infrequent 
Frequency —— 
‘Siles Ge ial cx] fev) 

G7 G2 Ga a1 

GB G&S 
UPGMA EFGH A c D 
Groupe 
Dorrwnant race Mecrowrachiun Thilcwrne Oecelissp Evntyes sp 
(SIMPER) dust abonioe padorinensle 

Crsuicean Gasimpod = Preciuiory —— ikphdy muatiiien 


Trenopteran predatary 


Biv 5S. Sites, UIRGMA itoups. and dormant tix Mren 
SIMPER in relation ty inuadatiua Frequeney, LIPCIMA 
eroup B has been omitted as i contains ont ore sacnple 
ind domindnt (Xu chniet be vendnited for one sample 
wroups using SIMPLER, Also the single sample from site 
Git (in LIPOMA feoup P78 Ongitied ae ie an edbier. 
being uniifrequently flooded Site that grouped with the 
Frequently Mouslevt sles. 


With regard to the jnstantaneous environmental 
variables measured, the sample distribution along the 
first axis of the SSH ordination was significantly 
correlated wilh Seechi depth, temperature and 
oxygen saturation (Table 4). Axis 2 showed 
significant correlations with salinity, temperature 
and oxygen saturation and all variables were 
correlated with sample chstbution along Axis 3. 


Functional feeding groups 


Collectors dominated the invertebrule assemblage 
of Goyder Lagoon, However, there was considerable 
variation in the FFG composition of the mesohabitats 
and of sites within mesohabitals (Fig. 9), Billabongs 
and waterholes contained similar numbers of 
collectors and predators. Plowever, collectors 
dominated the assemblage composition of the 
backwaters in both richness and abundanee, When 
the FFG composition of individual sites was further 


a 
Billabong 


@ Backwalor 
@ Walarhole 
© Billabong 
O Channel 


Aats 3 


ao 


hig G, SSH ploronases fab dy, Z(t by a le) 2, Sal 
auimples callected front Goyder Ligwom November 
(U9 Samples ure hibelled aecontine to the fiesphabitats 
from whieh (hey wereecoteerce 


26 F, SHELDON & J.T. PUCKRIDGE 


DNHAODOOKAGDH 


Qa 
S=OONTOBON + 
5 


oO 
e 
+ 
4 
a 
a 
im) 
° 
e 


* 
g 


Cliftan 
Hills 


i) 
Axis 2 


Fiy. 7. SSH plot on axes (a) 1 y. 2, (b) 1 y. 3, (¢) 2-y. 3 of 
samples collected from Goyder Lagoon, November 
1993, Samples are labelled according to the site from 
which they were collected, 


@ Leaf Litter 
Sand 


© Sedge 
&é Silt 
© Snag 
+ Rock 
@ Lignum 
Emergent 
Vegetation 
é Grass 


\) 
Axis 2 


Fig. 8. SSH plot on axes (a) | y. 2, (b) 1 y. 3, (¢) 2 y. 3 of 
samples collected from Goyder Lagoon, November 
1993. Samples are labelled according to the microhabitat 
from which they were collected. Emergent vegetation = 
Polysonum sp, and Hooded Cyperus spp. 


TABLE 4+. Spearman Rank correlation coefficients between environmental Variables and the sample scores on the first 
three axes of the SSH of faunal data from all samples callected Jrom habitats in Gavder Lagoon, November 1993. 


Significance levels are indicated as follows: ns = not significant, * = p<0.05 pe0.01: = p<0,001, 
Axis | Axis 2 Axis 3 Salinity Secchi Temp 
Axis 2 0,032 
ns 
Axis 3 -.143 (),324 
ns ae 
Salinity O.071 0,698 O.416 
ns ae a 
Secchi -0.658 0.029 0.375 O40) 
ns Hat eae 
femp 0.394 0). 302 -0,367 -0,237 -0,355 
sees yes # # * 
Dissolved -0,365 O14 0.409 0,835 0.688 0.261 


Oxyyen 


MACROINVERTEBRATES OF GOYDER LAGOON 


120 
Mesohabitats 


80 


Abundance 


40 


(a) 


water Channel Waterhole 


9 tb : 
Billabong Back 


75 250 


Sites 


\ 


160 


Mesohabitats 


Richness 
a 


fi 


0 : : 
Billabong Backwater Channel Waterhole 
Sites 


2 


Vig. 9, Mean (+SE) of (a) abundance and (b) richness for collectors (grey bars), predators (white bars) and scrapers (black 
bars) for mesohabitats and sites within mesohabitats, collected from Goyder Lagoon, November 1993. 


Git G3 Gi 


2M Bf SHELDON & f 1. PUCKRIDGE 


TANLE a. Pudertonal Feeding Group rats as indicators af damindnt ecosystem Processes. 
{ Auto = autotrophy: Hetero = Heterotrophy; CPOM = Coarse Parniculate Organic Mauer; PPOM = Fine Particulate Organic 
Matter; TFPOM = FPOM in transport; BFPOM = FPOM in substrate] (see Merritt & Cummins 1996). 


Eeosystem Functional Feeding Calculated Generul Criteria rao Evaluation 
Parameter Group Ratios Ratio levels 
Auto/Hetero Scrapers to Shredders + 0.21 Autotrophic Heterotrophic system, 
Votal Collectors >(),75 dependent on 
allochthonous organic 
matter inputs 
CPOM/ Shredders (o Total a Normal Shredder No shredders, riparian 
FPOM Collectors assoenition linked to zane functioning 
functioning riparian differently from thitt 
syslem predicted by Merritt and 
>t).25 Cummins, 
TEPOM/ Filtering Collectors to 12 FPOM transport (in low PROM in transport. 
BFPOM Gathering Collectors suspension) enriched = high particulate lou 
20.50 
Stable Serapers + Filtering (36 Stable substrates Limited stable substrates, 
Channel Collectors to Shredders plentiful habitats dominated hy 
+ Gathering Collectors >0.50 sand wn silt 
‘Top-down Predators to Total of all (ag Normal predator to Large mumber of 


control other groups 


explored (here were considerable differences among 
sites even within the same mesohubitat, The 
backwaters GJ and GS showed similar patlerns with 
both richness and ubundance dominated hy 
collectors. The two billubongs, however, differed 
with both the pichness and abundanee of G3 being 
dominated by predators compared with collectors in 
Gil (Fig, 4). Variation in PPG) composition of 
assemblages also differed between the different 
witerliole sites. The more temporary weterholes G4, 
G5, GY and G10 were dominated by invertebrate 
predarors whereas the more permanent walerholes 
G2 und G7 had aw larger number of collectors, 
Overall, there were few serupers. 

OF the total of S54 tisa collected from Goyder 
Lapoon sik were designated as ‘Scrapers’, three as 
“Filtering Collectors’, 25 45 “Guthering Collectors’ 
and 20 were ‘Preditors) no ‘Shredders’ were 
colleered from Goyder Ligoon (see Appendix), PPC 
ratios (Merritt & Cunmmins 1996), calculated using 
the PRG data indicate that Goyder Lagoon ts 
heterotrophic, has dow levels of invertebrute 
mediated leat litter breakdown, and the majority of 
PPOM in the system ts m storage (Pable 5), 


Discussion 


Samples taken ty Navember 1993 from habitats in 
Goyder Lagoon contiined a total of S4 macro- 
invertebrate bisa, Nearly all of these taxa have also 
been reeorded from the Coongie Lakes system 


prey balance 
<().15 


invertebrate predators, 
reflecting temporary 
habitats late in succession 


(Sheldon unpub.) and from the lower River Murray 
und Darling River (Sheldon & Walker [998 ). 
However, a striking feature of the Goyder Lagoon 
assemblage was the presence of a diverse vroup of 
Motlusea, including two gastropod taxa. Neropeli 
wwhlincata (Conrad) und Thieres beloomensix, that 
have become extremely rare, if not extinct, in the 
River Murray and Darling River (Sheldon & Walker 
1995a.b), 

Collectors were the dominant fecding group aeross 
all habitats du Goyder Lagoon, Funetional feeding 
group rahios (Merri{t & Cummins. 1996) suzgest that 
Goyder Lagoon is heterotrophic and dependent on 
dlochtionous organie matter asa carbon source 
(Table 5), This is not surprising sinee aquane plants 
were fare within the Lagoon. siall stiids of the 
mucrophyte Polygonum sp. being the nist 
conspicuous, The absenee of shredders iv the 
ussemmblave sugeests (hal for the invertebrates. the 
major allochittynous inputs do not come drrectly 
from ciparier lirter fall but rather from particulate 
organic mater presedbits PPOM withthe substrate. 
Decomposing plant material of inostly terrestrial 
floodplain origin would therefore form the dormant 
Food source for He agate (iagralhvertebrate. lowed 
webs. Organic mutter of Moodplain origin his been 
found to influence the strueture of invertebrate 
assemibhiwes mother lirge floodplain river systenns 
(Post & De La Crue 1977, Caley 188: Perry & 
Perry 99), Thorp & Delonge 1994) Meyer er at 
IQV7y 


MACROINVERTEBRATES OF GOYDER LAGOUS 19 


All samples were collecied when water levels 
within Goyder Lagoon were low. Low water levels 
lend jo maximise (he “heliveen mesohablea’ 
differences (ice. differences between channel, 
waterhole and billabong). At bigher flows. between 
habitat differences would become inereasingly 
blurred as billubangs and witterholes beedme parr oF 
the channel environment. Alihough we expected the 
main differences in assemblage vomposition to occur 
henveen different meso- and mitrohabitats 
(regardless of site). there were striking differences in 
the Tichness aod wbundance of macromyertebrates 
collected from the different sites, even belween siles 
from the same mesohabitat type (Pig. 3), These 
differenves may reflect different suocessional 
Inyeclones oeeurring withi the assemblage at cach 
site after hydrological isolation (Boulton & Lake 
}992). Thus. the recent Looding/dryiig history of 
each site may be a Significant factor in determining 
the structure of the assemblive at any time, 

Sites, regardless of mesohabitut type. at Andrewi lla 
and Koonchera Dune contained the largest number of 
individuals and the greatest number of nixa (Pig. 3). 
These sites ure Intermediate m the range of Mooding 
Frequency (Fig. 5), As Tlooding is uw form of 
ecological “disturbance” this supports the notion of 
“infermediate disturbances’ being a driving Loree tn 
mainwining animal diversity within ecosystems 
(Ward & Stanford 1963). The sites ob Andrewilli 
Watethiole and Koonchert Dune. falling wathin thus 
band of intermediate flooding trequeney. were 
characterised by taxa such as the prosebranch 
wastropad Chiara fatlonnensis and the predatory 
cuddistly Gecens sp, (Pig. 5). Both these taxa tended 


to be fare in the more permanent sections of the 
Lugoon. 

Goyder Lagoon isan ared of high conservation 
significance (Morton el ad T99Sa), It provides 
habiiat for a ovariely of aquatic and terrestrial 
Organisms idan otherwise arid chvironment (Morton 
etal 199Sb: ANCA 1996). The muacroinvertebrate 
assetiblage, although not unique to Goyder Lagoon, 
did contain a number of molluses that are becoming 
inereasmyly encingered in other river systems. [bis 
the Geomorphology of the Lagoon that grves rise to 
the different mesohabitat types. Overlying Uns 
morphological template is the hydrology of the 
system. The variable (ows characteristic of Goyder 
Lavoon are intrinsie in Maintaining macro- 
invertebrate diversity, The waters of wethinds in aril 
regions are Inereasingly in demand by water resouiree 
developers (Walker e7 ul 1997). 1 Goyder Lazoon ts 
to remain an area of high conservation value then it 
is imperative that the hydrological diversity 
ehiracteristic of the system i maintained. 


Acknowledgments 


This study was condueted as part of the Biolugreat! 
Survey of the Lower Diamantina Floodplalii. a joint 
project of the Conservation Council of South 
Australia and the South Australian Department of 
Environment amd Plunning. The project was funded 
by the Australian Heritage Commission. We thank R. 
Molsher, J. Arnold, A, Bingemer and Bunge for 
collection and sampling sorting assistance and Mr P, 
Schulty and ain anonymous referee for extremely 
helpful comments. 


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Appendix 


Species, functional feeding group [PC = filtering collectay, GiC = wathering collector: & = Scraper P= 


Predator] und total 


abundance for simples collected trom habits i Goyder Lagoon, Diamantina River, November 1994 


Spee FFG Total 
MOLLUSCOA 
BIVALMIA 
Sphacriielac Sphueriun sp. KC 5 
Corbieuhdie Cerio austniliy (Deshavesy FC an 
Lbyrifdae VWlesiuiin wilson? thew iC | 
EPASTRORPODA 
Aneylidiue frepisvic spp S | 
Manorbidke Glypraphysa sp. 5 4 
Vivipuiridae Veropale sublireant (Oonrad) ic II 
Phiri Chinn halaineisis Canis Cit! 1a 


OLIGOCHAETA 


HIRUDINEA 
Glossiphoniidae 


CRUSTACEA 
CONCHOSTRACA 
Cyzicidae 
DECAPODA 
Palaemonidae 
Parastacidae 


INSECTA 
EPHEMEROPTERA 
Caenidae 

Baetidae 

ODONATA 
Gomphidae 
Cordulidae 
HEMIPTERA 
Corixidae 


Ochteridae 
Notonectidae 


COLEOPTERA 
Dytiscidae 


Hydrophilidae 


Hydraenidae 


DIPTERA 
Tipulidae 
Chironomidae: Tanypodinac 


Chironomidae: Chironominae 


Chironomidae: Orthocladinae 
Ceratopogonidace 

Muscidae 

TRICHOPTERA 

Eenomidae 

Leptoceridae 


MACROINVERTEBRATES OF GOYDER LAGOON 


Indeterminate spp. 


Indeterminate sp. 


Cyzicus sp. 


Macrobrachium australiense (Ortmann) 
Cherax destructor (Clark) 


Tasmanocoenis arcuata Alba-Tercedor & Suter 
Cloeon sp. 


Austrogomphus australis Selys 
Hemicordulia tau Selys 


Micronecta spp 
Cymatia sp. 
Indeterminate sp. 
Anisops spp. 
Enithares sp. 


Antiporus femoralis (Boheman) 
Allodessus sp. 

Eretes australis Erichson 
Cybister sp. 

Rhantus sp. 

Hyderodes sp. 

Sfernopriscus sp. 

Enochrus sp. 

Berosus sp. 

Limnoxenus sp. 


Octhebius sp. 
Hydraena sp. 


Indeterminate sp. 
Ablabesmyia sp. 
Coelopynid sp. 
Procladius sp. 
Cladotanytarsus sp- 
Tanylarsus spp. 
Chironomus sp. 
Chironomus cleacalis Atchley & Martin 
Cryptochironamus sp. 
Stenochironomus sp. 
Parachironomus sp. 
Dicrotendipes sp 
Paratendipes sy. 
Cricotopus spp. 
Bescia sp. 
Indeterminate sp. 


Ecnomus sp. 

Triplectides australis Navas 
Triplectides clongatus Banks 
Ovcelis sp 


GC 


GC 


Qwun vised 


nn 


BREEDING BIOLOGY OF LITORIA BOOROOLONGENSIS 
(MOORE, 1961), AND LITORIA LESUEURI 
(DUMERIL & BIBRON, 1841) (ANURA: HYLIDAE) AND 
COMMENTS ON POPULATION DECLINES OF 
L. BOOROOLONGENSIS 


By MARION ANSTIS*, ROSS A. ALFORD}? & GRAEME R. GILLESPIE£ 


Summary 


Anstis, M., Alford, R. A. & Gillespie, G. R. (1998) Breeding biology of Litoria 
booroolongensis (Moore, 1961) and Litoria lesueuri (Duméril & Bibron, 1841) 
(Anura: Hylidae) and comments on population declines of L. booroolongensis. Trans. 
R. Soc. S. Aust. 122(1), 33-43, 29 May, 1998. 

The embryonic and larval development of Litoria booroolongensis are described and 
compared to those of the closely related Litoria lesueuri. The habitat, behaviour and 
distribution of each species are compared and indications of marked population 
declines of L. booroolongensis are discussed. 

Key Words: Litoria booroolongensis, Litoria lesueuri, embryology, larval 
development, habitat, aggregation, population decline. 


Transactions of the Raval Societe of SMa (9OR), P2200) A4-d 44 


BREEDING BIOLOGY OF LITORIA BOOROOLONGENSIS (MOORE, 1961), 
AND LITORIA LESUEURI (DUMERIL & BIBRON, 1841) (ANURA: HYLIDAE) AND 
COMMENTS ON POPULATION DECLINES OF L. BOOROOLONGENSIS 


hy Marion ANSTIS*. Ross A. ALFORDT & GRAEME R. GILLOSPIE: 


Summary 


Assis MM, Acrarn, R.A. & Girrrsme GR, 199%) Breeding biology of Lirarta aorodtongensiy (Moore. 
L06)) and Litera lesuen? (Dumeéril & Bibron. IS4)) (Anuew: Plytidae) and comments on populabon declines 
of LE, hoarowloreenves Trax 4 Sow S. Anst (2240), 33a, 29 May. 199k 

The erabryonic and fsrvail dewelopent af Litera: Becrnlongensns are deseribed and compared (0 those of the 
dlosely related Line feet’, The hubital, bebaviode dad: distribiition of each spews are compared and 
indicuGurms OF marked populition declines OF Ly owropfongensés ave discussed, 

Nhe two species have similar Jntic 1de-distories bin some ctifferences vecur ty body proportions, colyur in life, 
Dehavion! and habitat. iy both species the ewe nus isu compaer gelatinous clump, lypica) ot frogs breeding an 
vote environment. The tadpoles of both species are adapted to the lorie environment and hive streamlined 
bodies and suctoial moutb-parls. Adult Lo borroebuigenste vgerepate throughout the year and are wetve 


curnally (7 summer, 


Key Worns: Literla bourmelonmensiy, bite leidied, eoobryolopy, larval development habliit, dtgregatiott. 


popHlationd decline 
Introduction 


Literia, baorimlongensts was first desenbed by 
Copland (1957), us Ayia XY Moore. pending the type 
descdiption (Moore 1961). Moore deserbed tas an 
“uplind species” extending from the Armidale 
region to the Blue Mountains. in NSW. bur made no 
relerence to larval developotent, Anstis (/97+4) 
briehly described the tidpole as lotic and suctorial. 
with a footh cow formula of -/s und numerous oral 
papillae. The present paper provides a deseription ul 
embryonic und larval development or L- 
houwolongensis, With Comparisons to the sriibu’ 
tadpole of Lo leyueuri. Litarta hoeranlanensts is 
associated with flowing stretinis on the slopes and 
tablelands of the Great Dividing Range from the 
Queensland border to (he Victorian borer (Barker ef 
al, 1995). with the type locality (Guy Bawkes Creck 
near Bhor, NSW) and most records of this species 
indicaling that it commonly oecurs above S00 m in 
the region of the New England ‘Tablelands, NSW 
(Heatwole er ef. 1995), The most southern record of 
this species is from the Tumut River, Kosciusko 
National Pork and ithas revently bee found pr the 
adjacent Gooburragandea River, east of Tumult NSW 
(Hunter & Gillespie unpub.) Observations on the 
current shite of popularions (particularly in the New 
Lnghind region) ure discussed. 

Literia lesuened (Dumeril & Bibron) occurs in 
sunterh Australia tron) northern Queensland to 
Vittoria und almost vertiinly involves a comple of 


26 Wideyiew Ral Bepieca | Lehr NSW 2082 

» Department of Zoulogy aid Teapiwal Feelawy. Ties Cook 
Lnrversity Tawesville Ql dst 
Pepariniont of Addepy, Uivgedey of Metbornie Parkwitle Vie 
WU52, Anhur Rylud tostiowe HO os 147 Pecinelbere Wig. tO S44 


sibling species (Moore 1967: Barker ef af, 1995) 
Vhe type Joeality is) Port Jackson, NSW. 
Observations on current population trends at same 
southern sites are reported. ‘The eggs of the 
northeastern. Queensland form have been deseribed 
by Rivhurds & Alford (1992) and Barker ef af, (19954 
brielly describe the site of epe deposition by the 
populations found Fram southern Queensland to 
Victoria. The tadpole as found in the Melbourne ares 
has been brielly deseribed by Martin eral. (1966). As 
the distributions ef both species overlap Th places, 4 
more detailed deseription of Lo lesmerit tadpoles 
from NSW is provided here for the purposes of 
comparison with L, boaraolengensis and lo assist in 
distinguishing tbe species. Distribution maps for 
both species together with localities studied m the 
present paper, are presented as Fiyure (- Numbered 
localities on Fig. DA relate to 1. heatadivierests und 
those on Piss [Bota L. lesen 


Materials and Methods 


Material esanined 

Literty baeratoigensis larvae Atstahan 
Museum (AM) R119062-64. 119087. 119071, 
(19073 (Serpentine River, Point Lookout NSW), 
RIIY0S5, | LOUBO. 1 P9083. 1 L985, 119N87. LTRS 
(arvae drom Back Creek. Porm Lookout NSW). 
Embryo/larval descriptions ure based en one cee 
mass fronian amplectant pair collected at Serpentine 
River (near Point Lookout NSW) on. 3.91. 1973, The 
pair was placed im an inflated plastic bag containing 
stream water, teeds and a rock, Until after 
oviposition, Larvae were maintained for up to three 
months i containers of 40 crm diameter ab water 
temperatures of 4 -25° C. Only larvae tram strearns 


34 M.ANSTIS, RA, ALFORD & G, R. GILLESPIE 


i a a ee 


wir 


“i 


Vig 1A. Mup of eastern half of NSW showing distribution of Literia boorolvmgensts (shaded area = AM records: * = 
localities referred tin present paper). |, Back Creek, Point Lookout 30° 29' 20" 8. 152° 20° 48" E. 2. Serpentine Creek, 
Point Lookout 30° 28! 26" 8, 152° 19 26" B39, Near Tamworth 317 139° dO" 8. 150° (0 30" G4, tsaies Creek. Timor 
Caves 31°41 40'S, 151-07! 20" E44 Goobarragandra River 35° 24°02" §. 148" 26° 01" B. Populations no longer exist 
al loculities | a 2 (see tex), Bo Map of eastern hall of NSW showing distribution of Liria desuenet (shaded area = AM 
revords: #= localities referred to in present paper). 1, Quriimbalt Creek. 33° 19°45" 8, 151° 21' 38" B. 2, Near Palmadule 
AS (WATS, (St 20) 42" 3, Darkes Forest 34° 14" 02" 8, 190° 54 00" EL 4. Durkey Creek near Singleton 32" 44) 
36" 5. 150" 50° 32" B.S. Coco Creck BE. of Cupertee 33° 08° 15" S$, 150° Ue 40" E. 6. Mongarlowe River, Braidwood 
48° 29) 42"'S, 149° 57° 11" FO 7, Dingo Creek, Winghaty 31° 49° S50" 8. 152" E800" E, 8. Berrico 32° 04) 00" S, 151" 
19 79" 6.9. Tir Creek. Bulga State Forest 31° 3149" S$. 152708) 21" B. 10. The Busin, Watazan State Forest 33° (6! 
20" S, IST TP 4" B, LL. Bowong Creek 36° 21) 30" 5, 148° 2215" b, 12 Cotter River 39° 35'27"S, 148° 49! 0" BE. 
13, Goodradighy River 35° 25' 0S" S, 48° 44" 40" E. The MacDonald River is shown on both maps. 


where L. leyvewr does hot ocear (in the region of the 
type Incality) were inchided in the study. Lérerie 
lesueurt laryacc- NSW localities: AM RLIOTOL - 
Glen Davis, 119099 - Ourimbah: 119104 - Allens 
Creek Picton: 119106, |19107 - Dingo Creek near 


Winghun; (IOS = Berrico Creek, 2 km LE, of 


Bertico; AYAS - Witiuean State Forest; A724 - Tirnill 
Creek (collection of M.A.) 

Tadpoles were reared Lo metamorphosis to confirm 
ideouty, Embryos and hirvae were measured with 
vermier callipers und an ocular micrometer attached 
fod Wild M5 stereoscopic Microscope, Larvae were 
observed if the Natural environment, then examined 
while anuesthetized with chlorbutanol, The staging 
system of Gosner (1960) was used, For L, 
hooroolmivensis, embryos studied included stapes 2 
(n= 11), 8 (n= 15), 17- be (n=9), 19-21 (n=9), 22 
& 25 (= 8). hirval stages included 27 (n= 1), 28 (4 
= 4). 31 (n= 2), 32 (n = 1), 33 (m= 3). 34 (n = 4), 35 
(n = 3), 36( n =4), 37 (n=2), 38 (n= 1), 40 (= 1), 
41 () = 6) and inclamorphs al stipe 40 (n= 10). For 
L. fesueurt, larval stiges included 25 (n= 3), 27 (n= 
1), 41 (n= 2), 32 (n = 1 34 (n= 1). 35 (n = 3), 30 


(= 2). 37 (n =4). 38 in = 1), 39 (9 = 2), 4h (n= 2) 
and metamorpliy at stage 46 (n= 7). Ulustrations 
were made with the aid of drawing tube attached tu 
the microscope, The following abbreviations lor all 
morphometric variables are taken fram Anstis 
(1994), 
Lureral view: TL = total length, BL = body lenge 
BD = body depth: BTM = depth of tail musculature 
atbase of tails TD = maximum tail depth; DP =depth 
of dorsal fin (in tine with TD): VF = depth of ventritl 
ii Gu bine with TD); TM = depth of tail musculature 
(in line with TD); SS = snout to spiracte; S-E = np of 
snout lo anterior rim of eye: S-N = tp of snout to 
anterior ri of nutis; ED = diameter of eye. 
Dorsal view: BW = body width, EBW = body width 
at level of eyes; BTMW = width of tail musculature 
dt base of tail; 1O = interorbital spun, EN = edge ol 
eye to edge of haris; IN = internurial span. 
Ventral view: MW = maximum width of oral disc, 
The ubove morphometric variables. were logy, 
ronstormed prior to statistical aualysis, A principal 
components analysis (PCA) on the coviiriince matrix 
of these values was used to reduce the 


BREEDING BIOLOGY L. BOORQOLONGENSIS & (. LESUEURI 35 


dimensionality of the data set and remove the effects 
of overall size (which is extracted ds the lirst 
component (Marcus 1990). The scores of each 
unimal on the second and third component 
(representing Shape variables) were plotied in a 
hiplot (Digby & Kempton 1987) in which the sealed 
couffivients of each variable on the second and third 
PC axes. were overlaid on the data. Examining this 
plot enabled us to determine whether the species 
differed in shape, which size-independent shape 
variable was most important in this discrimination 
and io postulate which of the original variables 
contributed most to the score on. that yartuble, We 
then calculated the slopes and intercepts of the 
relationshipr between log,,-transformed beady length 
and wl other Jog) )-transformed morphometric 
variables for each species. We used separate analyses 
of covariance (ANCOVAs) for each morphometne 
varible 10 determine whether the slopes and 
inlercepts of the regressions differed significantly 
belween species, 


Results 


Distribunon/hahirat 
Lilorin boorovlongensis 

A map of the eastern half of NSW. shows the 
general distribution of this species. as determined 
from specimens cegistered in the Australian Museu 
and the localities referred to in the present study (1- 
5. Vis, PA), Hubrtats studied Were permanent Mowing 
seams running through wet or dry selerophyl| 
forest. or through semi-cleared grazing land in basalt 
Or granite range country (altitude range 450-1340 
in), Field observations On this Species were cated 
out annually in the summer breeding time of 
December/anuary (1965-1974) at Back Creek, Point 
Tookout NSW (loculuy 1 Fig. PAL 30° 29° 29" 8, 
152" 20) 38" EB), Serpentine Creek Clacality 2 Pie 
LA, 40" 28) 2608, (52° 19° 26" Ey and other neurby 
sirens al Potnl Lookout NSW (1250-134() om). 
{uring this period of nine years, gumerous tdpoles 
mehuniorphs and adults were readily lounct in the 
severil Streams inthe Pott Lookout and Arpaidialy 
rewion. A survey ol these same streams in December 
190d yeveuled No adults or larvae. 

The only southern loeality. surveyed was 
Goubarragandeat River (Ingality 5 Bis, TA, 38° 2 
O2" 5, 148° 2a’ O01" Ey where 13 adults were {once 
on Oxi 1996 along a BOO Mm stretch of the river. 


Litavia lesireiurt 

A nmap of the waster hall of NSW shows the 
eeneral distribution of this species as determined 
from speelmens registered in the Australian 
Museum, and a sample of localities studied trom 
1972-1996 (Pius. TB, 1-13), Habitits. varied: from 


flowing streams to large dams in sanestone, 
melunorphic or granite country in rain forest, dry 
yeleeophyll forest or heath land (40-| 1000 m), ‘The 
frog was encountered 1h fairly low numbers in recent 
surveys in most streams in NE Victoria apd was 
absent from several sireams Where it would be 
expected 1a occur, on the hasis of habitat and focal 
distribution. Surveys in the Kosciusko National Park 
in 1996 (Hunter & Gillespic unpub.) found the 
species to be present inoply 15 of 40 likely streams. 


Behavionr 
Literig hooroneangensis 

During summer months in the 1960s-70s, adults 
were observed bisking if the su on exposed basalt 
rocks in mid-stream and frequently (ree or more 
individoals were found onder the same rock 
Within/beside the stream, ut the sorther Back Creek 
and wt Serpentine Creek. When disturbed. they 
immediately leapt under the flowing water Males 
were observed calling at night while suung on 
exposed rocks in shallow, flowing seclions of the 
stream. Six females (four gravid) and seven males 
(six with nuptial pads) were found by day under 
stones on pebble banks at the southern 
Goobarragundra River on 3) November 1996, 

Numerous Jarvac and metamorphs from stages 25~ 
46 were observed at the two northern loculilies 
throughout December/January (1965-74). Tadpoles 
were commonly [ound on the substrite anonyse 
rocks. in shallow flowing sechons of stream 
including runs and riffles, and in shallow. slowly 
flowing inlets at the sides of the seam. The tadpoles 
pussessed at yiuimber of features typical of species 
inbubiting the lotic enyironment, including a 
suctorial oral dise fully surrounded by fhree or more 
rows Of papillite, a more streamlined hody form 
(especially the snout), a dicker tail musculature ung 
relatively shallow fajs, They were observed adhering 
to rocks with their suglorml mouths, tails bending tp 
the direction of water flow, 


Literia desueurt 

During winter, adults were found on ridve-lops in 
forest country away from streams. Adulis were 
ustully assecrited with [lowing streams. but alse 
bred ia completely isolated streamside pools ISPs) 
stots in bedrock shelwes. where the water was sul 
Males were observed calling beside dums in open 
forest near Ourimbah NSW (locality | Pig. 1B, 34" 
19°45" 8, 191° 23' 38" B). Eggs were found laid in 
the stream on the edge oF rons in Slow water in 
conneeicd still pools, arin ISPs. At Bogong Creek 
(locality 11 Pry. 1B. 36° 21 30" S) 148° 1215" haa 
series of perched ISPs was consistently selected as 
breeding sites by several Fermales froin 1994 ~ 1996. 
In the first vear all but wne ladpole from two 


mir] M. ANSTIS. ROA ALPORD & G.R. CALLLSPIE 


cluiches survived, in the second year the pools dria 
up aud in the third year, newly-hatehed tadpoles 
could not be found after a tigh spring fload, No 
tadpoles have been found in the stream iiself in any 
OF (hese years, 

Ip Streams or fivers. Gidpoles were most commonly 
fuund on the substrate in shallow, slowly Mlowime 
sections of streams, segregated back waters wit 
reduced flow and perched rock pools or isolated 
pools Git times stagnant) beside the stream. They 
were yery agile when disturbed, capable at’ fast 
movement darting under rocks or leat fitter. As ind. 
hoomelongensiy, hey possessed a suctorial mouth, 
shallow fins, thick (ail musculature and steeambined 
hocly 


Oviposition and embryonic development 
Litaria hoaraoloneensis 

‘The mean ege complement of four gravid females 
examined at Goobarragandra River near Tumut NSW 
was }S19 (range 1292-1784). A pair of frogs. foond 
in amplexas at Serpentine Creek on 3.x), 1973 was 
first observed at O740 he sitting in sunlight on an 
expased rock in a shallow inlet pool near the edge of 
the stream. After (O min they bad moved to a rock 
closer lo mid-streany ina shallow, Plowing seetion. 
The frogs were then collected ane phiced in i plastic 
bag, and at (300 ha single. compact exe muss was 
found partly adhering Co suspended veyenbon within 
the bag To avoid disturbance during development. 
eggs Were not counted. 

Embryos were al stage 2 when a Sample was first 
preserved at 1300 h. three or four hours after ees 
were lanl. There were two layers of jelly surrounding 
The embryo. In live emnbryas the animal pole was 
dark yrey aind the vevetal pole grey, The same 
preserved embryos examined in 1996, had a brown 
animal pole mening lo cream over the vegelal pole, 
A sertes of 1] preserved embryos at stage 2 ull had a 
diimeter of 146 mm. Approximately nine hours 
fier depostion they were at stige & and 15 embryos 
had a meun diameter of 15 mm_ The blastomeres 
were nore evenly divided tn the antinal pole than in 
the vegetal pole. Afier 36 h embryos were a stages 
|} and 12 and alter 56 h most were at stage | 
Stages 17-18 (lail-bud),. were reavhed after 67 by, 
Nine specimens at these stages ranged from 7.43- 
3.36 Tan (mewn 2.96). A typival stage 17 embryo 
(Hig. 2A), had a rounded snout, prom ipent V-shaped 
ddhesive organs stomedaeal pity small gill-plate 
bulge; indistinct! pronephrie swelling: slight optic 
bulge: indistinct nurial pit slightly delineated with 
pigment, und rudimentary fins along the tail bud, 
Live embryos at stage 19 were light grey above with 
i pale grey yolk sac. 

Some embryos burst through the capsules duping 
le Mosculiir response in stage (8S but most did not 


begin hatching vad! stage 20, An embryo al stage 20 
Measuring 5,76 mm (Pig. 2B) is deseribed us 
follows:- snout rounded; two pairs af externul gills - 
two branches on upper pair and 4-5 branches on 
lower; Optic and narial regions partly outlined with 
small crescent of pigment: adhesive organs well 
divided — remnant of V-shape below each: numerous 
fine museular tidges alone tail musculature: 
stontodaeal pit and fins both deepening, Dorsuni 
grey, yolk sac pale grey in live embryos, und pale 
brown/ycllow brown in preserved specimens, 

Live embryos at stages 20 und 2) were grey above 
with a lighter grey yolk suc and translucent grey fins. 
Nine specimens at these stitges ranged from 4.82 - 
5.07 yim (ean 5.43), Mitchlings adhered strongly 
ty the jelly mass. The external gdls were visible 
macroscopically and when fully developed by stage 
22 extended about 2/3 the length of the yolk sac, 

Ereht embryos ut early stage 25 ranged from 9.23 
957 im Cea 846 ) dorsal pigmentation of larval 
Stages developing; dorsal edge of tail museularure 
finely edged with melanophores: fins. external body 
wall (ind venter clear (in preservative); spiracle and 
vent lube both functional: jaw sheaths pigmented, 
hut as yet without shape of older larvae; 
keraunisation incomplete, Tooth rows almosi 
complete except for 3nd lower row. which was 
beginning fo develop m two more advanced Iprvac at 
suige 25, measuring 11.5 and 12.15 nin. respectively. 


Litorta lesnenrt 

Most clumps of eggs observed were loosely 
udbering 10 rock substitte but some alse adtiered bo 
bottom sediments or Sedwes. There was no indication 


——— B 


Fin. 2 Embreos of Liforne boerrploieensis A, Retnacee 
from capsule. stave (7, 3,24 nam, GB, Just hatched -stigee 
20, 5.76 mm, Soule bur = finn. Stuwes ace (rom Gosner 
( (960) 


BREEDING BIOLOGY L. BOOROOLONGENSIS & L. LESUEURI MY 


of nest-site excavation, as in the north-Queensland 
form, and 20% of clutches observed were deposued 
under, or partly under a smal! rock on the bedrock of 
the stream. Eygs were laid in a water depth of {0-20 
cm, Clutch sizes of 14 egg masses examined in 1995- 
1996 ranged from 610-3564 (mean 1878). The mean 
diameter of 10 eggs at stiges 43 (Gosner 1960) from 
Guoodradighy River (locality 13 Pig, 1B, 35° 25°05" 
5S, 148° 44" 40" E, was 17 mm. No embryos beyond 
this stage were ayailuble for study. 


hurvee 

Analysis of covariance showed that the two species 
follow very similar lrajectories through larval body 
sives und stiges (tests lor differences i slope and 
intercept both p >> 0.05). The relationships between 
developmental stage and body size for both species 
are Wustraled in Fig. 3, which makes it clear chi 
jhey cannot be distinguished on this basis. 

The results of the principal components analyses 
(PCA) ure presented in Big. 4Al with the 
contributions of cach variable to the second and third 
vivenvectors Shown Ww Table f, The first (general 
size) principal component accounted for 87% of the 
variauon ih the data set. 'Uhe seeond and third (shape) 
components accounted for 4.4 and 4.7%. 
respectively. Despite the relatively small proportions 
of the overall variance accounted for by these 
components, ib ts clear from Vig, 4A, that in 
combination they diseriminate very effectively 
between the two species, indicating that the species 


41 ss ! be 


39 


37 


35 


33 


Mu 


Gosner Stage 


29 


27 # A L. booroolongensis 


35 > Bai. lesueuri 
5 10 15 20 
Body Length (mm) 


Fig. 3. Gosner (1960) stives plotted agwinst body length 
for Litovia hooradloigensis and Lirarki lesneurt. Line 
represents pooled regression ol stage on length. y = 1.32 
6+ 15.77. = 0.84, p< 0,000, 


do differ strongly in’ shape. Most of the 
diseriminatary power resides in Component 2, This 
means that the niorphometric variables responsible 
for the eigenvector loadings which are most closely 
aligned with axis 2 of the PCA in the biplot, should 
be most important in discriminating bebween the 
shapes of the two species. 


Tank 1 Reyresston of logy of morphomeric variables on leg), eo] hedy length: foreach species, with pevalies for 
vignificance af diflerences in slope and tntercept berween species fron an ANCOVA, and eigenvectors far the second and 
third principal components of lov jy ansfarned morphometric veriibles, 


Lo bowen gets 


Lh, leaueuri 


ANCOVA pevalues Principal Component 


Eigenveelors 


Dependent Variable Intercept Slope Intereept Slope Hileree pt Slope pC? PCS 
TL (2588 11077 (280 L077 H.0012 HOO UWnss 19 HPISRG 
BL - - - (03202 “04520 
BW OTR ads 1623 DY19S (3654 WARIO Ud1304 NAME 
BD 0,123 (8400 (2574 (), 0625 0,000) (0327 27523 (LOSSSA 

BW “16 {hava4 “1512 (ASRGS UW (8t45 AY OS 2 ahs? 
WW) 04061 (8095 ASAT bleed? L000) PhO (HES (1245000 
IN -U) a8UM (5303 9237 L004 OAS 0001 AROLO72 0.24724 
LN A) hls4 (7780 9479 | Wo77 (1942 (0002 0.07379 UOTSHAT 

WM Ww 15500 1.795] 10958 1381 (5247 (00M ALUGITS 1), 4u54u 

BRIM 9476 1.2425 O.6718 |QQ07 O85) UA275 (i rset (423404 
Th ANIISI 77a0 04308 (MMe OSE (0001 () 22844 Qtaj2o 
Db 04379 WATT -O,4984 ).2369 0.0014 1.0002 U260154 1-454 
T™ (7799 04276 89%) hos7a O.U569 3348 (119224 OO 16840) 
ve (2827 ()4277 U9264 1udos (),7292 (0003 (20800) (34620 
SS OLA O 449 “U1 894 9647 0.0016 1), 7200 -W0S245 (UALS 
Eb -],22355 [2728 (727) OSTA OAWIOT HOU W2AIle AY AT394 
Vw 5 Lot “O19 8719 0.000) 0,006) “47415 0, OG78 
SE 0.4276 TLOTSo -OA735 O.YTOS ONO NYIas “UO I6108 0.02676 
SN O.N832 (RATS 7280 ORO72 O.0D01 1.5980, O,554d0 O11531 


aK M. ANSTIS. R.A. ALBORD & ©. R. GILLESPIE 


0.3 
ray Litoria hoaraclongensis yr 
BC aLitoria /esueuri 
0.2 
2 oA 
° 
a 
w 
o° 
oO 
a 0.0 
-0.1 
-0.2 
0.3 -0,2 -0.1 0.0 04 0.2 0.3 
PC2 Score 
N 


Pip + A, Results of Principal Component: Analyses. 
Vectors aire proportional to foadings of cach original 
morphometric variable in caleakiting scores on edieh PC 
axis. Bo Plots of the six (host Tiportant morplenmetric 
variables suggested by the PCA agaist body length 


Figure 4B presents plots of the six most inportant 
vuriubles suggested by the PCA against body length. 
These plots show that the species differ in these 
charactenstics, in the direction indicated by the 
PCA; L. heoravlongensis has relatively greater 
snout-naris length. mouth width, and snout-cye 
distance and relatively lower body depth, dorsal fin 
depth und eye diameter than £, fesneqad does at each 
body length, 


Literia bearvalongensis 

A composite description of wdpoles at stages 25- 
39 is given. Using some anaesthetized specimens still 
showing colour in dite. A tidpole at stage 37 (36.2 
mm) fron’ the egy mass laid at Serpentine Creek 
(locality 2 Pig. }A) is shown in Fig. 5 AVC. 

Dorsal; body ovoid, widest ucross branchial 
refion: snout broadly rounded: eyes corse-laterdl. 
positioned alinost '/3 along body length from cp ot 
snout, nares opening antero-literally: body wall 
uniforn rusty-brown with some darker mottling: 
darker browi band aeross urastylar region, brawt 
body colour continues along tal musculature as two 
longitudinal stripes, becoming lighter towards tail 
tip; young tadpoles ul stages 25-26 dark brown wilh 
irregular light grey band just anterior to darker 
urostylar tegion: Fibs iereasingly pigmented alter 
stave 34. 

Lateral: body streamlined, snout rounded, elongate; 
spiricle sinistral, moderately long and broad, 


aa 


ta 


an 
& 3 & 


Snout-Naris Length (mm) 


a0 
E E20 
Eu E 
= S 
2 = 26 
© 
= ie 
53 
2 Eel 
a 
op Sy 
4 4 a ow of} 


E ' 
3 = 
3 

5 = 
6 E 
s a 
¢ A 
FA a 
5 

4 o 3 te =] . . = “ “ 
Gedy Lange (rmeny Body Lengin (awn) 
B A Liroria nooreorongensie bitoni tomumurt 


tapering slightly from origin to postero-dorsal 
opening; oral disc directed ventrally: vent tube 
dextrak: body brown with part of golden ventral 
sheen visible, particularly over branchial region; itis 
golden: (ail musculature thick anteriorly, uniform 
brown with some darker patches, main antertor blood 
vessel and crevices between muscular ridges 
outlined with pigment: fins relatively shallow, dorsal 
lin rises eradually (or more acutely) to greatest depth 
at mid-peint of tail, or just posterior/anterior to i. 
yenural fin inereases slightly in depth in posterior 
half, but generally shallower than dorsal fin; fine 
hetwork of melanophore clusters (raced over 
Vaseuhie system on dorsal fi) and posterior’ hall of 
ventral tin (denser beyond stage 30); tal Up rounded, 
Venlral; venter with almost uniform copper/gokl 
sheen of iridophores, or pateliy sheen with darker 
areas showing through in between patches. Branchial 
region densely covered with — copper/golil 
Indopliores, 

Oral dise (Pig. 6B) oral dise wide, directed 
ventrally, band of papillae surrounding entice 
convoluted margin; 2 - 3 rows wound anterior, 4 - 6 
uround dateral and 3 - 4 around postertor margins 
inner papitie on posterior murgin larger and) more 
widely spaced: papillae diminish iy size and inercase 
in OuMber through to outeemost rows two complete 
unterior und three complete posterior rows of labial 
teeth, all equal in length; juw sheaths moderitely 
massive, quite heavily keratinised, upper sheath with 


BREEDING BIOLOGY L. BOOROOLONGENSIS & L. LESUEURI 39 


im, 
ot tier 


~ , | 


al Dp 


Pies S. Laryue of Liferie booreoloneensiy und Literia leswenri, each al stage 47. AL Lo boc madongetyis. lateral view. Bob. 
leaneru7, lateral view, C. Le beorevlamernsts, dorsal view, D, L. fesnearr dorsal view, Seule burs = | nim. Stiging systeni 
oF Gosner (1960), 


Av M ANSTIS, RAL ALFORD & GR. GILLESPHB 


a 


tt 
B 


byt Oral discs of Litera byerectoneensis and hirortay leaueni A. Le lestetet Coeulilyy 10, Tig 1B, Stige 37), BLL, 
hooroolengertily Coeality 2. Pig, LAL Shige 34) Seale bars =) min. 


cental potch and small Waderlying keratinised ledge: 
inner mining OF jaw sheaths serrated, (edge partly 
warn in some individuals and dows not appear before 
Stage 28 in spechineus examined. 

Colour tm preserved specimens (composite 
deseription, stives 33-39): pigment slightly darker 
over abdomen, cramil, vetlebral and post-narial 
regions, Lut museulature evenly pigmented (dorsal 
view); abdomen opuqne wirh fine 
melihophores over snout and branchial region: 
dermnil body wall clear-with uw few seattered clusters 
of melonophores; iris black (lileral view); fine dense 
layer ol melunoplores over heart. abdomen and just 
anterior tO caeh gill; intestine visible; rail 
muscularice ahd liipbs unpigmented (ventral view). 


Liraria lesueuri 

Composite deseriptiom, live specimens. saytes 41 - 
34 A radpote ar sie 37 (33-9 mm) from Watayain 
State Forest (loculuy LO Fig. 1B) is shown ia Pig, 3 
B,D. 
Dersal; body ova, Widest aross abdominal region: 
snout rounded; eyes dorso-luteral ynd pusitaned less 
than '/4 wong body length from tip oF snout, nares 
ppen uncerolateraiiy: abdooinal, cranial. post narial 
regions und eyes appear darker below dermnl layer of 
golden Wophores covering most OF dorsum. except 
for urostylar region. where ubsence of widophorcs 
creates hroud, Uniform or broken darker bund; body 
wall sparsely flecked with simall chisters Ot 
metinophores in seme Specimens; soe brows 
palches over hal musculature; golden indophores 
denser aver Gills, snout, posterior rim of iris and just 
interior to urostylar tegion; young Wdpales. at stages 
75-26 with darker dorsum (ess dridephares). and 


layer of 


cOnLUSHNg bund of pale grey pigment Just unlorior 16 
darker upoptylar tegtons tail musculature sandy gal, 
or with Slight salmon rage, dark brown: patules 
seutiered ulang length of taik especiaily vlone 
wutertar dah limbs iicreasingly premented trom 
stage 34. 

Lider: body oveid: stout rouided: spiracle 
sinistral. broad at origin tapering lowards postero- 
Jorsal opemiug; oral dise direeted vertrally: vent tube 
dextrl; snout appears golden, layer oF copper/golil 
iridophores oVer majority of iris and lower part ot 
abdomen, chanps of tridophores outline lower cdee 
ofeach oil, cantinuing over venter: Gail musculature 
fairly thick. anteriorly, some seattered melaiuphare 
closiers overall but anterior lower edge; maim blood 
veasels, creviees between nmudeulir  ridgys 
pigmented? fins relatively shallow to moderuely so. 
clear. with Joint wolden hue. line Seatleresd 
melunophore clusters over dorsal lin ane some dusky 
pigment over ventral fine-venation unpigmented brit 
Visible: dorsal fin rises gradually (or more deulely) te 
preatest depth anterior to, or dt, mid-pome ol tail, nui 
Lip narrowly rounded, 

Ventral opaque copper/gold sheen over abdomen, 
heart and some clusters of iridophores over stdes vl 
each gill, anterior half of venter otherwise clear 
hinbs-and tal oseulature unpiemented. 

Oni dise (Pig, GA. oral dise ventral in direenon 
fuirly wide: bund of pupiliae surrounding entue 
muraint 239 rews fine papillae around anterior, up ty 
six around lateral and 3-4 around posterior margin: 
inpermost row oF papilhie around the posterior 
margin very slightly larger; bwo complete anterior 
and three complete posterior labial touth rows, equil 
in Jength (PL row aw hte sherter in somrety jaw 


BREEDING BIOLOGY L. BOOROULUNGENSIS & Lo LESUEUWRI “y 


sheaths moderately missive, wath 
heratinisation greater on upper sheath; central notch 
On Upper Sheath, ti some individuals there is a small 
Underling keratinised ledge visible, similiur to tit 
shown in Fis. 6B for Le boervelongersis but less 
prominent, and absent or worn in ofhers; diner 
munis of jaw sheaths serrated. 

Coloar af preserved larvae (composite deseription 
stiges 3)-41)° body colour untlorn) brown uve 
abdomen, cranial, vertebral and: post-nacial regions. 
lighter elsewhere; dermal body wall clear, spirsely 
Nevked with small clusters of melanophores in some 
specimens. some brown patches aver ntl 
museulite (dorsal view); abdomen opaque, fine 
layer OF mekinophores over gill region, patehy in 
some specimens. becoming spurser over snouly pris 
hlaeke fins mostly elewe venauun uopigmented bu 
Visible (lateral view): fine layer of melanophores 


urpund sides Of abdoinen, (ntestine Visible, rest of 


venter clear: limbs and tal musculature unpigmented 
(ventral view } 


Metumerpheasts 
Lierice bern eeayin 

Dyes laid on 3.411973 were first metunorphosing 
from I8A1974, alter a larval life span of 21/4 
months. Some were sill metimorphosmy by 
14... 1974, Numerous metamorphs were observed 
aoninidly in Devember/ianuary at Back Creek and 
Serpentine Creck during 1965-74. No tadpoles or 
ielanorphs were observed a late auton at Back 
Creek on 1oy.1973. Ten newly mectumorphosed 
frogs trom Serpentine Creck (stage 46, (97+) ranged 
from $4.0 - 17.5 min (mean 15,23). 


Literia lesueurt 

Known observation dites lor metamorphosis ure 7 
Apnl 1974 at Coco Creek (locality 5 Fig, 1B, 33° 08! 
15S, (507 06'40" By god 1s Jamnary 1977 at Dingo 
Creek (locality 7 Pig. 1B, 31° 49°50", 152° [8 On" 
Ey. Tava lite spun is not known. Seven pewly 
Metunodphosed Frogs (staye 46) trom Dinga Creek 
ranwed fropy E17 = 15.0 anim (eat 14,94). 


Discussion 


Papnlation declines, hatitat 

While the present knowin distribution lor Litre 
Hoormlaiveiwes dong the mount range COU 
in NSWoos. tram the Queensland border to) the 


Viclooun border (he disinbutot and agius of 


Litarin lest aeeds clanfieation, with a likelilgod 
of omy of more species beth involved (Moore 
Oli Heatwole eo uf. (1995) report J, 
haoroalonieusis as “widespread” jo the New 
Englind region but the records they provide are only 
those of existing Museum specimens spain “a 


degree ol 


perjod from early 20th century ta 1990". 
Furthermore. extensive surveys for regional 
conservation planning in north-eastern NSW 
undertaken since 1991 through the  casiern 


escarpment forests around Tenterfield, Armidale and 
Glen Innes by NSW National Parks & Wildlile 
Service (NP&WS), have failed ty locate this species 
(H. Hines, Queensland Departrnent of Environment 
& Ueritage pers. comm. 1997), Field observations at 
Guy Fuwkes River and nearby well-known loealities 
where the species was similarly abundant prior to 
1980, have imdicuted few adults ever the past 17 
years: The species could not be found in recent 
surveys at Ebor (type locality) god a hirge number of 
rivers i the area (M- Mahony. University of 
Newcastle pers. commit. 1997). Intense searches 
during 1995-96 in this area were who to no aya CR. 
Lars, University of New Boglind pers. comm. 
1907). 

Surveys since (992 jn upland forests such as Glen 
Innes, Walcha, Mt Royal. Dorrige, Teatertietd. 
Coolah Tops and south to Tumul/Turbarurn ba 
(turgcting frogs in areas where £. hocuolimgenyts is 
likely to oceur), also resulted i hone al this Species 
being found (bh Lemekert, State Forests NSW pers. 
comin. 1997), 

Surveys in the southern region near Tumul NSW 
by Hunter & Gillespie (unpub. located only (3 frogs 
an 30 November 1996, along an BOO in stretch of the 
Goobarragandra River. Extensive searehies along the 
Tumut River. where L. beervaloneensiy had been 
recorded in the 1940s and in F987 failed to locate 
jany evidence of the Species (Hunter & Gillespie 
unpub). 

Examination of over L.Q00) specyinens of 1, 
honmotongensis in the Australian Museum revealed 
that uuly five specimens bave been vollected since 
HOO: One at Wombeyan Caves (34" 19S. 149° Su! 
EB) two at Canimbula, Blue Mountains (33° 41'S, 
150° 12°F), one at Cos River (43° 28 8, 150704) FE} 
ant ane at the Abercrombre River, Governors Fat 
(a4) OF S$. 149° 41 EB), where Lo lesen are 
sympatric (IK, SmalloSydney University pers. com. 
1997), Aceordingly, the spevies has heen Wominuted 
for inclusion in Part | of Sehedule | Cendtingered 
Species) OF the Threatened Species Conservation Act 
1995 (TSC), 

More studies on current population numbers of £. 
badrohmeenscs over a mich broader range of Ts 
distribution are required to assess fucther the curred 
alaius Of this species, mm the Ligh) oF very significant 
frow deelipes oy the northern ranges between 
latiudes 31° 30° S and 24° 30S (north of the 
Macdonwld River ~ Pig. TAI, Fron present 
indications. the species uppears to have disappeared 
at ibe very lease fron its type locality and all known 
lowalites an the AcmidaleyGuvrwPornt Lookout 


42 M. ANSTIS, ROA. ALFORD & G. ROGILLESPIL: 


regions where itonce was abundant. The only known 
northern area where adults have been observed ts 
locality 3 (Fig. 1A) near Tamworth (1994) in streams 
aha) altitude of 450-500 im (M, Mahony, Liniversity 
of Newcastle pers, comm. 1997), much lower than 
the Armidale/Point Lookout region. Further field 
surveys. will beed to be undertaken to verify the 
frog's continued existence here, but if so, this may 
relate to the data being gathered on frogs in perth 
Queensland whieh indicate that most declines are 
umongs| upland riparian species uboye 400 m 
(Richards ef al. 1993), 

Some observations in southernmost localities. near 
Turmul in 1996 also indicate a distinet drop in frag 
sightings (Hunter & Gillespie unpub.) sugvesting 
the species ts ikely to be endangered throughout is 
general distribution. 


Litorta lesueuri 

This species has a broader current distribution thin 
J. hooroatargensis. Barker etal. (1995) indicate thut 
UL desist LWO Species are currently included under L- 
lesnenri, “one confined (o northenstern Queensland 
une (he other extending down the couwst as fir is 
Vietoria.’ 

Comparative popakition studies on L. leven? are 
heeded, especially where the minges of the twa 
Speeres overlap, to determine whether this spectes 1s 
also undergoing a decline in certain localities. 
Surveys by Hunter & Gillespie in 1996 im Kosciusko 
National Park NSW showed that the species was 
present ip only 1S our of 40 likely streams, which 
may indicate a possible decline, While 4, 
boonmlongensts is restricted to Towing streants 
generally above about 400 in ZL. deswenry can breed 
in siréums. Strewmside pools und even shims. from 
1100 4) (River Murtay and Snowy Kiver, ML 
Kosciusko NSW) to 40 m (Ourmbuh NSW), This 
ereuer versuoliny of altitude und breeding site 
selocnon may lave helped more populivions renin 
Thun for lL. boeroealorueryin 


Behaviour 

Neither Le feurvelengemsis mor L, lesueur® bas u 
yoru sae and each produces a sofl. low call obi 
serves OF short. repeated notes. Adulis are similar 
Nuiphalowiedily. beth breed in usseeiatian with 
Hlowrny sireains, ud have a similar tialjate winelr ps 
Adluple do fhe lotic environment 

Liteiia bevntaledtinensis agorevates Uiiler stones i 
hivee nuinbers dueine wire A aragp op Tu 
mutivadiits, was founel unger (he same stone beste 
Enis Creek NSW (locity + Fig, LA, Fle 440s. 
oF 2" Bion, bhai eet by fe Parke 
Populariuns here were Jarge, wath tom ah 15) frags 
ball sive clusses ohserved a 19 ei 1970 durin the 
day, under stones th tie eteek bed Males called 


during the day and night. The aggregation of L. 
hddredlongensis under rocks both during colder 
months and also in the breeding season during 
spriiig/summer, has. nol been reported for other 
Australian bylid species, Winter aggregation has 
been reported for Litaria svubolandulasa (Tyler & 
Anstis. 1983). (Tyler & Anstis 1975) and for /. 
péarseniand (Copland, 1960) (McDonald. & Davies 
1990), but neither species has the same habit of 
commonly gathering in groups under rocks in the 
stream environment during its breeding season. 
Literia fesueuri adults, found on ridge-tops in forest 
uway from the stream during winter, are not known 
lo aggreaate. Lileria beorodlongensix is simular to 
the stream-dwelling 2. spencer’ (Spencer, 1901) 
(Watson ef af. 1991) in its diurnal behaviour. ofien 
basking in the sun on very warm rocks in mid- 
stream. Amplexus and oviposition also occurred 
diurnally in one obseryation for L. hoarrolongensts 
Lireria deyuteurt has been observed busking in sun. 
but not as frequently as bas been observed lor J. 
spencer) or L, houradlonze ysis. 


Ovipasition/embryes 

The egg inass.of L. booreolenuens’s found partly 
udhering 10 suspended vegetation in a plasie bag 
does not give a inte indication OF the mode af 
deposition in the natural environment, as the adults 
had separated and were swimming vigorously within 
the hag, constuntly disturbing the egy muss. Barker 
er af (1995) reported that the ege mass ot L, 
fooroclungensiy is deposited “among rocks. 

While the northeastern Queensland form of /. 
Jesuew lays caps ina single clump of two er more 
Tavers deposited in circular nests excavated in sund at 
the sides Of Mowing streanis. (Richards Ae Alton 
1997) ees Masses ure so deposited among mocks ty 
streams where (he substrate is nob sand, The number 
af eyes mone clutch was estimated hy Richards & 
Alford to be 1200. Two other cliitches counted by Ss. 
Richards and M.A. ut Elphinstone Creek norith-Qld 
On Alvi.l996, numbered 1.738 and 1,674, 
respectively. Referring to southern populations, 
Barker ef al. (1895) state (hat several Huadred esp 
ire Jepositec in a solid) gelutious etunip whieh 
idheres te submerged recks or rhe hatam sediments 
We hive seen 2-3 inasses Juid beside evaeh athe, 
partly comneei oe. 

Whilst cee deposition marily 
Queenslind form apf. desterad nay he exe hdtest 
Nests ji sand bats, eggs af the southern lori have 
ony been observed i mchy areas. wher such 
excivalion is mgt possible As [rows have been 
observed urauud dais (og, ub deealigy 2 Fig. bt 
Where Ho nicks Were present. ane cudpales Found i 
sundy atrediows (locubitnes 1. 3. 5. 1 Pige FBS, further 
studs of the mode of deposition mi the Somuliont for 


silts ot the 


BREEDING BIOLOGY L, BOOROQLONGENSIS & L. LESUEURI Wa 


References 


is needed (9 determine whether or not nest 
excavation may occur al suilable sites. The mean egg 
complement of L. fesmear? was ereater than in ZL. 
hooroolongensix. Both species have a compact. 
gelatinous egg mass. similar in form to that of other 
streani-breeding liylid foes such as L. pecrsentiahd 
(MeDonald & Davies 1990), L. wibeglandulosa (Anstis 
& Litdejohn 1996), L. genimeculata (Horst, 1883) (as 
L, encnemis, Davies 1989), andl. spencer. The egg 
mass of L, fesuewrt is loosely attached to the 
substrate, while those of 4. ysubelandulosa, L. 
spenverd and Lo pearyoniana adhere more strongly. 


Larvde 

Although larvae of bouh species are the sane size 
atany given stage (Pig. 3) and appear superficially 
similar they differ considerably in shape. he 
tadpole of L. hoeroelongensiy has a relatively larger, 
wider ofal dise afd broader, more elongated, 
streamlined snout than 4. lesuear? (Table 1. Pigs 
4.5). The distances from the tip of the snout to the 
anterior rim of the eye and to the anterior rim of the 
naris are relatively greater in L. booreclongensiy and 
the body depth, dorsal fia dept and eye diamerer are 


IMARANTILLL Ch. Gihonsmin Gok FesunG, S$. (106) 
Observilions Of oviposition sites of the spotted tree frog Litt 
spencert, “In vhe Spettieht 2 fd td 


less. The eyes are positioned more medially and 
directed a litte more dorsally. The presence of the 
small keratinised ledge underneath (he central noich 
of the upper jaw sheath was voted in populations of 
both species, but nor found as consistently qur us 
prominently in Z. desveut. This feature has not been 
recorded for larvae of any other Australian lrogs. 
The jaw sheaths ure commonly more heavily 
keratinised in L. bvorvoloneensis, but this feature 
may be variable umongst populations, us has been 
observed io L. desuenr?. add ip northern and southern 
populations of the tadpole of L. verreawsi (Dumeril, 
1853) (Anstis 1976), 


Acknowledgments 


The Australian Museum is acknowledged for the 
loan of specimens. A grunt to M.A. from the Peter 
Rankin Trust Fund in 1996 has assisted this study, 
and is gratetully acknowledged, Valuable 
Observations on population declines of  F. 
hoeredlongensix were provided by H. Hines. P. 
Webber, K. Harris, M. Mahony, F Lemekert, J. 
Reesei, K. Thumm and K. Small, Constructive 
comments on the munuscripl were given by $. 
Richards, FP. Parker, M. Mahony. G. Watson and M, 
Davies. 


References 


AwsTis, M. (J974) An introduetion ja the study of 
Austratian tadpoles. Herpetfaniad 7, 914, 

(1976) Breeding biology and birval development 
Ol Liferie verreauee (Anurd, Flylidae), Trans RO Sie §- 
Anish 100, 193-202, 

(1994) The lirval deyelopnient of Lirarie 

brevipalmete (Anura, Mylidae), Mem, Old Mis, 37, 1-4, 

— & Lerrecious. M_1.11996) ‘Phe breeding biolagy 
OY Litoriee Subghindutosa ind Le citrepe (Amur 
Hylidae). anda re-evaluation of thei!) geoeraphic 
tistabulion. Quen. ROA Ant 120, 83-99 

BarKwie, 1, GRIGG. GC & TYLER. Me J. (1995) 7A Held 
guide fo Australian frogs (Surrey Beathy & Sones. 
Chipping Norton NSW), 

Copnann, SI (1957) Australian ree frogs of the genus 
Myla. Pree. Loe She. NSW 829-108, 

Davin. M, LIY89) Developmental bielogy af the 
Avsttulopupuiin fiylid hon datura ehevenius C(Anure: 
Hylidhed. frais, RSet. 8 Aas PV TPS 220 

& Riiakos, Sod. (TYO) Developmental brolowy 
obibe Australian hy bd frog Ave cares lav? (Crlinther), 
Trans RK See 4 Aust, V4, 207-271, 

Hicry PG. NLR Kean RAL COST) Multivariate 
wnalysis of ceoloyicu) communities” (Chapman & Chill, 
Londons. 


Piercuer. JJ. (1889) Observations on the oviposion and 
habits of certain Australian hatrachians. Proc. Linn. See. 
NSW ser, 2. 357-387, 

Gossnk, BL (1960) A simplified table for shiwing aura 
embryos and durvae with nutes on tdenuticaion, 
Herpetelagica VW, V83-190. 

Hiavrworl, HO DL Ravay. 2. WeiHeR. Po & Wik, G, 
(1995) Fauna survey of New England. (Vo The Frogs, 
Mom, Qld Mus, 38, 229 249, 

Mancous. Lo L990) Traditional tmorphonetrics pp. 77 
122 In Roni, FI & Bookstuin, [Musi “Proceedings at 
the Michigan Morphatogical Workshop, Spee. Pub. no. 
27 (Li Mich Mus Zool, Ann Arbor Michi). 

Martin. A. A Lintneiniis, Mob & Rawirssen. PA 
(i966) A key to che dius ces oF the Melbourne arcs. 
and dn addition to the anurin (aun, Wer Nab 83. 312. 
FAS. 

MeBosarn, KR, & Poyins, MM. C1990) Morptiotoyy: and 
higlopy of the Australien tree Woe Liner pencils 
(Caphind Amur: bhyldiiel Cami. Ro Sues, Aust 04 
145-156. 

Moorn, J A CM6t THe Hogs al caster New South Wales 
Ball Amen Mies. Net Hivn 21, 149-486, 


44 M. ANSTIS, R. A. ALFORD & G. R. GILLESPIE 
RICHARDS, S. J. & ALFORD, R. A. (1992) Nest construction WATSON, G. F., LITTLEJOHN, M. J., HERO, J.- M. & 


by an Australian rainforest frog of the Litoria lesueuri ROBERTSON, P. (1991) Conservation status, ecology 
complex (Anura: Hylidae). Copeia 1992, 1120-1123. and management of the Spotted Tree Frog (Litoria 

, McDonaLp, K. R. & ALForD, R. A. (1993) spenceri). Tech. Rep. Series 116, Arthur Rylah 
Declines in populations of Australia’s endemic tropical Inst. Environ. Res., Dept. Cons. Environ., Vic., 40 


rainforest frogs. Pacific Conservation Biology 1, 66-77. + vi pp. 


TRANSACTIONS OF THE 


ROYAL SOCIETY 
OF SOUTH AUSTRALIA 


INCORPORATED 


VOL. 122, PART 2 


A NEW GENUS AND TWO NEW SPECIES OF GALL MIDGE 
(DIPTERA: CECIDOMYIIDAE) DAMAGING YOUNG BRANCHES 
OF EUCALYPTUS SPP. IN SOUTH AUSTRALIA 


By PETER KOLESIK* 


Summary 


Kolesik, P. (1998) A new genus and two new species of gall midge (Diptera: 
Cecidomyiidae) damaging young branches of Eucalyptus spp. in South Australia. Trans. 
R. Soc. S. Aust. 122(2), 45-53, 29 May, 1998. 

Two new gall midges are described from galls on young branches of two Eucalyptus 
species in South Australia and a new genus, Okriomyia, is described to contain them. 
The new genus belongs to the tribe Asphondyliini and the subtribe Schizomyiina. It 
differs from other Schizomyiina in the shape of the aedeagus, the solid tooth of the 
gonostylus and the cerci-like female tenth tergite. Okriomyia schwarzi gen. et. sp. nov. 
was found on Eucalyptus gracilis and O. flabellidentata sp. nov. on E. cosmophylla. 
Infested branches fracture at the site of the gall as the trees mature. Males, pupae, and 
larvae of both species and the female of O. schwarzi are described. The new species 
differ from each other in the morphology of the male genitalia, the pupal face, and the 
pupal prothoracic spiracle. A key to the Australian genera of the tribe Asphondyliini is 
given. 

Key Words: Gall midge, Cecidomyiidae, Okriomyia schwarzi, Okriomyia 
flabellidentata, Eucalyptus gracilis, Eucalyptus cosmophylla, South Australia. 


Transactions af the Reval Seviety of S. Aust (1Y98) 122(2), 45-53 


A NEW GENUS AND TWO NEW SPECIES OF GALL MIDGE (DIPTERA: 
CECIDOMYIIDAE) DAMAGING YOUNG BRANCHES OF EUCALYPTUS SPP. IN 
SOUTH AUSTRALIA 


by PETER KOLESIK* 


Summary 


Kovesin. P.( 1998) A new genus und two new species of eall midge (Diptera: Cecidomyridae) damaging youny 
branches of Kucelypris spp. in South Australia, Trans. A, Soe. S. Aust, 122(2). 45-53. 29 May. 1998. 

‘Two new gall midges ure deseribed from galls on young branches of two Licalvpius species in South Australia 
und anew genus. Okrmnwia, is described to contain Mem, The new genus belongs to the ibe Asphoniylinn 
und the subtribe Schizomyiina. lt differs from other Schizamytina in the shape of the aedeagus, the solid tooth 
of the vonostylus und the verci-like female tenth tergite, Okriomyta sehwers gen. cl sp. now was found on 
hawalvprs graciis an O. flabellidentata sp. noy. on E. cosmophylta. Infested branches fracture at the site of 
the wall as the trees mature, Males. pupae, and larvae of both species and the female of Q) sefnvarc are 
described. "The new species differ from each other in the morphology of the male genitalia, the pupal face, and 


the pupal prothoracie spiracle. A key to the Australian tenera of the tribe Asphondyliini is given, 


Rey Wokns: Gall midue, Cecidamyiidae, Okrionivia seliwarsi Okrionyia flebellidentata. Bucalyyitus 


grata, Eucuiypray cosmoplivila, South Australia 


Introduction 


Nacalypoes, the dominant genus of most Australian 
woodlinds and forests, hosts a whole suite of gall- 
lormings inseets, many of them undescribed. The 
present paper describes two gall midges, new to 
science. which were found damaging young 
branches of two eucalypts in South Australia, Galls 
of Okriomvia schwarei sp. nov. on Eucelyptus 
eracilis TF Muell, (Pig, 1) were found at two 
localities: Nadda. in the southern part of South 
Australia near the Victorian border and Porestville, a 
suburb south-west ol Adeluide. Galls of O. 
Mabellidentita sp. nov. on LL cosmophylla P. Muell. 
(Mig. 2) were found at Cleland Conservation Park, 
near Adelaide. The newly-described gall midges 
were found only in moderate abundance. However, 
heavy infestations could have the potential to impact 
seriously on the population dynamics of their hosts, 
since (he infested brinches fracture at the site of the 
galls as the trees mature 

Eucalyptus gracilis sa 3 - [2m high shrub or tree 
distributed through the mallee belt of continental 
southern Australia. [is an ari zone species usetul 
for firewood und crosion control and is) highly 
revarded for honey production (Cunningham e7 al. 
1981, Chippendale 1988). Ibis offen used in urban 
planting. 


© Department oF Horticulture. Viticuliure ind Oenology Faculty of 
Nerieultucal diel Nateral Resource Serences, Che University. ut 
Adeline PME OT Glett Osmond S. Aust. S064. 


Fig. |, Gall of Okriomvia schwerce sp. ney, on 
brinch of Arecelypris gracilis, Seale bur = 20 min, 


young 


at 1 KOLDSIR 


nN = 
Pik, > Gallo Ohkrionvie flabellidentata sp. nov, on youne 
brinch of Rac yyas cosmaplivila, Seale bar = 20 niet 


Eivalvprs cosmophylla is a South Australian 
shrub or tree, usualy 5 LO high, that oceurs from 
the Mount Lolly Range to the Fleurieu Peninsula and 
Kangaroo [sland in open shrubland. low. open forest 
and heathland nearthe sea (Chippendale TO88) Tis 
Widely dsed in urban planting, 

The new gall midses do not resemble any Known 
eens so a new genus has been erected for then. 
Ohrfomyia becomes Australia’s fourth known gers 
of the ibe Asphondyliint und the 
venem endemic to Australia. A key fo the Australian 
eeperd of Asphondy lint ts given in the present paper, 


Material and Methods 


Gls on branches Of Evealvyptus viacilis were 
Porestville (19477999) abel Naddy 
(b2eyi T9960), Two, one, Hires. Tour and one galls 
Irom branches of A. cesmophyila were collected at 
Clehind and Morintta Conservation Parks 27,41 1992. 
TS 199F 5S. und 12 P99S. and 23.1 1997, 
respectively. Ja the labortory the pulls were cut open 
and the larviie processed i two ways. “Ao small 
number was preserved in 7O% ethanel AO darter 


collected at 


Wind of 
Sulnzomyiini. ad subtribe consisdnge exclusively of 


number Was Wansterred ilo tearing pots where Ure 
larvae dug themselves into wet sand. Pupation took 
plice i (he sand, Several qales and ternales 
emerged from the galls from Ay graeiliv, OF the galls 
collected from E. vesmoplydie adults emerged only 
from the suinple collected on 23.11.1997 12 mules 
und ong femules. Emerged adulis were preserved 
together with their pupal skins i 70% cthaeiel 
Microscope mounts of the type series were prepared 
weording to the feehnique outlined by Kolesik 
(19950). The type series and other material retained 
in 70% ethanol, together with dried galls, ae 
deposited inthe South Austithian Muscum, Adelnide 
{SAMA}, the Australian National Inseet Colleetion, 
Canberra [ANLC| and the State Herbarium of South 
Australia, Adehiude [SHUSAJ. Descriptions aid 
measurements reler to the holotypes and paratypes 
Terminology of adult tnorpholosy follows That of 
Gaené (DOS) ): larval terminology Totows that ot 
Cane (1989) 


Genus Okriomyia wen. nov. 


Type species: Gkrioivic schwirel sp. mov. 


Adtilty 

Heeul. Fye haces heaasonoid, eye bride H-8 
fieets longs medially, Antenna with 12 Thigelloneres, 
distal ones nol shortened. Plagellomeres eylinadrical. 
sessile. first ane second not fised. with short sehuw 
ind hearing low, finely reticulate circumlila. Suape 
4s long as wide, pedicel hulfias long as wide, Labella 
hemispherical, each wilh several setae, Palpus with 4 
segments. 

Theres Wings: Rs joing Coat apex. sliehtly 
bowed anteriorly, Rs absent. Ry joining C neve mia 
length, Cu forked, birst Girsomere lacking: vente 
distal spine, tarsal claws Simple, as long as empodia 

\hdomen. Tergites | o- & with selie eventy 
distribuied, forming dense row posteriorly, Sterna 
| Hot sclerotized, aselose; stermites 2 - 8 willl seuie mM 
iWO separate areas: wide, anterior field and narrow, 
posterior band. Pemale abdominal sternite 7 15) % 
stermile @ Male lemminialis gonocoxite with apico 
ventral lobe: genostvlus short and wide. with tool in 
form ot serrate plate no more strongly pigmientea 
than remuinder: cereus hilobed, deeply divided 
medially, eniutyinuted posteriorly, with several 
posterior setoe; parameres. small, sefuses hypoproct 
WITH posterion Mari coneuve, each faberd lobe with 
J ~ 2 apteal setae, aedeagus comprising Iwo pits; 
dorsal part robust, conical, ventrally eovered with 
selerotived Vili on apical thicek ventral part sinmeerh 
Thin ie katerul view, shallowly enurgnded apieatly 
in dorso-yventral view, usetose. Pemule terminalia 
OVIPastlor short, Peshy, tergunmy 9 and stevia, 9 
sclerotizeds terguin LO in form of two hinge, cory 


NEW Cb 


like lobes. more selerottzed uinteriorly, evenly selose; 
verer large, diserele, more selerotiaed: posteriorly, 
evenly selose: hypoprect small hilabed. cach lobe 
with apical sen. 


Pipi 

Aen hors strongly piemented, cephalic 
swellings, fugral protuberances. prothoracic spiracle, 
dorsal spines of abdomen stghily pigmented: 
abdominal Skin not pnented Adtennal horns blunt 
onantlerior suchice, produced anteto-ventally Hitoau 
aeule ridge, Cephalie selenite wilh pair of swellings 
shorter (hin antennal horns. Cephalic pair ef papillae 
With long serie. Moos with one or two sclerouzed 
protuberdnees on cach side, One Of two lower facial 
papllic With seni, one Of three lateral papillae with 
sel. Abdominal seginents | - 7 awilh pair of setose 
ventral papilie. 2 pairs of setose pleural pupilhw, 2 
pairs OF usectose and pair of setose dorsal papillae, 
Abdominal seements with pair of ventral papillie, 2 
pits Ol pleural pupiite, purr of dorsal papillae, all 
selose, Abdominal seunients 2 - & dorsally with field 
cf strong. one- or tweepetited spies on auiterior 
tall, 


Lave 

Inlewument covered wilh finy, Sparse Spiculac, 
Head: strongly selerotived, postern-luleral apodemes 
longer than head capsule, antennae 2 x Jomger than 
basal width. Neck segment with pair of dersul 
papillie, Thomeie segment, with pair of ventral 
pupils, 2 pairs oF pleural papillae. pair oF sternal 
papilhie. 3 pairs of lateral papillite. 2 pins of dorsal 


pupillae, Spatula bilobed, with shut. Abdominal 
segments. |= 7 with paitol veuteal papillae. 2 pais of 
pleural papitlae, 3 pairs of dorsal papilfie, 


Abdominal segment 8 with pair of ventral papithie, 2 
pues Gb pleural papillae. puir of dorsal papilhue, 
Terniindl segment with pair of aml papillae on short 
lobes, pair of terminal papillite on prolonged lobes, 
ATT pupillie aselose, 


Livintelasy 

The prefix, “Okfio?” ts fram the Greek okries. 
Meanie roughness, referting to the pigeed ventral 
surfice OF the dedeavus dnd distinguishing the new 
genus trom other Sehimomyimi The suttix -myiae 
is Cireek for fly, 


Bematks 

Okrtonvie gen. hoy, belongs ta the trike 
Asphondyliini on the basis of the following shared 
dpomorphies: the presence Of a ventrosupied! lobe on 
he SondeoXite with wonestylis consequently 
sihdatcal dorso-ventrally; the short, quadrate 
pomosty lis, He presence of puramercs: and the lange 
female slempite 7 thal iy 1.5 4 is long as sterniie 6. 


TDOMYTDAE PROM EUCALYPTUS 47 


The new genus belongs tothe subtiibe Sehivorny iit 
beewise il lacks a yentro-apical spine on the first 
lursomere. bas male parameres. hus a short Meshy 
ovipostor and the pupil miegumentis unpigmented, 
The new genus is unique among the Sehivomyiina 
because of the divided acdeapus. the solid loath on 
the gonostylus and (he cerci-lke female tergumt 1 
The Australian gents. Leeinoricariio Fell the oat 
other genus assocniued with galls on /vealyptan spp. 
(Kolesik 1995a). most closely resembles Gkrionie 
wen. now OGdrtoumia shires wilh Keciedicarna the 
lone Tobes on the lerminal larval segment and the 
Neshy ovipositor with divided ceren. whieh 
represents the most plesiomoerphic ovipusitar in 
Asphondyliini. 

The tibe Asphondy lit is Known in Australia front 
12) species distributed among four gener 


Ayphandviia Loew, a large. eosmiopolitin genus. 


belonging lo the subiribe Asphondyhina, contains 
sever species: A, aithocercidty Kolesik (Kolesik er 
a V99T), A. dodonevae Kolesiky (19950). A, 
a arimis Kolesik (1997), A, lal Fava (1916). 

\. inte Kolestk (L997. AL loews) Skuse (LASS) and 
A mibieunda Skuse (1888). The other three genera, 
all belonging (oO the subteibe Schizornyiina. are 
Knows only from Australi and contain five species: 
Bnvineticornitoustalasiae Felt (POU) Eo malarska 
Kolesik (1995u). Shuserurtee atlecusuarimee Kolesik 
(1995b) and the new species. Okrinnvid selec 
and Ch fubellidenrata 


Key to Australian genera of Asphondytiini 


1. Pirst farsomere With spur male pumimiere absent) fenile 
with pair or dorsal Tobes at base ob needle tite 
eV positon, ptipal slain conmplerely piaiiented 

as = 1 Asphandylia 
First tursomere without spur: male paramere presen 
female witholt pur Of clarsal febes at base ol ovipostlot 
or owiposrlor not oeedle-like; pupal skig mat preme med 
ona leastubdomen., (al owepiefoes email estate re seebitven so 


1 Pitee fteriniinal female Thigelotieres slickossively anu 
progressively shorten; ovipostor needle-like, mule 
purneres Tiree, us wide as postcrion lobes of coerce, 
pupal cephalie swellings longer tian antennal horns 
p cobkinens otto eres porrerma acto comeaphracerna vorcpon xo egypt toe REELS FAL 
Three tering female Hagellomeres subequal in length: 
ovipasitor shert, fleshy, wath untiised cerch mate 
purdnieres smth meh narkower than posterior lobes at 
cerci! pupal cephulie swellings shorter than antennal 
TIGHT PT ns coceccenenncasiiilers conse nsaitidesMensriestleescesibintecen gonbhiidenn al 


‘aa 


Tooth on sonosty lis ConsEstiqg ol seven separate leah, 
fermale with pair at smal) dorsi lobes posterior (0 
eighth terete: fareest dorsi sping vn pupa serrated 
Upiclly on ved EOC Orne 
‘Tooth an AYNsIYLUS “sot im tai ut Wo pllwen female 
withbut pir of dorsal lobes pustercar Wo with wererbe 
Uiiesal spines: cn PPA ONE Ort wed-perntedl A krienive 


1s P KOLRSIR 


Okriomyia schwarzi sp. ny, 
(PIGS 1, 3-6, 19-23, 27-30) 


Hoalowpe: &. Nadda. South Australia {34° 37" 
40° 54° FE], Lali, (990, reared by PL Kolesik fram 
branch gallon Aucalyptas gracitiy b Muell, larva 
calleeted [2vii 1996 by J. Sehware, [21338 
|JSAMA| 

Paratypes; 255, 3 9°. 4 pupal skins [SAMA, 
L21SA9-121347], 2 8A, 2 FP, 3 pupal skins 
[ANIC], same ditt but emerged 12.-20.Vin, Love: 4 
larvae |SAMA, T21348421350). 2 larvae [ANIC], 


collected with holarype 


hie. 3, Gall OF Oknaniiin sofivercr sp. nov. on Bucalypins 


wis ~ longitudinal section. Arrows mirk larval exit 


hiles., Scale Py 10 na. 


Other niuterial (SAMA: 4 od. 24 23 pupal 
skins, 3 pupuc. sania datas 17 larvae, gall collected 
Will holotype: 2 9°22 pupal skins, Porestwille 
South Australia |44'56°$. 14836 Ef. 23.1, 199s. P. 
Kolesik, reared from branch galls on 2. ereesliy 
larvae collected POAT 993; 4 gulls collected with 
holotype [SHSA I, 


Deseriprion 
Male (bigs 4-6, 10-15) 
Colour antennae grey; head black: thotax brown: 
lees yellows ahdomen with selerotized parts and 
Wing 
length 2.2 mm (19 9 25). Genitals gonocoxite 
covered wilh short setae, with 2 short, thin, posterior 
lobes; apico-ventral lobe oon 
wonoeoesite long, deleulate: tooth oon gonostylus 


setae black, Hon-sclerotized parts orange 


dorso- medial 


narcow. finely serrated: aedeaetis narrow distally in 
lateral view; hypoproct with durge lobes, ats loug its 


Hedeagus. 


Fennile (Fuss 10-20) 

Wine length 3.0 mn (2.8 © 3.3). Circumlila oo 
Nagellomeres uboul ball density of mile ones. 
15 x (63 -b.0) lonver than 


sy 


Abdumind stern 7 


sternilc 6, Setac on cere 2 X shorter and much denser 


thin on lergite 10, Ovipositor as Jong as lergites 7 
und § towether, Colour as in male. 


Pupii (Figs 21-23) 

Colour: Antennal horns brown, cephalic swellings. 
facial protuberances, prothoraeie spiracle. dorsal 
spines pale brown, abdominal skin grey. Total length 
4.3 mm (3.8 - 4.0). Antennal hortis 86 pin (77 ~ 109) 
long. Cephalic sewe lol tin (138 - 18h) longs 
Cephalic swellings 46 Win (36 © 65) Jong, Upper five 
with 2 pairs of sclerotized protuberiunees, inner pail 
SI pin (48 ~ 54) long, outer pair 30 pam (29 — 38). 
Selae on lower facial papillae 122 pin (103-143) 
long. Prothoracic spiracle with slight, eridual curve, 
244 Hin (206 267) long, trachea ending al uper, 
Learvai (Hiss 27.30) 

Colours pink to ofange. Total length 3.9 mo (44 
7.8), Head capsule width at base 92 fim (90 ~ 94), 
lengih 70 pm (63 - 74). length of postero-laleral 
upodemes 116 tin (110-127). Antenna 26 pum (25 - 
27) long, Sternal spatula 445 po (461 - 543) lone. 
With apical enlargement (OO pin (83 — 130) wide 
depth of incision 46 Lim (29 - 68), Termmal lobes 
160 um (113-233) long 


Etyinology 

The speaies is named after the collector of the 
lurval stage of the Wpe specimens. Julie Schwarz, 
Department of Plant Serenee, University of Adelaide 


Gall and bielows 

Young branches of Anecelvplus wraciiy are swollen 
lo form galls 8 - 20mm in length and 7 - 9 mim in 
diameter. with outer walls 1 - 3 rim thick (Fig. 1). 
The gall outer surface ts scabrous, reddish brown i 
colour, Inside there dire 1 - 5 ovoid chambers, each 
occupied by | = 13 larvae, Larval colour may yviry 
from pink to orange between chambers of the site 
wall but is the same within a chamber, No association 
berween the colour and the age of Jarvae was 
apparent. Gall walls contain less woody Ussue than 
Wnaffected parts Of the branch, which results i the 
will being springy lo the touch and cruhehy when cut 
with a kaife., This characterisuc is shared with alls 
OO. flabellidentata. When the kirvae are Tally 
#rown, they Jeave the galls through one or (we 
wireular openings that develop in cach chomber 
(hig. 3). Pupation tikes place tn the stil, 


Okriomyia flabellidentata sp. nov 
(FIGS 2, 7-9, 24-26. 41-34) 


Holotype: &. Cleland Conservation Park. South 
Australia [34° 58° 8. Tak! 42" Ri, boa bee? P 
Kolesik. reared trem beaneh gall on Brealyprits 


NEW CECIDOMYIIDAE FROM EUCALYPTUS AY 


Fives 4-6. Male of Okriomyia sclwwersi sp. nay. Pig. 4. Acdeagus in frontal view, Fig, 5, Aedeagus in lateral view. Pig. 6, 
Genitalia in dorsal view (inner part of cerci diagrammatically cut out for beter clarity). Pigs 7-9. Male of Okrionn te 
flabellidentate sp. noy. Fig. 7. Acdeagus in frontal view. Pig. 8. Aedeagus in literal view. Fig. 8. Genitalia in dorsal view 
(imer part of ceret diagrammatically cut out), Seale bars = 100 tim. Abbrev.: a. aedeagus: e. cercus: ga, gonacaxal 
apodeme; ge. gonocoxite: gs, gonostylus; h. hypoproct: 9, paramere, pa. parameral apodeme. 


50 P. KOLESIK 


Figs 10-20. Okbionrvia scliwarel sp. nov, 10-15 male, 16-20 female. Piz. 10. Head in froma view. Pig. Pl. Wine, Pig. 12. 
Last three Magellomeres. Fig. 13. Sixth flagellomere, Fig. 14, First tarsomere. Fig. 15. Last tarsomere with claw and 
empodium. Fig. 16, Sixth flugellomere. Pig. 17. Last three flagellomeres. Fig, 1S. End of abdomen in lateral view. big. 
19, Oyipositor in lateral view. Pig. 20. Ovipositer in yentval view. Seale bars = LOO im 10. 12. 17. 19.20: S00 pm |: 
50 pon 14-16: 200 tm Ls, Abbrev; c, cercus: h. hypaproet: 5. sternite: & tergite. 


NEW CECIDOMYIIDAE FROM EUCALYPTUS 


\ \ 
. y ) dN ) ° 
\ 7 \\ 
\ | \ . 8 
\ \ (oe) os \ @ D ms 
* Sa . 9 Ne 
f \ " 
\ | fret 
{ \ I, | 
i SS. \ , 
i | \ if 
| y i 
32-——_ 33-— 


ce 


Vigs 21-23. Pupa of Okriomyia schwurci sp. noy. Fig, 21, Anterior part in dorsal view. Pig. 22. Anterior part in lateral view, 
Pig. 23. Prothoracie spiracle. Figs 24-26. Pupa of Okriomyia flabellidentata sp. noy. Fig. 24. Anterior part in dorsal view. 
Pig. 25. Anterior part in lateral view. Fig. 26. Prothoracic spiracle. Figs 27-30. Larva of Okrioniia schwarci sp. nov, Pig. 
27. Wighth and terminal abdominal segments in ventral view. Pig, 28. Eighth and terminal abdominal segments in dorsal 
view. Fig. 29. Head in ventral view, Fig. 30. Spatula with adjacent papillae. Figs 31-34. Larva of Okriomyta 
flabellidemata sp. nov. Pig. 31, Bighth and terminal abdominal segments in ventral view. Fig. 32. Eighth and terminal 
abdominal segments in dorsal view, Fig, 33, Head in ventral view. Fig. 34. Spatula with adjacent papillae. Scale bars = 


200 bun 21, 22, 24, 25, 27, 28, 31, 32: 100 Lm 23, 26, 30, 34; 50 um 29, 33. 


52 P-KOLESTR 


casnuiplivilea b Muell.. larva collected 23.17.1997, 
121351 |SAMA],. 

Puratypese 2.33, 3 pupal skins (SAMA, 121352- 
121356). 2 22,3 pupal skins [ANIC], same dita bul 
emerged! (5-17.11, 19972 3 larvae [SAMA, J21357- 
121359]. 2 larvae [ANIC], collected. with holotype. 
Oren material (SAMA): (all collected: fron, branch 
sally on A. casmophylla by Po Kolesikhe 7 oct. 4 
pupal skins, sume data bul emerged |-f4iv. 997: 12 
larvae, gull collected with holotypes 3° larvie. 
Morialta Conservation Park [34° 54° 8. 138° 44 0], 
27K, 1992; 9 hurwac, Clehinal Conservation Park. 3, 
& FQN, 1995; wall, Cleland Conservation Park, 
73.) 1943 [SHSA]. 


Deseviprion 
Male (bigs 7-9) 

Colour: as in QO. sehwears Wing length 2.90 in 
27-40). Genitalia: gonueoxtte covered with long 
sche. WHT Awa Shork, posterg-dorsal lobes, one thin, 
One Wider apleo-wentral labe on gonocoxite stot, 
routed, tooth on gsonestytas wide. coursely 
serred: acdeags wide distilly ja lateral view: 
liypoproct with thin lobes. much shorter chin 
Wedeagus, 


Fennile 
Unknowea 


Pupe (Pigs 24-206) 

‘Total length 3.9 fain (3.7 © 4.0), Antennal hors 62 
um (St - (15) long. Cephalic setae 147 um (137 
1605) long. Cephalic swellings 25 tm (20 - 29) Jong. 
Upper lave with parrof sclerotized protuberances, 31 
jn (25 - 34) long. Setae an Jawer facial paplac 38 
frm (32 - 45) lone, Prothorieie spiriele bowed: il 
distal third, MO um (174 204) Jong, trachea ending 
abapex. Otherwise as in Qn se/warst. 


hein Wigs 31-34) 

Colour pink (o orange. Total length 4.4 pam (3,7 - 
5.0), Head eapstile width ait base 99 Lh (95 = 102), 
length 7O Lin (69 81). Teneth of posteroshiteral 
apodemes 125 pny (LOO - 140), Antenme 24 pn (24 - 
25) Jong. Sternal spatula 425 pm (398 - 475) long, 
With upiedl enlargement 127 pun (LOS - 14) wide, 
depth of ineision S4 ta (50-59), Terinmal lohes [48 
pun (2b = 160) long, 


Livros 

The name "tlabellidentata™ is i eompound Palin 
ddjecrive Tram “flabellun’’ meaning lao. and 
“dennis. meaning toothed. refering ta the shape ol 
The tooth on the gonostylits. 


Gall wid dielagy 
Youn branches of Auacelyday cosmoplvila are 


swollen ti form galls 10-70 tim in length and (0 = 
1S mm in diameter, with outer walls 2-4 imo thick 
(hig. 2), The gall outer surface is smooth to scubrous. 
green to brown i colour, Inside there ure | - 4 
inegulirly-shauped Chambers, each vecupied by 5 - 
IS larvae. Pupation takes place in the soil. The gulls 
ren tecornisable on the branches for several 
years afler (hey have been formed. Many branches 
hater fravtuwe at the site of the gall since the gall 
Hissue as less rigid than that of the tree. The same 
phenomenon was observed iW On yefwerci, The galls 
OF O fibellidentata on i. cosmoply tie are common 
in the nature conservation parks around Adelaide, 


Remarks 


The two new species difler frome cach other in 
several characters. The males of OAriemiyta selves 
Haye wnarrew loth on the 2onosiy lis, the hypoprect 
is as long as the aedeagus, the gonocoxite has two 
Wil, posterior lobes dorsa-medially. wid the apices 
ventral lobe om the pgonocoxile is aceite: Thre 
mules of OQ. flebelfedenane have w wide tooth oa the 
gonustylus, (he bypoproek is mueh shorter (at che 
vedas, the gondcoxite has no posterior lobes 
dorsa-mediilly bul tas one thin und one wide lobe 
dorsally, and the upice-verntral lobe oan the 
gonccoxie is short and rounded. The pupae ab c, 
vohwurs! have bwo pairs ol sclerotized protuberances 
on the upper faee, long setae on the lower facil 
pipilac, and an evenly-henl prothorucic spiracle 
The pupae of O. ffebellidentata have one paw of 
sclerotized protuberunees on the upper luce, short 
setae on the lower facial papillae, dnd a distally 
bowed prothoricie spiracle, 

Thal as many as 12 males and ne females were 
reared from the one gallon kieahyres cosmophaila 
eolleeted JAI 1997 suggests that females of G. 
flabelidentata produce unisextual progeny, a 
phenomenon found im Centarinne vorghicole 
(Coquilletty (Baxendale & Teetes TYh)) and 
Cystiphora sonchi (Brent) (MeClay 1996). In arder 
lo verily (he production of unisesual progeny iO, 
flabelhidentata, and perhaps ©. sehwerg. inure 
adults have fo be reared from: separate galls. This 
Indy require rearing larvae rom a larger number of 
Wills ds OL flabel/idertata seeiis Hot lo be an easily 
reurcd species. Prom some 1500 lirvae originating 
frome TO galls inehided in this work only the }2 mates 
emerged. 


Acknowledoments 


The Department of Environment and Natural 
Resourees, South Australia kindly permitted 
Collecting in the Cleland and Morialia Conservation 
Parks, M,C, O'Leury, Stale Herbariin of South 


NEW CECIDOMYIIDAE FROM EUCALYPTUS 


Australia Adelaide, courteously identified the host 
plant species. Special thanks go to J. D. Gray, 
Department of Horticulture, Viticulture and 
Oenology University of Adelaide and R. J. Gagné. 


4 


Systematic Entomology Laboratory USDA 
Washington DC USA, for commenting on an early 
draft of the mansucript. 


WwW 


References 


BAXENDALL, EP. & Trevis, G. L. (1981) Production of 
unisexual progenies by (he sorghum midge, Contarinie 
sorehicola, Auns. Entom. Soc, Am. 74, 412-413, 

CHIPPENDALE. Go M. (1988) Eucalyptus, Anguphora 
(Myrtaceae) pp. 1-447 In George, AS, (Ed.) “Flora of 
Australia” Vol. 19 (Australian Government Publishing 
Service, Canberra), 

CUNNINGHAM, G. ML, Mutinam, W. £., Manriorer, PL. & 
Lreicu, L. H. (198l) “Phints of Western New South 
Wales” (New South Wales Government Printing Office, 
Sydney). 

Epwarps, FW. (1916) Two new Australian Diptera, Av. 
May. Net. Hist. 103, 496-502. 

Ping, LP. (1915) New genera and species of gall-midges. 
Proc, US Nath Mus, 48, 195-211, 

GaGnh. Ro J (L981) Cecidomyiidae pp. 257-292 In 
MeAlpine, J.P, Peterson, B. V.. Shewell, G. O., Teskey. 
HI. Voekeroth, J. R. & Wood, P.M. (Rds) “Manual of 
Nearetic Diptera.” Vol. 1 (Canadian Government 
Publishing Centre, Quebec). 

_ —— (1989) “The Plant-Meeding Gall Midges of 
North America” (Cornell University Press. Ithaca, New 
York), 


KOLestk, P. (1995a) A new species of Lecineticornia Pelt 
(Diptera: Cecidomyiidae) on kucalyprus fasciculasa in 
South Australia. /. Aust ent. Sov. 34, 147-152. 

(1995b) Shusemvia alocusitarinde, a new genus 
and species of Cecidomyndae (Diptera) damaging lateral 
branch buds of drooping sheoak, Al/locesuariid 
verticilata in Australia. Trans. R. Soc. S, Aust, 119.4140, 

(1995¢) Asphondvlia dodonacue, anew species 
of Cecidomyiidae (Diptera) damaging leaves and 
branches of hop-bush, Dodonaea viscose (Sapindaceae) 
in Australia. Zoid. 119, 171-176. 

~ + 6 il (1997) Two new species of Aspliondylta 
(Diptera: Cecidomyiidie) trom Iilosercia spp. 
(Chenopodiaceae) in South Australia. hid. 121, 59-66. 

_ Wiirtremore, R, & Stach, H, M. (1997) 
Asphonidylia athocercidis, a vew species of Cecidomyndac 
(Diptera) galling flowers of Anthoverciy  litterce 
(Solanaceae) in Western Australia. /bzel, 121, 157-162. 

McCay. A. S. (1996) Unisexual broods im the gall midge 
Cystiphora senchi (Bremi) (Diptera, Cecidomyiidae), 
Canad. Entom, 128, 775-776. 

Skusy, FA. A. (ESS88) Diptera of Australta. Part 1. Prac 
Linn. Soo, NSW. (2nd Series) 3, 17-145. 


WITHIN-NEST BEHAVIOUR IN A EUSOCIAL AUSTRALIAN 
ALLODAPINE BEE EXONEURA (EXONEURELLA) 
TRIDENTATA HOUSTON (APIDAE: XYLOCOPINAE) 


By ZETA STEEN* & MICHAEL P. SCHWARZ* 


Summary 


Steen, Z. & Schwarz, M. P. (1998) Within-nest behaviour in a eusocial Australian 
allodapine bee Exoneura (Exoneurella) tridentata Houston (Apidae: Xylocopinae). 
Trans. R. Soc. S. Aust. 122(2), 55-63, 29 May, 1998. 

Understanding the processes involved in the evolution of social behaviour has become 
one of the most challenging areas of modern biology. Since bees and wasps exhibit a 
variety of social organisations they are particularly useful for addressing social 
evolutionary questions. Allodapine bees are especially useful for examining social 
evolution, since species display varying forms of social organisation from solitary to 
eusocial. This study examines within-nest behaviour of Exoneura (Exoneurella) 
tridentata, a native Australian allodapine bee. This species has the largest known 
colony sizes of any allodapine bee and exhibits striking size variation among female 
nestmates suggesting that sociality may be regarded as highly eusocial. 

Key Words: Exoneura tridentata, social behaviour, allodapine bees, aggression. 


Piawurrions of de Navel Nentety ab hive (1QO8L 122(2). 55-63 5S 


WITHIN-NEST BEHAVIOUR IN A EUSOCIAL AUSTRALIAN 
ALLODAPINE BEE EXONEURA (ENONEURELLA) 
TRIDENTATA HOUSTON (APIDAE: XYLOCOPINAE) 


by ZPTA STEE 


’& Mictiart P Scuwarz 


Summary 


Sihtn. Z. & Scuwans MOT Cfg9Ss Withinenest behavioul ia etsicial Australian alledapmne bee fe tenenc 
(vareuredta) iidentatoa Houston tb Apidae: Xylocopiuie), Tris A Seo SN Advi 92202), 55-04, 29 May, 198, 

Understanding the processes invelved in the evolution of sociil behaviour has become ene of the most 
Chillensing areas of modern biglowy, Sinee bees and wasps exhibit a variety of sect) orgunrsations they are 
paicuherly Use for addressing soetl evalitioniny questions. Stlodapime bees are especially sel for 
WT Soci evolution, since species displiy varying forms of sachil ordination fron solitary 1 eusacial. 
This Study examines WHh Tn nest hebaiviodl ol Bievewia (Evonearete) tridentate, a niative Austertian allodapiic 
boo. This species his fhe lamest Known colony sizes of any allodapioe bee anc exhibits siriking size vuriaiion 
among female nestimtes susgesting thal sockliiy may be regarded as bighly cusoeil, Here we assemble a 
bobavivical cutulosie Lor this species atid show dat although many behaviors are similar la those recorded: (or 
other allidupines. this species differs by the marked presence oPovertaesression displayed in the form of biting. 
Overly dgonshc bebaviodi's have fet been recorded Om ather Australian allodapiaes and lave been recordca 
ony rarely ay offer allodapine (iin, Evenedaye fridentaie appedes Wo differ from ether bizhly eusocil speetes 
where (here s uscally HAle or no aggression bul iste “gentle despatisnn', 


Key Worbs, Jowioie: pide, social beliavinur, allodupie bees. aeeresston 


Introduction 


The atlodapine bees provide opportunities for 
LOIpUvE approwehes to the evolugan af social 
behaviour becouse of the wide range Of social 
Orunsaiion WHE the betwee spectes and genera 
One sinall and endenie Australi subgenus 
Prone. contitins fours species that ringe from 
the predominantly solitary fiveaenra densa 
Rayment (Michener 1965) to the eusecial & 
fedemate (louston (977: Hurst & Setware 1996), 

In most comparative studies of insect social 
evolution (bere is an impheit assumption that 
small coluny sive is assoeiated woth flexible und 
hehaviourally mediated reproductive skew. The 
ailehance of dominance hrerarehies vie phy steal 
awoniSnh is conmsmWercd wi opranmiiye trail (Wilson 
O71), Correspondingly, Jurge colony sizes with 
strom repraduaive skew und) pon-ugonmsrieally 
mintiined hierarehics are usually regarded as move 
derived traits. Wilson (1971) stiggested that less 
Sophisticated Forms of social ordbisation would 
iwolve physreal neelinisms OF control such as 
igeression Wwilhimew colony: bul (hal this ts rephiced by 
“gentle despotism" in more advanced lorms of 
soenilty. [lis alse generally assumed that a high 
level of behavioural specialisation wa more derived 
Ira and that this ean lead to higher levels of volony 
eHlhieney (leanne P98So) However. dhe idea that 
dillerent formas oF sacl orginisutiog cin be 


Seal al Biolomel Seieniss, Flinders University of South 
MSHI GHD Wax STD) Adlethoidle’ So Aviat SCH TD tibial: 
Aa Steen Ge Tider scabies 


dFunged ina sequence of ‘prmnidve to Tadwaneed? 
his been questioned (Kukuk 1995) bul few studies 
have explicitly investigated whether jpronitive’ or 
‘udvaneed! Loris OF soctulity Within tixu correspond 
10 husal or distal positians within phylogenetic trees, 

Eronenre tridentata is ao Austrabir allodapine hee 
thal lives in semi-arid environments. This species hits 
the hirgest Known colony stees of inv allodaprne bee 
and exhibits morphologicul differentiation. between 
putative castes (Houston 1977, Hurst & Selwirs 
1996), Much of the information aboul social 
organisation bas been inferred frou disseetion af 
Hest ovcupants and brief observations of females 
ouside of their nests Clouston 1977, Hurst 
unpub), Tas suspected that this species exhibits 
caste dilferenliation, where large females 
(lermed ‘Miajors') ure queen-like and smatler 
Females ('Minors') act as workers within the 
colonies (Homston 1977: Horst 1996). Llowever: 
wilhih-fest behavioural studies have not heen curried 
Ou) Loouaséss Whether These Iwo trorpls realhy ane 
behaviourally distinet, Colony size and the wsseenition 
between morphology une reproductive shitty sugeest 
thal (his species more closely approaches the bightly 
cusecial fon oF organisation characterishe obapinie. 
micliponine and highly eusocial hahietme hees. than 
uny other allodaping hee. 

This study Mvestigates \witiichest behaviour rn 
observation colonies ob /. tridentata. A repertoire ol 
hehuviours ts presented fiere in the term of i 
bebaytoural catalogue and compared with other 
behavioural studies ob wlodipines. These data well 
Who be used for specitic anulysis of behavioural 
speculation, which wilhappear ing (uti series of 


a0 ¥ STERN & MP SCHWARZ 


poblications. Ty addition, the Wea that morphological 
caste differentiation and linge colony size are 
ussocniled with low levels ob agenisat im colony 
intevration is discussed in relulion lo the social 
ormimisuhion ol A. prredemtaret. 


Materials and Methods 


Stitely vite 

Exonenra tridentart tests were collected fram 
Lake Gilles Conservation Park (13048 bk, 32 54 8) 
loaded in the ori east of Eyre Peninsula. South 
Austrulig. In this aren, 22. aidentde Wests were prin 
oipally ty disused beetle burrows excavated in Aewe fr 
pupyrecarpo Benth, (Western Myall) and Halyerryou 
aleiafoliun (Dest). (Bulloek Bushy Deal branches 
ol both tree species were examined for nest 
enifinces ie. the exit holes made by the original 
beetle oecupalts Trtaet colonies were collected 
daring February 1995, Field collection of nests 
took place when lemperatures were cool (12 C-20) 
Cy. 0 ensure Thiet all Gecupinty were present. 
Onee un entrance hole was locwted, the brunch was 
removed, entrances were blocked With tHssuic paper, 
fhe brinch was placed ina watemrool bay and 
stared qh ai insuhured container with jee for 
Hransport to blinders University. 

At Flinders University the nests were stored ana 
constant Lempertture roon ab approximilely 1 ¢ 
for provessing. Nests were opened using a knife 
cil ull nest aceupants. meludiig brood and: test 
contents sdch us pollen. were transferred to a Perr 
dish. Adults were individually marked) using 
Uambrol!™ and Testors! enamel pauls apphed to 
The thorax and merisond, Bee colonies were then 
Prubsterred to arlifienl observation Wests, 

Arniticial nests were similar in design te thoxe 
described by Selwear? & Overholt (1993) hut were 
imade Ob pine wooed tstead ob balsas Bach dest 
consisted uf a rectingulie peewee of untreated: pine 
woo 7108 20% [Ss mint A gropye was gouged inte 
one longitudinal fee (3 rm dian x 200 mm tent), 
The groove wus smobtiicd out wilh a mend rod ta 
remove any Splintdrs ab woud. A piece of glass. 210 
X Shon, Was placed Mushy suuiinst the groove aid 
secured! al both ends with iipulador gape. A black 
curdbourd cover was placed over (he hiss Wy exehidhe 
Hivht henween observation periods. 

Observalion nests were set up oo subhori4ontal trays 
ina shade house at Flinders Unrversity. One end of the 
shade hotise wis open so (hab bees could. fords 
freely outside, Nest entraiees faced fhe oper end ol 
the shade house, A inuxinian gl four nests wats 
plased on each dray with approstmately TS ern 
herween cael) nest Observalinn nests were list 
placed in the shade house at disk Sy days athe 
colleetion and opening. This ensured that the bees 


hud approximately b2 hours tinue aruhenl nese to 
allow their odours 10 permeate the nest before i wits 
possible for them to leave (the next morning) Sticks 
were haphazardly placed near nests ty wet as visual 
cues for returning bees, 


Behavioural observations 

Ofice Observation Nests Were ser tip Aves were 
alowed to adjust Lo their new enviroment lor one 
week before observations began. Data collection 
ivolved ‘sean’ and ‘focal’ sampling techniques 
(Altman 1974). Sean sampling invalved recording 
the position of cach individual a the observation 
nest. using a S prim seale whong the gliss and wos 
vonducted immediately before and alter focal 
simpling. This was done to determing whether 
certain bees were spending more Gine than oliers 1 
verlain areas of the nest. for example, uew the 
entrance or near the hrood. Focal sumpliny iiwolyed 
2 min observations of cach bee mat nest. Nests aod 
individuals were randomly Selected euch day for 
order of observations. A headband magnifier (4 5,2 
magnification) was used to observe (he behuytour ol 
individuals. All behaviours pertormed ina 2 min 
period for each individual were recorded into a 
voice opened recorder Observations were trie 
sermbed on to data sheets ata later date. These 
behavioural dala were used to eonsiruel te bebay 
foul catalogue ind liter to caumine behavioural 
speciilisanion. 

Behavioural observations look place ia the after 
noon, (1300-1700 hp, when temperatures were = 20 
Cound bees were active Tn total, 1 nests were 
observed with up to lotr bests here Gbserved i any 
one sessian. Table | provides information about 
whieh nests were observed. when Wey Were observed 
ind how miny minutes of observation each bee per 
vest received) In addition, the numbers of bees (hal 
Were present for the ital and Gaal observatin 
Periods are saved. 


Results 


Field-callected Wess 

The contents of nests collected TH February Ln 
are summarised in Table 2. Durtne these simpli 
peniods. colonies used for behavioural observations 
were ceubings brood, dT curly Februiry colanios 
contimed brood Gf all developmental siames. Le, 
owes, larvae, prepupae and pupae By bite February 
fondle bees i tire colonies: haa aliiost ceuacd ces 
hiyin ind brood mostly comprised: Taryae, prepupyie 
und pupa, There was a great deal of vartabion i tie 
number ol adult lernales present ima nesh rine 
Mun T-TR (Pie TS 


BEHAVIOUR IN AN ALLODAPINE BEE ST 


TABLE |. Details for nests of Exoneura tridentata obyerved in this study. 


Nest First Last Total number Total minutes — Initial no. of Final no, of 
observations observations of of Individuals Individuals 
observation observation 
periods per nest per bee per nest 
I 7 Mar: 14 Apr. 15 30 8 a 
6 7 Mat. 14 Apr. 15 30 9 Sa 
y 7 Mat. 14 Apr. 15 30 ) 12h 
12 7 Mar. I4 Apr. 15 30 13 15h 
3 5 Apr. 4 May i) 38 5 5 
4 5 Apr 4 May 19 a8 4 5b 
20 5 Apr 4 May 19 38 4 5¢ 
30 26 Apr. 16 May 2) 40 6 x 
43 26 Apr. 16 May 1) 4) 3 4b 
56 29 Apr. 16 May 20) 40 4 4 


Decreases in the number of indivduals were probably due to death whilst foraging or dispersal to other nests“ Increases 
were due to the addition of newly eclosed bees?, or intruders which swapped nests°. 


Taser 2. Summary af nest contents for colonies of Exoneura tridentata collected in February 1993 from Lake Gilles, 


Soutlt Australie. 


Mean value (+ S.E.) 
for early February 


Nest contents 


Mean value (+S.E.) 
for late February 


(N=24) (N=13) 
Eggs 1.21 (0.57) 0.08 (0.08) 
Larvae 75 (0,03) 0.62 (0,27) 
Prepupue 0.67 (0.28) OAT (O13) 
Mupiae 242 (0,72) 2.23: (0,70) 
Majors L.L7 (O16) 1.23 (34) 
Minors 4.17 (0.83) 4.48 (1.40) 
Males 0.33 (O13) (LA8 (0.21) 


Behavieural repertoire 

In the following section behayiours observed during 
ihe study ure presented as a behavioural catalogue. 
Observed behaytours are classified into four functional 
groups (often interconnected or overlapping): (1) sell 
maintenance behaviours, (1) nest maintenance 
behaviours. (ii) inter-adull behaviours. and (iy) 
aduli-brood interactions, 
StL MAINTENANCE BEHAVIOURS 
INACTIVE 


Bees were recorded us being "inactive" when no 


Number of nests 


other behaviour was being performed, Inactivity 
often occurred within a behavioural sequence. For 
example. a bee could stop grooming, be inactive [or 
same time. and then travel forward in the nest. Bees 
could ether be stitnding ‘upright’ or they could be 
lying ‘upside down' on the floor of the nest. Maeta ef 
af. (1992) included slight movements in their | 
description of a similar behaviour, "Resting" 
However, in this study bees were only recorded as 


20- 


10- 


16-18 


I 2-3 4-6 7-1) A1-15 


Females per nest 


‘a. 1. bistegran of colony sizes (number of females per 
nest) of Aveneure tridentate collected from Lake Giles 
South Australia, Bebruary 1995 


3K 4, STEEN & M,. P. SCIIWARZ 


inuehive When Wey were motionless, veers 
iWdentata spent a large amount OF Lime inactive, 
Since inaciyvily can oecur within and between 
behavioural sequences i) is dilficull ta) show 
Humerigally the wmuunt ob time spent juctive 
hecutise of the way the dite were colleeted, 
Generally, though, We bees were more active Wher 
lemperilures were 220°C and/or when a lorager 
returned, 


SELL GROOMING 

“Croomine” wits observedt frequently, and included 
wy uelivily where the body surface was cleaned 
Sequenves lop clean different areas of the body 
wore similar to those reported: for Braaiseps leiierr 
Cameron and Cerufina spp. (Mactieral 1992), The 
most Common sequences were: (a) head cleaned by 
imtialhy wipta a foreles with the probascis then 
lorcloy wised ta wipe the leneth of the antennae. 
beginning dt the base: foreleg (ain wiped with the 
probosers, followed by the wiprig of the head 
wilt the forelegs, (bh) the metasomi was cleanee! by 
using the Tibial spurs ae the hindlegs to serupe all 
dust/pollen, (&) the thors was cleaned with the mid 
fous (the metasoma and the thorax were often 
groomed al the sine lime wath the different legs) (a) 
ihe wing surfaces were groomed by drigging the 
wins under the metisoma with the bind lees, 
Wiping then between the metasoma and hind legs, 
une then Thiekite then baek inte position, 
Grooming did not oceur as one long uninterrupted 
sequenve as hus been observed Tor Bo hewirt 
(Macta ef af, 1992), Grooming could be brie! or 
last for the whole 2 mi observation period, 


SHICHI BODY SION EMEA S 

This was intermittent behaviours whieh was fie 
Gbserved dung lone bouts ol naetivity, and bebiy- 
OUP Comprised SHANE Movements OF head, body or 
lees which diel nob anvelye any other type ol behav- 
TOU, 


PAWEL LING 

“Teeelling’ tnvalved movin: forwards Gt back sands 
up er down the west for 1-20 en, Bees that were 
travelling Were usdally very detive but the wavelliny: 
speed varied, Travelling forward offen resulted ina 
bee coming inlo contact with others and was usually 
followed by "passing! (see below), 


VURENING 
"Turning was used to desermbe a change of 
direction in the nest. Turning involved cuming the 
body and semersaultina. resulting the bee facing 
the opposite direction. Both Majors and Minors 
uppeared fo turn with equal ease, This behaviour 
vwevirred anywhere in the nese, uolike that in 


Cerainad spp. whieh have ov luring burrow 
enlurgement nea the nest ertynee (Maki en al, 
1992), Thiming ollen ocenrred as parlor a sequence of 
behaviours during interactions between idividiials, 
ie. ih Gould becur daring sequenees which involved 
“nudging”, “passing” Or “uvoidance™ (see below), (fa 
hee upproached but avoided another bee, it mish 
either "Wavel” up Lo the bee, and then back away or il 
might "tare" and “travel” ithe e@yposite deen, 


MECTAR DEHYDRATION 

Individuals were observed flexing and bending the 
proboseis and. although droplets of neeiar would hat 
be seen with at the magnifications used, i wes 
assumed that they were dehydrating eet! as fas 
been observed in other allodapines idler leecdime 
(Michener 1972; Macta ered, (992), Same bees slow 
ly fully extended and retieted the whole pru- 
hoseis without bending if. The proboseis was 
extended and held out lor about 20 see thes 
retracted before being extended again. Some 
individuals spent the whole two mio ubservation 
period perlorming: this behaviour 


NEST ABSENTEEISM 

When individuals were regularly absent (rer the 
Hest Ho owits assumed that they were forage. 
However, i they were absent for more dni 3 obser 
JUUON sesslobs in a cow. He Was assured: (hat they 
were elther dead or had dispersed, Absenieeisi (ur 
forging aclivity) was only observed when tenper- 
atures Were 2 25°C. Foriwers were dentitied when 
they were seen returning to (he pest, Upen return. 
Hg, forrers usiidly worked (herr way dlown 
(he mesh pissing: dnc interucting. wilh) ether job 
Vidiels. often having “bueeal contac!" with vtber 
muds. presumably providing thei with mee- 
tir (see inber-adult behovieurs) Often such wv 
bee Would then leave the nest again amd retien 
filer Poragers were not observed feeding bir 
vue, 


Nest MOMNTENANEL BEHAVIOURS 


GL ARDING 

A bee was recorded iis “suurding" when ifoceupicd 
The position closest lo the nest entrance wilh ils body 
oriented se that its head wats facing away trom the 
entrance, Suelo position atiows the metisomu ta 
block the nest entrance from intruders, as reearded 
for other allodapine bees Bo lewitn (Maeur er al, 
1999), Al miata (Batra er at 1993) aind L. biealor 
(Melia & Schwary 1993), During guarding the hee 
wus tnactive either on ths back or standing upright, i 
a bee was closest fo and facing the vest enivance, it 
Wus mot tecorded os guarding, sinee bees i thes 
postion would offen be we the process ol leaving 


BEHAVIOUR IN AN ALLODAPINE BEB ) 


the Hest, Minors were often seen guarding und in 
done fests, Majors, particulirly ege-layers, were aot 
seen to atid at all, 

Guarding did) hot always oevur near the nest 
cuirance, In some nests the “wuard" was stutioned 
hel ol the wiry down the nest bul was the bee 
closest to the nest entrinee, These guards were 
somedimes seen tn patrol’ the nest fron thitt secuon 
Up to the catrance. This involved the bec rapidly 
“travelling” forward, whilst rapidly aptennating 
(“Inspeetine”) the nest linen before returning to the 
guard position, Tn some nesis Th also appeared thal 
(wo tadividuals would) gud alternately or one ii 
frontal the orien Although there were times when 
More Than cone Hel vidal Was Seen ith The g@etara 
postion, there were individuals who ever 
"ouurded". During the study. no other invertebrates 
Were Observed entering the Hests. Sinve there was my 
interference from other invertebrate predators in the 
CAPUVe SIQATION, guardiig te this study tay ot 
reflect natin! behaviour ob this speectes. 


INSPECTING 

This hebaviour invelved a bee alternately anter- 
nate Obyects. lor example the nest will or 
brood, Epes were frequently antennated fin this 
way, Sometimes bees travelled up and down the 
Hestrospeetine the lumen wall Dirring this beh 
jour bees moved their Heads shehuly aed apiclly 
Hoved antennae, 


MOVING HUBRIS 

Debris in (he vest was moved by passing it under 
the body with the forelegs to the bind legs ther 
pushing backwards with the hind legs or metasorn 
This behaviour was rare (approx, Q.39 oF the 
observation Time) sinve the nests avere in hard, 
Mie-vriammed wood wich required litthe main- 
ftenanes, Debris observed Tm) the nest eluded 
cauynie jie. oecusronally, dead individuals, 
"Moving debris" was not usually observed doless 
lumperutires were = 25°C. 


PAPER ADE TT AE TAMIGUIES 


WOTPARC 

“Avondinee', a combination OF other behaviours, 
iInvelved Gne dodividdal daawelling: towards another 
mdividual dnd “Untcueiing either the metusonn 
or lace of thatindividdal and thea suddenly bucking 
away oe Tinie an tavellinges Ti the opposite 
alirection, 


ANTE AAT CON TACT 

"Antennal contaet® accompanied most itereadul 
behaviours, When wt Trdividiael Game tn te contact 
With another mndividtial a criti “antermiited" the 


ofher's metasona or faee. HW tidiyidtids were hice 
losfaee the two individuals tapped each other's 
antennae, 


PASSING 

“Passing” is the eschanve of positions by nest 
mates. Passing oeeurred when individuals were 
either ficing euch other or the "passer" was lieing 
the tnetasonw of the individual she intended to piss. 
In cach cause, iudividttals ooented (hentselves ven 
ler-lo-venter, essentially walking over each other A 
pass was either simple or complex. “Simple pass- 
ing” Gnvelved the smouth exchange el) positians, 
with individnals usually flattening thei bodies 
ugainpt The nest wall, “Comples. passing” involved 
one individual biting al anether medividualls buaty 
parts. und/or strupeling and grasping cueh other 
with the legs. Either one or both individials would 
Hits, Sometiones one midiyidial would bile the othe 
on the ventral sig between the ietusonit ane the 
(horas. near the articuluion between the tochunte 
und the thors. Passing somehimes mvalved briel 
"buveal camtuce” between the two individials. 
Although it Was often difficult to determine clearly 
whether buecal contiet tad actually geeurred. Ue 
Wan not always cusy to Uistinuiish between the 
passer adind the "passed", except when onde was 
intbially stationary and doarher wes teevelli pe 
HEICCAL CONTAC EP 

lnelivaduiils were often observed to toueh cuch 
other's open mundibles with their ewrn épen 
aindibles: this was termed "buccal cantict. When 
Hhdividuadls were Involved dn such intenichons. one 
individual was standing upright und the other was 
positioned: upside down, Tnehvidtals dibse engaieed tn 
briel hoceal contact during passing. During 
Approximately 54 oF biiweal contact iileractions. 
neekiur flow between the mouth purty of individuals 
Was Observed ancl individuals were observed placing 
thew probosers bebween the mandibles oF another 
individual, Protflering of lobules of nectar (Meloa & 
Sehiwirs 1999). wits nolohperved in by drideniale. 


UDO Ne! 

"Sudging"” involved one individual using Us fice 
to nudge or hut) the metisoma or face of another 
individual, The hee that "nudged" was usually 
upright. Nudging usually resulted in one af the 
Following: 

a) The nudged individual (ined and (he nudser 
retreated, which sometimes involved (he nudged 
hee opening iis mandibles. 

bh) UP nudged frewr behind. the bee beige gudved 
would sometiines position its antennmie laterully 
(oul tothe side). then 7 nudecd again ib mnght open 
is mandibles, ‘This eventially resulted in the hee 


a Z. STEEN & M, P.SCIIWARA 


juming, jnvesigaling the "nudger’, and thea simple 
or complex passing and/or buceal contact 

¢) The nudged or the nudger pissing and "bring" 
uel oer, 


MANDIBULATISG 

Mandibulating, Le. the opening und closing of the 
mandibles not associated with eating, appeared i 
occur before biting enconnters. I some cases 1 
Uppeared that mandibulating was a signal thal one 
individual was reject an approuch from another 
individual, Por example “Ab approached ‘Bt, ‘A’ 
nudeed "BY "BR then opened mandibles, “Ab (hen 
retreated. “Biting” encounters someuimes followed, 
Similarly. an imdividual was nudged rom behind 
it sometimes opened its mandibles and/or turned aed 
faved the nudger often opening the mandibles wan 
In wddition, Matlening of the antennie laterally often 
ocenrred during mandibulating, This sometimes 


oevurred when midividiails came face to face or il 


one was nudged from behind, 


BEEING 
In this Siidy aggressive cheounters were Observed 
for oh. weldentata. These 


the thor, around the coxue ung metusoma. Olen 
When one individdal tried te escape from such in 
eheounter the other bee would pull it buek osing ts 
forelegs. "Bilin encounters were olten vomples, 
For exwihple. (A> used its fie fe nudge "B'S lave. 
Then one or hath bees opened the mandibles and a 
complicated pass followed. Whilst the bees were 
venien oO venter and strigeling (holding cach other 
with leas; one would hite the other ou the ventral 
side ol the thorwa, After a sttiiggle the bitten bee 
wis oflen observed on dis hack while the biter hele 
the other bee's antennae in is mandibles, ia “ug 
Whawar" Gheounter This tug-of-war could: last far 
120 see, botlowiie a fie-olewyr encounter the 
Tdividiial whtel bad daiiaited (he pass (he bition) 
SOMetiMmes ullenipledl (er puss ius and Ofer a 
siiiple pass would follow, 


ADRUEIE BROOD IN TR ACTIONS 


PY ANINATION OF WROOH 

Excunation of brood was acceanplished with ve 
ailennae, aid, faa desser exten, ie ieuth putts 
(open and clostime mimdibles on che bread) 
Individuals Lippe pupae. larvae or ees. WIE cael 
HAW 


NEEDING HROOD 

Brood Were sainetimes nudged before (hey were 
moved, This belivioie dich dot fesull, Hewever, iy 
jhe bro ipprestibly chanutas: position 


invelved bituay ab 
mandibles, auitennae, peek, legs, the ventral side of 


MOVING: BROOD 

Older brood (ate instar larvae, prepupae and 
pupae) were usually moved in a way similar to the 
way debris was moved in the nest. In A. iridentate. 
simik to EL biedlor (PS. VMurst pers. comm, 1995). 
the bee initiiy held the brood with the fore (arsi 
then passed them under the body and pushed ther 
backwards using the hind legs. Repositioning of 
brood occurred allen within the nests of A irideniua. 
Sometines a bee would move edeh pupa until it 
reached the end of the nest, then it would move then 
all back again: seeonds later another individual 
sometimes did the sane thing. Some Minors which 
consistently stayed near the broad were oflen 
observed performing this behaviour, fh addition, 
hees sometimes simply handled the pupae with the 
fore lows bur did pot aetially reposition them. 


GROOMING BROOD 

Bees ovcasromuly extended the proboseis to the 
hrood or bit gently al the brood with thei 
mandibles: such bebaviour wes eateporised as 
“grooming broad’. This behaviour was rarely 
observed. Greating jay have occurred during 
movirns or with Tiindlings but i owas difficult to 
observe the finer movernents of such behaviour 
becuuse of the speed oF movement of the prohoseis 
and the lirited magnification, 


OVIPOSTTION 

When "Ovipositing”. the female oriented herself so 
that the head pormted towards the nest entrance 
During eve laying hees were observed fone of thice 
positions: vennal sartice lyeing Upwards, dorsal 
Surface facing upwards und lateral surface facie 
upwards, Prior ound during "Oviposition” the sting 
Was Gaternled. Onee an ene had been deposited qa 
the floae or the nest, Ure bee retracted the sai 
Approximately ba punt passed betore the fenyale 
iuirovel around and inspected (We eee willy (he aalennie. 

Oviposition @ecurred close La the mest end (5 
and was vibserveel ford Majors and | Miner ¢§ 
sepacdte colonies) Tadividhials Tok approx inidtely 
SO Wii fo discharpe un cep Plowever oie Majer 
look F8 ni to Tay at eas. 


Discussion 


Beliviouwr has prev daily heen stodied i denial far 
RB. lrewinli (Maeta er iil. 1999), A ote, AL kallawe 
(Baumer af (99%), AMlodapie exelent Strand) 
(Mason lOX8)} dind 2 bleelor iMehine & Seliweure 
1093), Bvonenme (rrdentiata wenerallyospencdd iy Lite 
HMOURL OT Ce Tachive, SHH Teather Rees CN aot y 
eral (902: Batra ela (998), Activity temled lor pre 
wicuter ua dlays wheo the temperahines were uhove 


BEHAVIOUR IN ANS ALL ODAPISE BEE ol 


28 C. Sinihily, wher the temperdture was warmer 
bees tended to forge more und, especially ater 
return of a forager to the mest. gener) activity 
appeared lo increase. 

Exon iidednita was bat dbserved to exhibit 
(he types of fest maintenance behaviours found in 
other allodapines. probably due to the hard nature of 
he nest substrate. Most allodapines excivate (here 
OWE NOSES I pithy Sudstrite imiterial, whereas &. 
Iridentatd do pot Although observation nests 
provided no opportunity for nest walls to decay 
during the course of (he sludy, natural nests are 
WAG unbRely to require fepines to the nese wall or 
entre, Since Lhey also ocetir in line grammed wood. 
fhe contrasts With ak, biceley which pertoriis 
varius nest miintenanee aetivities suely as 
Clearing aod famping (removing loose material 
(rom the nest wall and shape gest lumen) 
extending the nest lumen (excuvaling rear of the 
burrow). collar Construction (ping Wood imto a 
collar near nest entrance) und remoying debris 
dwood strimds. Zienewne rridenta may exhibit 
HOSE IUTIeNUNee SeOVTes Lod prenter degree when 
Hew nests ane lotinded and tere is a need to remove 
brine Tet behind by been larva, 

This study is the first lo deserbe eee Tayroy ip at 
Aagnenre species. Bee laying was only observed 
duting, (he diy althotirh iarkiy idsi have oeeurred 
at feht (obserwtions Were only tnade dupe the 
iliy) Meo living wieyscoilier bo that deseribed far B 
mrte (Batra oral WOD) un Bde (Maer et 
1907). However, bwo of the three 2, tridmire 
majors That were observed OVIPOST Te were runely or 
never seen euardun. Phe thin mijor was seen bo 
sajurd bol she was sully S em fron the base of Hie 
nestamd fot pear the entrance. This differs from &, 
Jeovien (Maen et al O07) aad bo dicaler 
(Howerdoory de Selivare P9988: Buller atin pless) 
where reproductive glominants wre guards, Exe 
faving in his species appears da be uw yery slow 
process compared wath other bees (G8 see. Bo tear 
tH) (Maeta ef al 1992). in terms Of both the Time 
Taken to deposd af ems and (he freqdeney ob eee 
laying, Que tenile, i particuhae spent 4s min 
depositing aces whieh tray have heen related to 
the [acl that the femperture was low that dia ¢< 
WC). and bees were generally less active al 
lower temperadiunes. However, these ohservariins 
did pot cover the perrod of imaasrtal eee prduction 
and stow be treated with edution. 


Aggressive belityiour bas net been reported far 


olber allodipine bees except rarely henveon fH 
wiylitanel its Social parasite B halfage (Batti ef af 
(W984) and jnlrequently for A. eveforuee wed B. freee 
(Mason L988). Phe ayonistic behaviour described 
for These speeies rnuinly coosisned of nudging, biting 
Ol legs amd bodies ind blocking passage. hut alse 


included singing (Baten er vf. (993. Mason TOSR) 
Agoniste behaviour between a hast and fs parasite 
is nol uncommon and Often results i sither host G1 
parasile being removed from the nest (Batra el af. 
19034), Ageressive ieteractions are alse found in 
social Species OF the bee tibes Halictini and 
Nylocopiny (Breed et ah 1978) Michener 1990). 
However, 2. frfdeniiir Was offen observed ti 
cngave in gegressive encnunlers which inyolycu a 
prea dewl of bidne ound strugeling, with seme 
eneouniers Becoming quite savupe, Sach cneounters 
were offen preceded by nudging ind folliwed by 
passing. The miandibilating that ecard same- 
limes. eiiber prior lo er in response to nudging iil 
Hite. might idso be aggressive oy mature Cane Xe 
Michener (1983) found that some 2eeavewre: spp. 
produce iratuis whieh cligit Vigeroos geoming 
responses in predatory als, Batra ed af, (b984) 
deserthed oo mundihulating  durrog  aeeression 
heiween Bo ate und its sucnil parasite BL keviiges 
wil sugeested (hat mandibilar seerenons were 
involved, Hinay therefore be suggested that when & 
Tridemate mandibulate al eweh other, (ley alse 
release chemical seervetions which rity be weorishic 
or relay information wheut donee: stctus. 

The agonistie behaviours observed mL. ident 
sugeest Wal dominance hierirehies nny he present 
wilbin colonies, Te appears (hat some ijdividddls 
engave im cern oypes af behayinur whieh could he 
IMerpreted as assertion oF duminunce. Bees that are 
offer nudged or bitten and those that exhibit avonlance 
behaviour muy have more subordiiate roles in the 
nest. Dilferences in the way ined;viduals respon be 
other individuals in terms of these behaviours pay 
he related to dominance (he. when Some individuals 
wre nudwed (hey cheaee Ha simple: pass. whercus 
when other individuals ure nudged and/or bitben they 
Copuge iw complicated puss), Brothers & Michener 
(1974 found that ‘qneens’ ob Lenvreadiassan 
sephyran) were the maximal nudgers in the colony 
They steeestecdh (hal nodeing belie indicates 
dominance sinilar to hal observed in other prin 
Hively cusuen wasps amd bees Brothers & 
Michener (197-4) experimentally showed toe 7. 
séphyriin. that mudenig by the quicen pris ae role in 
the division of Libour anion the workers hy imbibing 
evar development, 

During this study gourding behastorr was not We 
sete us thucobserved in field studies ol Ey frfefeyitetia, 
Pewith the abdomen curled and used ta block the 
entrance from predidors suchaits anes UP Hirse unpub.) 
This may be rekited ta the Piet that there savas no 
predation pressure in the shade house environment 
unlike studies of AL Pieefar conducted in shide 
houses where ants were at problem (Bulls Furst") 
However females that were guardine were always 
facing the betitom of the nest which sugeests du 


ot 4 STUEN & M, PSCTIWARA, 


they were dna posmon to block the nest a the need 
arose. 

Trophalladis as altruistic behaviour, foragers 
engage in energetically cosdy and risky behaviour to 
vhiiin food whieh they reloigiuish to others. 
Trophallaxis isamportant in (he soci! organisation 
OF iiny seeial insects (Wilson 1971), Thy allodapines 
(here nay be differences m the way in whieh 
Wophallaais is performed. Evaneura brealer Wave 


been Observed te engage i solicition behaviour 


before rophallasis oeeurs (Melia & Schwarg 1993), 
Solieurion dvelved: individuals rapidly: stroking 
cach other's antennae prior to buecal eontaer, 
Trophatlasis in 2. bieular can adso invalve one 
iidividiel proflering a slobuile oF liquid: ty another 
(Melia & Seliware 1993). Peotfering of globules 
was not obperved in“. tridentera und UP solicitation 
occurred, ib was Loo fast to he identiticed, However, 1 
is Hikeby that midividtals which engaged i "huecul 
conuiel where heel flow was observed, were 
frequently engaging i trophylhasis, Trophiallaxis 
Allows tenes to feed without leaving the nest. The 
presence Of Wrophullaxis in AL trideviie therefore 
allows behavioncal specialisation where only some 
Ol the females huve to lorige and other females ean 
perform other duties in the nest. 


Axonenura tridenieio exhibits a similar repertoire of 


behaviours loother allodapines (Maeta er ef, 1992, 
Bawa oe wh 193; Melna & Schwarze 1993), 
Behaviours recorded in (his study, ineludiie adult 
Udult fiteractions and adull-brood inleractions, ure 


TBC. Nr 1994) Puseehility ia Neatidand: populition 
ofan Astrid Atlodiuping bee, Eveneure bicolor Smith 
(Apidie. Xylocopime), BSe Cons) thesia, Plindens 
University of South Australia (unpub). 

Hers, POS (1004) Repradtictive hiamarehies by 
on Austealian ATodapine bee. Evaneiiid Bical 
Sith (Anthaphenrtdae, Sylogapaned, BSe (tons 
thesis, La’Probe University. (ipl), 


HIP Simihie to those found for other species, sugeestiny 
thal such behaviours are likely lo be ancestral an 
thal development of novel behavioural elements is 
Hot Heeessury Tor soci) oruinisation to evolve tron 
small family groups to large groups wilh morphological 
ifferennation among colony members, 

However, unlike other allodapines, 4. piedenraore 
esbibirs frequent und overt ugonistie behaviours 
among nest mates. Such agonistic behaviour tas 
offen been associated wall more primitively soci 
speeres, According la Wilson's (1971) erileria, & 
wridemiata can be lussed as highly eusocial because 
there female morphalogiea! dimorphisin associated 
with reproductive division Of labour, Pherelore, &. 
frdeweta doesn't cortorn to Wilson's (197 |) sugeestion 
hith averession within a colony can be replied by 
"gentle despotism’ as socklity involves larger eroup 
sive und requires a greater degree of integration, 
Most other highly eisociit species display distinct 
morphs WAH are directly associted with diserele 
behavioural costes. invelving Minimal or noageression 
Considering the presenee OF aeyressive iilenielions 
with) 2. trident colonies, if would seeny (hut 
incrensedvolony size ane the developracnl af mor 
Polo ditfercotiaton among colony mernhers 
need not he acconmpanied by decreased levels ol 
over iitra-eolony dye ression, 


Acknowledgments 


We would like to thank friends. and laboritory 
Inembers who assisted with field work and N. Bull, 
S. Reyes. PB Tuest, J. Bird aid bwo anonyitous 
relerees foriuviee on the manuseripl. This researeh 
was puirtially finded by wats from the Ansteatian 
Research Couneil to M, POS. Field work was curricd 
oul WH permission from (he South) Australian 
Depurtimeal af Environment and Natural Resources. 
Per Ney, Q23256-03. issued to M.S, 


Reterenves 


Alona fe CIM 74 Observational stuely al lelavieue 
sapling methoels. Bedaeine 49, 227267. 

Baton, So Wo To Sakagiiin S22. & Moen Yo boos) 
Relwivinwn al the Tndicn alladapine bec. retina 
Adie. i SeTUE psi dn he festh uh AD iste 
(Hy menoptenn: Anthoptiaridieh 2 Neascas brrtenid Sen 
Of, 345-460. 

Broce M1. Silverman. I ML ode hell, Wo (17s) 
Aponistiv Behwiour soci infericuons, aml 
hehavioueal specialisation im a primitively eusocil bee 
frsecten Sactaia, Cavis, IS, 351-364, 

brothers, Do & Michenern ©. 1D. (1994) Ttentetions ia 
colonies al perontively soc bees TH Ethalowy al 
division owl hike ine Lentiiglossi se plivrane 
(ymehopteri Halide) 2 Comp. Physi a |e 
Jan. 


Bick Nd. Minti. ACC. NORTMAIST, Yoo ubvEsIVN. B 
Lo & Souwary, M. PO (in press) Caving your 
duliehters Wie esve he quenthing reproductive 
dominance in a oprimilively wackil bee, fre. Ney, 
Soe Set (Serres B), 

Combo Hook Micueare, cof) (1983) Chvanistey aod 
Hichon of nanidibular ehind praduets af bees of the 
Lenus Laenenra (HY ienopterd: Anthaphoridieh A. Chen, 
Evol, 8 (825-1531, 

Tlonbsbeurn, Kd SonWwane. MT (it press) Chardin 
Spee saion jn pre-repramuenve Colonies Pine etl bochap yi ie 
bee Kavtenre bicalvn dal boul eval WW 

Hrnsres oT, F197 7) Nesting: Woloy of fhtec alocappite: 
hoes WE THE subgenis Crema Michener Pins, RL Sey 
SN day TOE G1 ba 


BEHAVIOUR IN AN ALLODAPINE BEE 63 


Hurst, P. S. & SCHWARZ, M. P. (1996) Morphological 
differences among females of the eusocial allodapine 
bee, Exoneura trideniata (Hymenoptera: Apidae). 


Proceedings of the XX International Congress of 


Entomology, Firenze Italy 25-31 August, p. 413. 

JEANNE, R. L. (1986) The evolution of the organisation of 
work in social insects. Monitore zool. Ital. 20, 119-133. 

Maeta, Y., SAKAGAMI, S. F. & MICHENER, C. D. (1992) 
Laboratory studies on the behaviour and colony 
structure of Braunsapis hewitti, a Xylocopine bee from 
Taiwan (Hymenoptera: Anthophoridae). Univ. Kansas 
Sci. Bull. 54, 289-333. 

Mason, C. A (1988) Division of labour and adult interactions 
in eusocial colonies of two allodapine bee species 
(Hymenoptera: Anthophoridae). J. Kansas Entomol. Soc. 
61, 477-491. 

MELNA, P. A. & SCHWARZ, M. P. (1993) Behavioural 
specialisation in pre-reproductive colonies of the allodapine 
bee Exoneura bicolor (Hymenoptera: Anthophoridae). 
Insect. Soc. 38, 1-18. 


MICHENER, C. D. (1965) The life cycle and social organisation 
of bees of the genus Exoneura and their parasite, /nquilina. 
Univ. Kansas Sci. Bull. 46, 317-358. 

(1972) Activities within artificial nests of an 
allodapine bee. J. Kansas Entomol. Soc. 45, 263-268. 
——— (1974) "The social behaviour of bees: a 
comparative study" (The Belknap Press of Harvard 

University Press, Cambridge, Massachusetts). 

(1975) A taxonomic study of African allodapine 
bees. Bull. Amer. Mus. Nat. Hist. 155, 67-240. 

SCHWARZ, M. P. (1994) Female biased sex ratios ina 
facultatively social bee and their implications for social 
evolution. Evolution 48, 1684-1697. 

& OVERHOLT, L. A. (1993) Methods for rearing 
allodapine bees in artificial nests (Hymenoptera: 
Anthophoridae). Aust. ent. Soc. 32, 357-363. 

WILSON, E. O. (1971) "The insect societies" (The Belknap 
Press of Harvard University Press, Cambridge, 
Massachusetts). 


FIELD ECOLOGY AND BEHAVIOUR OF THE EGG PARASITOID 
TRISSOLCUS BASALIS (WOLLASTON) 
(HYMENOPTERA: SCELIONIDAE) 


By S. A. FIELD*, M. A. KELLER*® & A. D. AUSTIN® 


Summary 


Field, S. A., Keller, M. A. & Austin, A. D. Field ecology and behaviour of the egg 
parasitoid Trissolcus basalis (Wollaston) (Hymenoptera: Scelionidae). Trans. R. Soc. 
S. Aust. (1998). 122(2), 65-71, 29 May, 1998. 

The ecology and behaviour of Trissolcus basalis (Wollaston), a parasitoid of the eggs 
of the horehound bug Agonoscelis rutila (F.) and numerous other species of 
pentatomid bug, were studied in the field over two years near Adelaide, South 
Australia. The adult bug population declined sharply early in summer due to the 
combined effects of senescence of host plants, egg predation and parasitism by T. 
basalis and a sympatric species, Trissolcus ogyges (Noble). Hyperparasitoids of T. 
basalis were recorded for the first time in South Australia. 

Key Words: Trissolcus basalis, Scelionidae, egg parasitoid, horehound bug, 
Agonoscelis rutila, egg masses, defensive behaviour. 


Treailvdetionts af.the Bowl socrety of S Mest C2998), 122¢2), OS-71 O35 


FIELD ECOLOGY AND BEHAVIOUR OF THE EGG PARASITOID TRISSOLCUS 
BASALIS (WOLLASTON) (HYMENOPTERA; SCELIONIDAE) 


by S.A. Finep*®, M.A, KELLER® & A. D. AUSTIN ® 


Summary 


Rib boS AL KELL ML Aw ADs in, AOD. Meld ecology and behaviourof the euy parasitoid Disses Masalis 
(Wollaston) (Hymenoptera Scehontlaes 7s. A, See, S. Aust. (YOK). 12242). 65-71. 29 May. 1998, 

The veylovy and belay of Frissadens baseless (Wollaston, i pairisitaid af the eggs af (he horehound buw 
Agenoscelts ratily (VO aid numerous other species oF pentatamid bug, were studied ithe Held aver hwo years 
ned Adehtide. South Mustialia. The adult bus population declined sharply curly in summer due to (he Combined 
GHects of senescence of host phints cae prechition and parasitism by 7 bawafis and a sympatric species. 
Trisyolcny peyges (Noble). Ayperparisitolds of 2 basis were reconded for the tiest tine in South Auseratia, 
Compennon among femule purusitonds for ueeess io host age masses differed widely between the Gwe years 
(sume season). and lemiles displived uduptitions ( earpatition, They patrolled host cae lasses where alone 
and defended then ugeressively in divect coests with conspeciics. These ohsenwitions rejaforye previous 
liboialory Work ond suaoest hither avenics al peseueh on the behavipural sinilegies used by f. brasalis during 


Helerce al cue miisnins 


Ky Wovon: Cielo busulis. Scelionidie. eee parasitoid, horehound bug, Agonaseedis eunila, eo Wiisses. 


Uelensive behaviaur 


Introduction 


frissoleuy bavalix (Wollaston) is uo solitary 
parasitoid of fhe wees of the itrodueed preen 
yeectuble bus, Nesare viridila (Loy (Hemiptera: 
Pentitomidae), dnd aw number of other pentatornied 
species (Cimber 1964), ineludim Lhe qurive 
horehound bug. Avancayceliy come (EL, Sinee its first 
WMpertation tito Austria in 1933 (Noble 1937), 7 
basaliy has heen released a number of times (Clarke 
ISM) Due lo is perceives! importince as a 
biocontrol avent worldwide. many aspects af its 
hinlogy trave been documented (e.g. Wilson L96t, 
Cumber 164: Powell & Shepard 1982: Bin ey af. 
19s6; Volkol & Colazza 192: Matiaeci ef al 
1904), 

Although the field eculogy of 7 haselis in 
Australi is best Known Hon its assocnition with WV, 
Write We Turner 1983, Clirke 1990), its biolagy 
when parusitising A. futile on the iitrodueed weed 
Horehound, Merrubriun vulgare (yh has also been 
Tivestimsited in view of tts potential for maintain 
purus numbers in cropping areas (Kelly |987) 
When feeding on horehound. reprosuetive maturity 


of AL varity is dependeat pow the availabiliy of 


Hlowers, and so both hust and parasitoid populavon 


dynamics ire closely linked to Seasonal eyeles of 


plant growl, Although activity and populion peaks 
Ol both host and purasitonl comeide with the major 
srowth phase of (he plant in spring and early 
sumer neither species aippears to unter dhapause 


Departinit al Crap Protection. Waite Coonyins Pho Lirtiversityot 
Adelie VO Cer Csengiud Si Nest SClie, 


over winter and a high rate ab parasitism (> 70% ) is 
maimained throughout the year (Kelly }987). 

Due to the smallsize and rapid movement of 7 
hevelis. field observations are difficult and dati have 
only been collected foi) one study on host searching 
under semi-field conditions (Turner 1983) 
Oviposition behaviour, exploiiwion of host egy 
Intsses (= pulches) und competition have not been 
studied in the field. This paper reports fell data on 
the ceology and behaviour of 7) basalis parasitising 
Ac ruil, Us most common hosr un the Adeliide 
region OF South Australia (Pig. 1). In spring and 
summer OF 1994-5 und 1995.6, date were collected 
on the seasonal Tuetations of fost plant and host 
populations. sources OF host mortality and behaviour 
Of parasitoids as (hey exploited, and competed tur. 
masses of lost cees. The purpose of this work was 
ulso to provide the Fetundkition Tor more detaded 
laboratory-based stidies ol patel exploitation and 
defence (Micld ef af. 1907). 


Materials snd Methods 


Hast plants anel tiases 

Data on the ecolowy and behaviour al 7 basalis i 
(he held were collected trom late October to late 
March in 1994-5 ind 1995-6 in the Brownhill Creek 
Conservation Park, in the Adelaide foethitls, 
Sampling sites were selected by taking Lwo 500m 
Iradsects dloug candor direchions through a patch ol 
horchound, The transcets were divided inte 10) 
intervals of equal length and a random paint was 
taken along cach fferval, The nearest horchound 
pling, or discrete cluster of plants. to cach of these 
porns was marked ws a sampling site, [stems veased 


fy S.A, PIELD.M, A. KELLER & A.B, AUSTIN 


Competing agq parasitoid: 


Thssaleus ogyges 


yb ey *S : 
*! Host plant 
14 Horehound 4 
Marru brtum vulgare 


4 ruiiis.eaa mess 
{hest patch) —— 


‘Alternative host plant: 
Salvaton Jane. Ecthum plantagnaeurnt 


| 


Vis. 1. Surmoiary of the natura history of the A. radile 


to show any green foliage during sampling. the 
marker was removed and replaved on the nearest 
stem bearing foliage. When no phints with eight or 
inore stems With green folipge remuined, sampling 
was discontinued, On 21 days between 26 October 
and 2) Mareh 1994-5. dala on host plant and host 
population, and parasitoid behaviour were collected, 
The numbers of open towers on stems were 
recorded and ai ides of the A. Afi population was 
obtained by counting the total number of adults on 
all simple stems at LO.00 am. 


Parustioid egy lee 

The nuinbers of eges earried by female parasitoids 
(n= 31) were assessed by collecting wild 7) fetsetis 
females on seven days between 30 November and 20 
December in 1994, and dissecting them in the 
laboratory, In addition, exe maturation under 
laboratory conditions was studied by determining 
parasitoid egg load when females were between one 
and 10 do old. Prior to disseetion, wasps were held 
individually in vials supplied with honey solution for 
L-10 do at 25° CL without ovipositing, before being 
frozen at -60°C, The metasoma of individual wasps 
was removed and dissected ina drop of water on a 
eavily slide and the number of eves in the ovaries 
wournted, 


Behavioral observations 

In both years, behavioural observations on 
parusiloid oviposition behaviour, patch exploitation 
und competion were made. Host patches were 
created hy glueing O-1 d-old ege mitsses of 
laboratory-reared AL vad, each containing between 


Eqgq parasitoid wasp: 
Trissoleus basalis 
— 


Hyperparasitoid wasy: 
Acres es sp. 


we 


Main host insect: 
Horehound bug, Agonoscelis rutila 


‘Natural! host insect, 


Green vegelable bug, Nezara virdula \ 
Alternative hostinsccts | ~-~~—__ 
Cermatuius nasal 
Oeshalla solinellanterg: 


L leiscilis system used in this study (see text for deserption) 


12 and 24 eggs, on to small squares of green 
cardboard and stapling them to leaves on randomly 
chosen sample stems, Patches were laid ont belween 
O.O0 um. and 2.30 pm. and the oumber of female 
wasps on cach ege mass was recorded every 30 min 
until O30 pam. tn 1994, between one and 12 exp 
masses were observed on each of five ditys between 
26 November and 12 December. In 1995, 16 exe 
asses Were Observed on cach of eight days between 
& November and 26 December, An index of daily 
competition for eye masses was obtained by taking 
ihe maximum number of wasps observed in any one 
sumple during the diy for each exe mass and 
calculating (he mean aeross all egg masses, To 
facilitate Comparison between data sets for the 
dillerent years, hese means were taken it (he period 
2.30 p.m. to 6.30 pam. as some data from 1994 were 
collected only during these times of day, To compare 
rates of disvovery of egg miusses, Kaphin-Merer 
vstintes Of survivor functions (Haccou & Meelis 
1994) lor the time until discovery of egg masses 
were caleuhued for data pooled within seasons, ‘The 
survivor functions plat the cumulative proportion of 
weg masses discovered its a function of time, and this 
provide an estimate of the instantaneous rate ol 
discovery of cap masses. 

To observe pateh exploitation and) defence 
behaviour in detiil, pitch visits by single wasps (= 
4). amd by pairs of wasps (n= 6) to randomly 
selectad artificial patches were videotaped und 
converted to behaviour sequenee records in the 
laboratory using a TRS-3O Model 100) portable 
computer programmed with event recording 
software (The Observer, Noldus Information 


BEEAVIOU RAL ECOLOGY OF TRISSOLCUS BASALIS 


Technology Wageningen The Netherlands). 
Behaviour was divided into categories representing 
host @aimination, oviposiuon. puleb-leaving and, for 
pales of fenmes.agonistie behaviour (Field!: Field in 
press) When patch contests between two femules 
one individual (he resident’) usually 
estiblishes dominance and aggressively exclides the 
other (ihe intruder) (Watson L961, Pield'). The 
Mttuder then waits nearby and periodically returps to 
(he cue mass fo allempl further oviposition, Where 
bout deneth simple sizes pernrilted (n> 20), intruder 
‘retreat’ behaviour (detined ay the time between 
being driven away Hom the ege Mass and returning) 
Wils tested Tor abrupt chinges: (i bout lengh using a 
HON pPUVMNetne multiple change point lest (Haceou 
& Mechs 1994). Where changes were sinifeunt at 
the udjusted levels saweested by Haccou & Meelis 
(1994), they are usteated with cumulative bout 
fongth plots. 


Mec, 


Results 


Hast pitts, hosts, porasttords aned liyperparasitoiads 
I) 1994 ink 1995, host plants und host insect 
populations underwent marked HMuetuations. 
Natibers of lowers peaked in late November, and 
therwuller declined steadily Wich mid January (Pie. 
2) when all plants showed very Jide or ne green 
yerehition und no flowers or leaves. Popukiion 
counts of A. rita deereased in’ parallel wath: the 
deeline in plant quality, stabilising (ic low devels in 
mid—Kinuary (Pig, 2) Addit A, cidi/a also appeared to 
be susceplible lo high femperuiures, as many died 
during dhol. windy spell carly in December. The lirst 
Hyiiphs appeared at this time, indicwting the 
vibergence OF The first weneration of the season, 
Numbers of adull ad, rafiiae over summer remained 
nich lower thin the peaks observed in sprog, 

Weeds other than horehound. in particulir 
Salyvaiion Jane, Mehiane planteaginiia (L.) were alse 
abundant at the fieldbsite in (995-6, and were utilised 
for feeding ane reproduction by A. rutile (Fig 1), 
Adults were observed leeding. mating and eying 
eyes On 7 plieeinesen, allhounl ip wits uitelear 
whether pymphs were able to complete development 
solely on this phint. 7rixsoleus basealiv were also 
vbserved foraging on A. plantergineuin plants, 

Apart fram 4 puffle. other pentatomisdts 
accusioially observed ou horchound at the study site 
were N.oniridite, Cermeatulis neasaley (Westwood) 
uml Gechalia Schieber (Cuérin-Méneville) 
(Fig 1) alhof whieh have previously been recorded 
vs hosts Tor 7 havelis. Rees Ob ON, viridider cid C. 


I Bae S.A CU) Patel eQphoruiid cmt dletemeas (ip the exe 
Pibiitonl Frases Hii Wolken Up iteepiene 
Sev Honthue) MAD Hiisis. Phe Criversity al Adehiishe Canputs 


Total bugs/80 stems 


o 


— 


—— Adult bugs 
= Flowerheads 


Mean no, flowerheads/stem 


1) OS Fem A 7 


Vig. 2. Numbers of a raila adults sind mean jurbers ef 
Nowerheads (47 SD) on horchound sfetos in 1994-95, 


10 


it) 2 


6 4 
Age (days) 


Fin. 4 Mean number of eaes (+1 SD) ii the ovaies at 
lomitle 7 baxalis trom [10d after enemenee Nunihers 
below error hues indieale sample sivas 


nasdliy were Nol seen bul those of CL wehaedenbury 
were colleetud and both To beaselis and Trisseleres 
avyues (Noble) were reared from them (Fig. 1) 
Trivsoleuy oavges did not complete vevelupment i 
A. ridley eves, so O. selnellenbergd appears to have 
been its min hostal the site. Triwoleuws ovy ees could 
be distinguished from 7 easaliy in the Held and 
laboratory by @ distinet difference in host marking 
behaviour, Ruther than dragging the ovipoesitor 
smoothly ever the hosteve ia igure 87 mation as 
in TD) basaliy (Wilson 1961, Weber etal L996), 7 
egvges Woved the ovipostlor horizontally aeross the 
oge With a “bouncy, jagged” motion, Females 
defended eye misses similarly lo 2 bavelis and 
Hilerspeciie vontests were observed. bul nor 
recorded in detail As ne objective of This study wits 
fy gauge the overall levels oF competion far aveess 
lo hosts among 7. hawaliv, the data presented below 
inglude Observations in which J auvees was alse 
present Tp 1994, The proportion of loti observiligns 
in Which 7 veyves occurred was nol recorded but in 
JOYS TL was upproximilely |O%, 

In addition to the primary parisituieds. females of 
the hyperparasitoid devevdiseides sp. (Ciraule & 
Dodd) — (Hymenopteri: — Pleronmalidae) — were 
vecusionilly Observed silting an egy Masses fon) 
December 1904 to Linuary 1985 (ria. 1), 


OA SA PIBLD. M.A. KELLER & A.D. AUSTIN 


Parasitotd eae lente 
Disseetions of Liboratory-reared female 7 hasaliy 
revealed (hal they emerged with a substantial 


complement of eyes and then slowly mittured firther 


cogs over lie (Pig. 4). showing that 2 basis isa 
syHoVigere species, 

Most of the parasitoids collected From the field 
were carrying substantial numbers of midure egys 
(Fig. 4). indicating that ege-limitation was not 
common inthe field at this time, In curly November 
in 1994, three different fembles that discovered egy 
inasses inthe afternoon reniined On then overbight 
alibough no intruding eonspecilies were observed, 
Two of these females were disseeted and found te 
have exe loads of one and three eggs, respectively, 


Behavioural observations 

Yirastioids searched for hosts by flying between 
horchound stems, and then walking rapidly up and 
down the stems, palpating the surface with thei 
antennae until they had located an cee mass. While 
scurching a stem, they offen passed within a few 
centimetres OF an ese mass without detecting if and 
so-did nol appear to be detecting exe miusses using 
visual orchenical cues, Thus, location seemed to he 
by physival contuch Upon contiaeling un ese mits, 
Wusps examined unly one ob a few host ewes. then 


commenced oviposition (Bin ef al. 1994). 1 one or 


more conspeeiies were present, wasjys engaged in 
agonistic behaviour (Wilson 1961), 

Competnion for egg masses differed widely 
herween the (wo years (Pigs 5.6). Tn 1994, there was 
a peak of piwastlold activity Orn December 2 (ig. 5). 


On this dale several remarkable Observations af 


parasitoid competition were recorded, Th one 
instanee. maximo oF 14 parasitoids was observed 
simultaneously Competing for aceess loa single ege 
Hass. dn another observation, five purasituids bad 
discovered an ese mass as it was being laid by the 
female AL rutile. They were following the bug, 
parasitising the eaes immediately they were laid aod 


Eqg load 


Fig. 4. Mature exe loads of female To basaliy collected 
benveen 30 November aml 20 December pn (994, 


were fighting each other for possession of the 
incipient egg muss. Lhree other cases of immediate 
patch discoveries were recorded on the sume din. 
Avenoseelis ratila showed only rudimentary 
delensive behaviour, occasionally kicking at the 
parasitoid but this had no deterrent effect on the 7 
dasaliy, Instead, the parasitoid sometimes responded 
by directing its aggressive behaviour toward A, 
rulilid. 

In contrast to the high peak of competition 
observed a (99d. gompetition remained Tow 
thronghout the entire sampling period m 1995 (Pig. 
5), This difference in intensity of parasitoid aeivity 
between the Iwo years is alse reflected i the time 
until disvovery of patehes (Fig, ©) In 1904-95, 
almost allege masses were discovered within 7 hat 
being laid, whereas in 1995-96, the vast majority of 
eve masses remained undiscovered jy the same time 
period, resulting ina highly significant difference 
between the vu curves (Log-rank test, e* = 100.9. | 
dh. p< 0.001), 

Sample sizes for continuous time records of patch 
exploitation and defence were small and 
observations were sometimes incomplete. precluding 
un extensive anmulysis of the time and sequence 
strueture of behaviour Flowever, the observatrons 
did confirm that the patterns of behaviour seen in 
previous laboratory studies (Wilson 1461, Cuniber 
1964) rellected those oceurring under Tield 
conditions. When alone, wasps successively 
examined, oviposited in and marked hosts hefore 
examining the surrounds of (he cee mass and finally 
leaving. dn cach of the four observitions. sell: 
superparasitisen (i.e. double oviposition in the same 
host cog by the same femule parasitoid) did not oecer 
before the egg mass Was fully depleted (ic. all host 
eves in the eve muss were parasitised), and only 
becurted after that in one observation, when the wisp 
sell-superparasinsed three times before leaving, 
Upon depletion of the pateh, in two of the Tour 
observations wasps embarked on penods ol 


Mean (Max. no of wasps) 


Hig. 5. Comparison at patel competition herween years 
mean (+) SD) af maximum noniber al 2 devalis onary 
Ce THUNS 


BEHAVIOURAL ECOLOGY OF TRISSOLCUS BASALIS oH 


—— 1994/95 
—— 1995/96 


% Discovered 


Time (hours) 


lig, @. Comparison of pate discovery limes between years 
Tor 1994 (26 November -12 December) and 195 (23 
Novernber- 14 December). 


‘defence’ af (he egg muss. These defence periods 
consisted oF alternating bouts of ‘stabonuary? 
behaviour (motionless, sitting on the ege mass) aod 
‘patrolling’ behaviour Gripily darting from one side 
of the ega mass to the other). This apparently pre- 
cnipuve pateh defence behaviour continued for 
upproximately Tl omin and 2h 30 min in the two 
observations, respectively, despite the fact that no 
comipelilors Were present. 

When contests between two individuals occurred, 
sequences Of agonistic behaviour developed. These 
sequences exhibited the same major characterises 
as those observed by Wilson (1961) and Cumber 
(1964), including the establishment of resident 
intruder roles. Fights occurred cither on the first 
encounter, or after a brie! period of mutual tolerance, 
The tendency for individuals to light appeared to 
Ncrease alter successlul completion of one of more 
ovipositiens, although occasionally individuals 
becume aggressive jmmediately upon arriving alan 
eer miss, and before examining the host eggs or 
oviposifing, Following the onset of ageression tind 
role establishment, the itrider usually retreated tor 
the underside of the leaf when attacked by the 
resident, out of view of the resident (eae fapses 
were always phiced on the upper side of the teal), 


Cumulalive boul lenatty (fH) 


18] 
n wu ao iu HU (io hb 
Flout number 


hig. Comiiative beat leverh plat fin crete belay vai 
OF omiculer im opuiewise vantest showed two ahnapt 
Chee Th Bout ena Tesi an Hieeedise Chere grraw yen 
a deereused Gripe arrow), 


Cumulative bout length (h) 
c 
[=> 
_ 


iv] § 10 15 20 25 
Boul number 


Mig. 7 Cumulative bout length plot for retreat behaviour 
of the iitruder ti a paiiwise contest. showing a single 
ubrupt increase in bout lengahy Garrow )- 


searched the 
or fed on nectar in 


und cither sul motionless, groomed, 
surrounds OF the eee ass, 
Nowerheuds. 

In two cases Of pairwise contests, abrupt changes 
in bout length were observed. Inthe first observation 
(Fig. 7), the intruder switched from short to long 
retreat bout lengths, suggesting that it had ceased to 
compete for possession of the cue iass, and had 
begun wailing for the resident lo leave, In the second 
observation (Fig, 8). which lasted 36 min longer the 
intruder swilched twice: lirst to longer relteat bout 
lengths, and then back to shorter ones, which were 
SUT longer than those inthe initial period, 

The intensity of bouts of aggression varied widely: 
from no-contest encounters in which one individual 
retreated without retaliation, to infense escalated 
fights. in whieh individuals locked together and 
rolled across the leal for up to 300s. Although no 
obvious injuries were observed, sometimes both 
individuals tell from the teal, temporarily losing 
Opportunities to exploit the egy mass amd leaving i 
unvturded, Although some individdsls recovered 
their position on (he egg mass quite rapidly. others 
did not return, showing that in addition too the 
possible risk af injury, engaging in an escalated fight 
also entitled the risk OF permanently losing HWecess 1 
the eee pass, 

Conspecilic superparusilisny (oe, oviposition tn the 
same host eee previnusly parasidised Once or mare by 
a different female parasitoid) was common, either 
Wiel Wasps Visited cae iasses sequently or 
simualiineously, In one extreme case, a mass of 20 
set oul oon the day of peak competion 
(December 2) in 1994 recenved 126 ovipositions over 
p period of 9 hh, 


Discussion 


The host popuhition observed im this study 
underwent marked Puctuarions 
Populitions peaked i ospriig. bot eges hed ly the 
overwintering population ob adulis ab this time 


seasonal 


vw SN PIED MA KEELER A A DD ALS TIN 


tiered high levels of predition und parisitisn aid 
few pyniphs cmerged to form the new generation, 
Hhis mortdity, combined with) the deeline tn food 
avaihibihity resulting frond the senescence of host 
plants, Ted tou sharp decrease To host population 
levels by Tie nd Gf the December Therefore, bor the 
parasitoid Gere was a peak iy absolute plumbers ot 
host exes dyailable for approximately tWwer ionths 
front late Qetober to early Deeember Alita 
depletion ol wee reserves mi 7D hesaliy was rare 
during this period Th inay hive had significant efheets 
On forum Behoyiour when (lh oecupted, eaustie 
wasps fo shay leger and fund ese misses. rather 
Thun Contin Faring. 

The eteunetnce oF  hyperparasitism ly 
Avrrcliveides sp. may also hive exerted G seloetve 
pressure Or wisps awit low cae load. causine 
twendeney rewired staying dnd giuinding egg passes. 
Aerlyiidey spi previously been reebvercd Tair 
eve jWasses ool Vo viride it custern Australia by 
Churke & Seyntone (1992), These quthars shawed 
(hil the species isa fy perparusttont of 7) bawalin, bul 
sugeested iLmay be only casually associated with 7, 
hasalis, Clarke & Seymour (1992) did not repert 
Which stage of Z drisetiy the hyperpanisitoid attacks 
bul in observations made i Wie present study: wasps 
only sal on (he exe masses withbul probing or 
OVIPOSMp, Suggesting that they may have heen 
wilting for the immuiine 7 beweliy da reach a hater 
larval stage, ar the pupal stage. Attempts to Observe 
behaviour oF the dereefiveides: sp. incl cour it in Whe 
lubaratory were vosuceesstil, as ik was only 
dbserved a few Uines in the Jield and: was oever 
recovered from A, itil Gee MUSKeS 

The Trequent ocearrenee of superparasitsny by 7 
beiliv in Whis study provides lurther eymence that 
superparasitien) is vominTon iT nature Canssen [Yad 
van Alphen & Visser 1990). tis eeolowival 
importinee far Lo hitsaliy as underscored hy the 
Hichine That (he probability of The superparasttis ie 
Terie oblige offspring (oii a host is very high vt 
she Superparasiises soon aller the other fennale 
oviposits inthe host Wielledal 1997), As Wis leaves 
the oftspring of the first female ta oviposit al 
subsuintiil ask. the high frequeney of vecarrenee af 
stiperparasidisin and its high fitness pay-allrnay Have 
favoured the evolution af pateh defence in F baselis. 
Th adition lo the Rack thal Gee pousses are of a 
delendable size, us noted by Winige (1982), The drop 
in the pay-ofl fren) superparisilisny lowards vero 
ifter & ob (Kield er a 1997) may explain the 
observation that fenules aiborted Gviposiiign in an 
eae uss tit had Been parisitised as much as 9 h 
earlier Still this raises the question of the 
Mechanisin af host discrimination, which needs ta be 
pursticd ie Pitare: studies, 

Observahions of puleh exploit ancl delence 


revedled behaviour patterns simi kin te those seen i 
previous laboratory studies (Field!) and sugwest init 
belaviote is adapted to bial levels of competition, 
When alone on an ee miss, wasps engaged in 
extended periods of defensive behaviour boll before 
and alter the eve irtss wits Fully depleted. suggesting 
a high ie expectation” of conspecifics. arriving 
and superparusiising. Chis behaviour could result 
from oa combination of dinate expectation of 
competing set by natural selection. aid a flexible 
response based On experience from previous eez 
messes, Alted  restdentypteuder tales were 
established tn pairwise contests, Tilruders showesl 
ubrupt vacredses in periads of rethaat behaviow 
CUFT COMENES wh Walled OUL OL Visto Of residents 
(ander he dear), suggesting Whey were waite for 
residents Wy leave the eee tnass. Alihiweh patuder 
hehavinne was pot followed jo deri. tn the second 
comeust analysed (rig. ®t ts probable tat the fest 
switel resulied from the intruder rikiligh an 
extensive search Of the sUPOU ine are Wy check Cer 
other Uinoceupied: cyan rasses, uneb the second: (roy 
its retuuming to the origimal patch, Exploring the 
SUPPOUNd ws Lor aHeniiive HOST Gee masses 1 he 
current one May be particularly important wher 
sihy (nubvidaals ure Competing Tor a) Ege rns ard 
giining access iS difficult: a Siteition thal was 
Hequently Observed in-times af peak eompetirioan 
The dynamics of patch delence im such sittiitions 
invelving more thap lwo individualssire contplex dn! 
difHeull lo fifer trom: the present study, Altiougl 
subslantial Muetuations in the levels of compelition 
(lo beet and conipetitian is someimes less iitense, 
periods of extreme Competition muy nevertheless 
have played a dmpertiadt tole ta the evalytion at 
foraging behaviour. 

The Observations made in ibis siidy reveal sere 
Hovel patterns TH Lie agonistie belmwiour of 2. Leselty 
dnd indicute the Appropriate ceolovicn) context a 
WHICH Lo tivestigane them Tureen ie the liberatapy 
The factors leading to the onset ofigeression did the 
Mechwisiis Ob Coplest fesoliton Che. eslablistiment 
Ol resident-intruder roles) hiwe rarely been studied ny 
parasitoids (bul see Petersen & Tardy T296) and are 
the subjeet oF current jvesdivation iin 7 heryeelia 
(Pield & Calbert unpub, Also, a qiuntibative 
Mi Vsis. OF The Hime-struetire OF interactions betwee 
Hehting purasitioids fas nol preyiwstly beer 
allemnpled. and the present results mmaicute (hat (hs 
muy he both possible and ol wreat interes! to 7 
Pasatin. 


Acknowledements 


feo wish to thank G. Thoms fer the tse ol bis 
property, C. Culbert for programining the multiple 
chinge-point test and 1. Mound, M. Kakkinn and [ 
Bird for eriticisms of the manuscript 


BEHAVIOURAL ECOLOGY OF TRISSOLCUS BASALIS TI 


References 


VAN Atpibn, J. J, Mo & Vissrk, M. E. (1990) 
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parasitioids, A, Rev Ent. 35, 59-79. 

Rin, F, Vinson, S$. B. STRAND, M. Ru. Conazaza, 8. & 
Jones. W. A.J. (1993) Source of an egy kairomone Sor 
Trissedcus bayalis, a parasitoid of Nesara viridule, 
Physiol, Ent, WB. 7-15, 

Charkh. A, R, (1990) The control of Nesura viridula 1 


with introduced eggs parasitoids in Australia, A review of 


a ‘landmark’ example of classical biological control 
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— & Srymour., J, &. 
Aerocliseides Girault and Dodd (Hymenoptera: 
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viridila (lL) (Hemiptera: Pentatomidae). 4 Ausi Ene 
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Cimapk, Ro A, (1964) The egg pitrasite complex 
(Seelionidae: — Tymehoptera) of shield bugs 
(Pentatomidae. Acanthosomidae: Heteroptera) in| New 
Zealund. NZ J) Sei, 7. 536-554. 

Piteb, S.A. (in press) Pateh exploitation. patch-leaving and 
pre-emptive patch defence jn the parasitoid wasp 
Trisseleus basaliy (nisecta: Scelionidie), [liology: 


———. Kenner, M.A. & CALBERY, G. (1997) The pay-off 


from superparasitism in the cee parasitoid Trryselcus 
basatis, invelation to patch defence. Aeol Kur, 22, 142 
149. 

Goprray, A. C. J. (1994) “Parasitoids, Behavioral und 
Fyolutionary Leology” (Princeton University Press, 
Princeton, New Jersey). 

JANSsiN, AL (1989) Optimal host selection by Drosophila 
parusitoids inthe field. funet Beal, 3, 469-479, 


(1992) "Two species of 


Marriacet, 1... VINSON, S. Bo WILLIAMS, HJ, ALbrici, J, 
Ro & Bin, (1993) A long-range attructamt kairomone 
for egg parasitoid Trrsselous basalis, isolated trom 
defensive secretion of is host, Nesara virtdula.  Chene. 
Evol, 19, (167-1181. 

Nosik, N.S. (1937) An egy parasite of the green vegetable 
bug. Agric. Gaz. N.S.W. 48. 337-341. 

Prrirsin, G. & Harpy, 1 C. W. (1996) The importance of 
being larger: parasitoid intruder-owner contests and theie 
implications tor clutch size. Ania Behe $1, 1363- 
1373. 

PowenLl. JE. & Sumparp. M. (1982) Biology of Australian 
and United States strains of Trisyeleuy Paxalis, a 
parasitoid of the green vegetable bug, Nesara viridisse. 
Aust. J, bel 7. 81-186. 

Turner, J. Wo (1983) Influence of plant species on the 
movement OF Triwelcus basaliy Wollaston (Hyimen- 
optera: Scehionidae) - a parasite of Mesure viridula Let. 
Aust. ent. Soc. 22, 271-272. 

VotgKork, N, & Conazza, S, (1992) Growth patterns of 
teratocytes Mm the inimiature stages of Trissadouy bewelis 
(Woll.) (Hymenoptera: Scelionidue). an eye parasitoid of 
Nezara viridula (1) (Heteroptera: Pentatomidae}. fat. J. 
Insect Morphol. & Enbrval. 21, 323-336. 

Waact, £K, (1982) Sibmating and sex ratio strategies in 
scelionid wasps. Keol But. 7, 103-012. 

Weber, C. A. SMILANICK. J. ML, Ennnr, LL, EL & ZALoM. B. 
G. (1996) Ovipositional behavior and host dis- 
crimination in three scelionid egg parasitoids of stink 
bugs. Biol. Control 6, 245-252, 

Wir son, (1961) Adult reproductive behaviour in Aweleus 
hasalis (Hymenoptera: Scelionidae). Aust Zool. 9, 
739-751. 


WOODWARDOSTRONGYLUS PETROGALE SP. NOV. 
(NEMATODA: CLOACINIDAE), FROM THE STOMACHS OF 
ROCK WALLABIES (PETROGALE SPP.) FROM ARNHEM LAND 


By I. BEVERIDGE* 


Summary 


Beveridge, I. (1998) Woodwardostrongylus petrogale sp. nov. (Nematoda: 
Cloacinidae), from the stomachs of rock wallabies (Petrogale spp.) from Arnhem Land. 
Trans. R. Soc. S. Aust. 122(2), 73-78, 29 May, 1998. 

Woodwardostrongylus petrogale sp. noy. is described from the stomachs of two 
species of rock wallaby (Marsupialia: Macropodidae), Petrogale concinna Gould, 1842 
(type host) and Petrogale brachyotis (Gould, 1841) from Arnhem Land, Northern 
Territory. The new species is distinguished from congeners, W. woodwardi (Wood, 
1931) and W. obendorfi Mawson, 1976 by the presence of four pairs of oral denticles 
compared with six pairs in W. obendorfi and 16 pairs in W. woodwardi, by the spicule 
length which is 0.90-1,07 mm in W. petrogale sp. nov. compared with 1.4 mm in W. 
woodwardi and 1.7-2.1 mm in W. obendorfi, by the length of the female tail which is 
0.22-0.23 mm in W. woodwardi, 0.18-0.22 mm in W. obendorfi and 0.11-0.17 mm in 
W. petrogale sp. nov. In addition, the vagina is 0.7-1.0 mm in W. woodwardi, 0.8 mm 
in W. obendorfi but only 0.31-0.48 mm in W. petrogale sp. nov. The characteristics of 
the genus are considered as well as its relationships within the Cloacinidae. 

Key Words: Nematoda, marsupials, rock-wallabies, Petrogale, Woodwardostrongylus, 
new species. 


Tre ans ef the Revel Somaya So Auge, (L998), 9222) 73-78 7 


WOODWARDOSTRONGYLUS 
CLOACINIDAE), FROM THE 
(PETROGALE 


PETROGALE SP. NOV. (NEMATODA: 
STOMACHS OF ROCK WALLABIES 
SPP.) FROM ARNHEM LAND 


by J, Bhyeringn® 


Sumnuiry 


BEVPRIDOD, TP (TOUR) Wanelivcdostndiy petoagale sp, now (Nematodes Cloacinnlie), fromthe stomachs 0 
rock wallabies (Perrouele sppo bron Armbent Land, Tres We Soe. XS Meare, 122 (2) 75-78 29 May Ys. 
Woodiidosnuney los petal sp. nov ops desernthed: from the stemaehs ol iwo species a? roek walhiby 


(Marsupiatia : 


Maeropodidie), Perogale conenuid Gotld, 1842 (ype hosti and Pariwdle becliveniy (Gol, 


PO) trom Arnhem Lund, Northern Territory. Phe new species is dintingiished from Gongeners. Wo wrcedtvesdi 


(Wood, LOS 7) ae Wi endian de M 


vwaert L976 by Te presence of four pps al onl denticles Compared willy sty 


pens ie WE oPbeydeedi and 16 pars in Wo woedword:, by the spicule tone th whitch ts 0.90-L07 min in i perraueate 
ap ney, conipared with Frm Wo ypeodiweritt and 17-20 im in Wo obentorfi, by the lentil of the female 


Toth whieh is O. 22-0, 23 lo We teed. OTK 


“27 jan in WE ebcudoeti and QE, Pim WW, petreigale 


sponoy tnaddibonm the vagimiiis (7 DOs We hod Ue inn in Wo abendor? bitonty 131 a8 iin 
WW. petiagele sp. ney. The characteristics of the genus-are considered as wells ibs relationships within (he 


Cloweurilie 


Kiev Woks. 


tiiroduction 

One ol the most dnusudl eenerd of nematodes 
OoC Une Wy the stomaehs ot Kirigarous und 
willibies is the cloacinid genus Weeewurdn 
Mraneviiy Wahid, 1964 which ts found in tannels i 
the superficial squamous epithelinan of the stamach 
und Oesophagns rather than ja the lane ob the 
stomuch oor hare maitestine oor coiled around 
vesuphageal papilliwas is the case with most of phe 
other members of the Eimily (Woed 193.2 Miwsen 
1971, 1976; Beveridoe Ac Spratt 1996). Two species 
are cirently Keawe willl this genus, the: type 
species, WM weerdwird? (Wood, 193.1) origtimally 
deseribed From Woodward's wallaroa. Moers 
mens Weeder? Thormias. LOOT, frond the north 
Wo Western Austrilia (Wood [98d. Wahi 964), 
slibsequcnlly redesenbed from Pearson Island: rock 
wilhibies, Petrogale lafeclis pearson’ (Thos. 
1927) Troms Soul Australie (Mawson LU7 by ane We 
dhendori Mawson. 1976 froin tre whiptiiled 
williby, Maeropis pares) Beret, ES3S (ype host, 
theewalllaroo, AL robust rebuatis, Gould, (S41 and 
the red-neeked wallaby, AA rifowerseus (Desmurest, 
IS17), trom: north-eastern New South Wales ane 
souti-eastend Queenshind (Maiwsoo 1976). The litter 
species Wits subsequently reported as a comiion 
parasite Of brush-railed rock wallabies, Mmefibers of 
the Mucovule poeniciiaa complex (Po aystiniliy 
Kuinsay. TA77. 2 eodman Thomas. 1923, 2 
herberti Thoms, 1926, 2 iernate Gould. 1842. 7 
muireeba Eldridge & Close, 1992, BO penned 
(Gray. 1825) and Po shaman’ Eldridge & Close, 
J992) Tron eastern Queensland by Beveridge etal. 


Deparment of Verrinery Semmes, The Liveenty al Melbourne 
Purkville View A062 


Neruda, (hirsupluls, ruck-willubtes. evade. Weed iiyreiiis. new species 


(O89) aind bus since heen found alser in the agile 
walhiby, Mecropus cudis (Gould, TS842), und (he 
swamp wallaby, Willabia bicolor (Desmurest, S04) 
(see Spratheral 1991). Spratt eral (1991) dsted ain 
additional andescribed species Ob Werle: 
stiome vis from the nabarlek, Petourle concn 
Gould, [S42 and the shori-eured tock walliuhy, P 
hrachyenis (Gould. E84), fram the Northern 
‘Territory. Tnothe present paper the undeseribed 
species fisted by Spratt es a (1991) is deseribed, te 
didanostic feattites Of the eens are reassessed und 
iis poston within the existing classification is 
discussed since the venus bas in the past heen 
variously allocated to the Strongyloiden (Walid 
1064) and the Trichostrongyloided (Mawson 197 1), 


Materials and Methods 


Nematodes were recovered fran’ dhe preserved 
cureases OF rovk wallabies provided by Dr J. bk, 
Nelson, Monash University Melbourne Vie, 
Immediately afer shooting, curcases were perlusca 
with TO% formol saline yin the Jel ventricle 
lullowed by immersion of the entire carease in LO% 
formalin, Nenitodes revovered trom the gastric 
Mucosa Were slored in 70 ethanol and were clawed 
in hiclophenol for cyanitiation. Drawings were made 
with the amd of a drawing tbe atlached to an 
Olyinpus BH microscope, Measurements are 
presented in mmous the range fer 5 specimens 
followed by the mean in parentheses. To examine the 
localisation OF nematodes within the SUstric MUCOSA 
small pieces of purasitised stomach wall were 
embedded i ip was and sections, cubata thickness of 
S in, were stuned with haematox sti and ¢osin, All 
nemalode specimens have been deposited im the 
South Australian Museum. Adelaide (SAMA). 


id | BEYERIDGE 


Woodwardostrongylus petrogale sp. ny. 
(FIGS 1.13) 


Melitype : 
Atehem Land vt 
SAMA AHC 30592. 


sume dita. SAMA ATIC 30593, 
Tarai ype BO ue dite. SAMA ATIC | 47490, 


Giher mitrerial exanrinert (ron stommieh ot 
Felravale brachyots, Artem Land NV) edt dk, 
Netson, Dandeye i ee, 20-99, SAMA ATIC 
17, 


5 from Stomach ol Pedragale cence, 
IxXtl977. coll do bo Nelson, 


Allan. <) 


Devseriphion 

Slender elongate nenmlodes, cuticle covered wath 
numerous fine transverse anniliiions. Mouth 
opening Uny. dorsoventrally elongate, apparently 
rigid Joteral murgins of mouth opening, each with 
row of four relraetile, dome-shiapeu denticles: 
adhhionil parrot hoy denticles al dorsal and ventral 
extremities of aich row. Two amphids and) four 
suibmediin papullie lateral to rows. of denticles; verve 
tissue extending posteriorly and literally fran 
scuisury pilpiiae. Sabcutieular region of anterior 
extremity heavily selerotised. Buccal capsule thick 
walled with Liint triisverse striaitlans: anlerilin part 
of buceal capsule dorsoventrully clongate in apical 
vielvs of Dead, become ciecolir posteciorly, buceal 
castle Supported externally by LO promiment bands 
of muscle running from external surface ob buccal 
capsule ta longitudinal somatic musculature. two 
bundles of inuseles present dorsally and two 
ventradiy, lwo thick lateral musele bundles anid: four 
slender submediin bundles. Oesophagus long und 
slender corpus eylindtieal oarrowing slightly to 
form short isthimuss isthmus merging tito elongate 
bulb. Nerve ring surrounding 
OesOphagenl Corpus ail SHIT: secretory excretory 
pore in mid+egion ol oesophageal bulb: deirid at 
level of seeretory-esereLory pore: 


Male 

Total length 131-155 (140): maximonm width 
0.1 7-0,23 (0.20%; buceal capsule O.020-0,037 (0.032 } 
in length, wrth pn biter views O01 7.020 (0.019), 
in dorso-ventral views (L0TO-OQ.013 (0.012), 
oesophagus 0. 79-086 (O83)2 nerve ring lo anterior 
end O4A0-048 (42): sveretory-exuretory pore tu 
anterior end Q.46-0.57 (0.52). deirid to anterior end 
0.60 10.60). 

Bursal lobes short, of approximately equal length; 
ventral lohes joined yentealiy; dorsal lohe small, 
slightly shorter than lateral lobes, nol clearly 
demarcated from lateral lobes. Ventral rays slender. 
upposed. reach margin of bursa. extemo-lateral tay 
shehtly stonter than other literal rays, divergent [rom 


Vineiien of 


them, shefiily recurved peur extremity nol reachine 
margin of bursay medio-kiteral and veatro lateral 
rays slender apposed. reaching margin of bursa 
extemo-literal riy arises from lateral trumk, serail, 
does nobrewch pabeny of bursa. Dorsal cay brtireites 
close to Origine branches long, Slender, arctate: 
seoundury Hfavision into branchlets Geeurs nedr 
cxremily abrayy external branchlets short, direeted 
postero-hitevally, da not teach matin of bupsy: 
internal brinchlets longer, dirceted posterorly, 
almost reach margin of bursa. Genial cone 
prominent. anterior lobe hatte. conical extenus 
almost to lini oF veutral lobes: posterior lip spall 
Will) parr oof prominent postermorly  directes 
appendages; pubernaculanm absenk; central cordite 
and lateral paired elongate thigkenings ol spicule 
shewths present at then junctions, spicules wlungale. 
alie, O90 1,07 (0.97) long: anterior extremities 
ircceuhurly Knobbeds distal Gps blunts die diminishes 
Th width towards spice tip, 


Hemale 

Length T64-22.4 (200) masini width Obs 
O29 10,27), buccal capsule O.027 0.087 (030) in 
Tenth. width in lateral views 0.020. in dorse-ventral 
views (010-0015 (L013), desophigis 0.92 [28 
(LOLS nerve ring te anterior end O,24-044 (O39) 
seercloryeexerclory pore to anterior end 0,470.65 
(0,62); deirid (o anterior end O.S58-0,63 (0.62), Tat 
short. confeal (LT 1-OLE7 (05). vulva inomectiarely 
anterior to anus, 023-0. 42 (26) trom posterior ond, 
Vaging short, straight. direeted anteriorly, 031-048 
(0.39); vestibule ta form of Ye wath thick pmiscutae 
wills, O.1S-0:20 (0.174) Tong: sphineler aril 
INT bolo GP approximately equal lengths. very 
variable in length, each O.08-0.25 longs utent parallel. 
ru anteriorly trom intundibula ege ellipsoidal 
(). 03-0, 1A (O13) 4 0,072.09 (0,08), 


Locetisation within sramach 

Nemiutodes lig in small sinuous tunnels tm the 
superticral ayer Ob the stratified squamous 
epithelium of the forestomach. Nematodes were not 
found penetrating as deeply as the lavine propre 
and the presence of the nerpatodes in the squires 
epithelium provoked no inflammatory response. The 
umerior ends of the nematodes were buried i 
tunnels while the posterior ends lity (ree in the gastric 
lumen, 


Discussion 


The nematode species described above belongs to 
the Cloacininae Stossich, 1899 beeaise it possesses 
a cylindrical bueewl expsule. a burs in whieh the 
dorsal lobe has four branches. the externe-dorsal ray 
arises from the kiteral truink and a cervical groove js 


NEW NEMATODE FROM ROCK WALLABIES TS 


LX 


.\ea 


3 
| | 


Pigs 1-12. Weodwardosrongvius petrogale sp. voy, 1, Anterior end. Tateral view. 2. Cephalic extremity, Jateral view. 3. 
Cephalic extremity, dorsal view. 4. Cephalic extremity, apical view, 5. Transverse optical section through anterior end al’ 
buccal cupsule. 6, Transverse optical section through posterior part of buccal capsule showing muscle bands running 
midrally from outer wall of capsule, 7. Bursa, lateral yiew, §, Bursa. dorsal view, 9. Bursa, ventral view. 10, Posterior end 
OF inale, dorsal view showing spicules and thickenings of spicule sheaths at their junction. U1. Female tail lateral view. 
12. Pemiale genital system, lateral view. Seale bars = 0.1 mm 1, 7-12; 0.01 mm 2-6. Legend : a, amphid: d, denticle: de. 
deinids e, seeretory-exerelory pores i infundibulum: UW, lateral thickening of spicule sheath; m, mouth opening; a, nerve 
ring: s. submedian cephalic papilli: sp. sphineter: t, cential thickening at junction of spicule sheaths: v. vasina: ve. 
vestibule, 


Vy 1 BEYER 


= ’ " er oe Tuy 


~~} se 


Pip LA Histilogigal keaton af othe: stimngel wall al 
Perrugade ciniiane —oshrowing — laeuliatient al 
Wialivedeaviaie is poravele spo nove Cy Tt ol 
super Hepat aquumels epithelial timely eros! formed 
by the nematode dad the lack of any leitlannniory 
reavnon ithe epitteliian, Seale bare dh barn 


hicking, Ho belongs to the genus Woodwarde- 
Mromgylay becuse 1 possesses a heavily selerotised 
nonil rezion. av Gransversely stated buccal capsule 
and a vow of sclerotised denticles on cither side at 
(he mouth opening. the latter characteristic being the 
INust obvieus featipe defining the genus. The 
specnmens deseribed above ure distinguishable fron 
Woowemlward) and Woobenederfi primarily by the 
Humber ol pairs of selerotised ural denucles, 
Woodiedrdostraneylis \woddwerdl possesses: 16 pairs 
Of denticles wile Wo ofencdori possesses six pars, 
Inailbol the specimens from Pelrageale spp. Tron 
Arnhem Land there are four large pairs of denticles: 
Avcaeh end of the cows oF dentteles. there ys a pair 
of tiny denticles which has hot been meludedl in the 
determmation of the punmber of pairs of denticles 
becuuse these denticles ure oot obvious in baler 
views and are only vlearly visible in an upical view 
ol the motth revion, The same terminal pairs oF tiny 
denticles ure evident mp ihe seunning electron 
Wicroeruph af the cephalic extremity al Wovhendarfi 
published hy Mawson (1976, fill) although the 
feature is Hat entioned ip (he deseription and was 
nob taken into considerbon when determining (he 
number of pairs of denticles on each side of (he 
mou opening, Whether these same terminal pairs 
ol tiny denticles ure present in Wo Wreeelwerned? 1s not 
known, In addition. the specimens described above 
differ fron eongeners in spicule length, being 14 
THM HL WO seeeediverredé (Wood 1931), 172.7 mine i 
Wo ebendorfi (see Mawson 1976) compared with 
090-107 nim in the species described above, There 
is.alsow corresponding difference in the length of the 
yagi which is O.7-L.0 mim tong in Wo weenie, 
O68 mm in Wo ahenderi and O31-0.48 mim in the 
species deseribed here. Therefore, the material from 


rovk wallabies trom the Northern ‘Lerriory ts 
considered Lo represent a distinet species amd tle 
ANG Weeedwerdastnaneyiay peirorale sp. roy is 
proposed for al, the name being derived fron) the 
generic narne of the hosts. 

Th compuniig the deserpuon of We new species 
wilh those of its Congeners, several morphological 
characterises of the genus warrant comment dae te 
ipparent inconsistencies or errors ip publishes 
Jeseriplions, 

The vesophagus isdeseribed in the other species ius 
heing slender und clavate yet in We perragele there is 
a distinel constnction ab tie level of te nerve ring 
und the oesophagus is clearly divisible jalo corps, 
isthmus and oan elongate bulb. Specimens af ie 
dhendorfi were examined und the same subalivisiats 
Of The oesophywus wre evident althotigh they have tet 
been dlisted or desertbed in the Titermtuce, The 
structure of dhe nesephagus is of considerable 
hiagnuine sizmficanee heeause i is ahayaie 
woneru ol the tribe Cloucininea (Stossich, (ROO) af 
the Cloawciiinae bat i subdivided tite corpus, 
mihnnus and bulb in the tribe Pharyneostroney lines 
Popova, 1952 into whieh Wheewerdes ay is bas 
been placed (Liehtenfels TORO) The revised 
Wlerpretaion Of these morpholowieal charaeters 
Therefore beeares eoncorduit with the: earrent 
laxononne position of the genius, 

Th the original deseription of the genus, Wahn 
(Mod, fig, Th) desetibed and Uhistrated a 
“wubermiculin’ fh the type species, We Waoeeiver till 
Th the orginal desemptton of the same species, 
however, Wood (199-1) had heen muh more caulions 
qd had deseribed ~* yin accessory piece... present is 
iin frresulur shaped siueiure’ which “uppeats: to 
finetion as a auide for the spicules’. Mawson (1976) 
slated That a SUDerNACUTUNY Wis present uy 
Pbendorfi ul did not Whustrabe the stricture, Durele 
Desset & Beveridve (M980) illustrated the 
“gubermuculan’ ob Wodhenderfi but iiselour (rei 
the tHusteation thi the structure 4s not 
gubernaculin but the cordate dhiekening at the 


junction af the spicile sheaths (Beveridge |982) A 


woberpaculit is absent in Wo perreauade, allhougly at 
Joes possess a cordate thickening and paired Tater 
thickenings of the spicile sheaths af their junction fo 
vommnon with all other cloaemids iad ehubertiids 
that have been examined for the presenee ot tlie 
structures (Beveridge |9O87) I therefore seems thost 
likely that a gubernaculum is absent in this genus and 
the cordiule central thiekening at the junction of the 
spicule sheaths hus been mistaken fora 
vubernaculum jn the past. This problem pertains to 
several genera of the Pharyngostrongylined and has 
becn discussed in detail by Beveridge (1982). 

The morphology of the feniale genital systenr hus 
not been described in detail for Wo wacedivereli, 


NEW SEMAPODE PROM ROCK WALLALIES i? 


allhough Mawson (M7 by rllusteated an essentially: Y- 
shaped ovejeeloe with parallel, wmphidelphie uleri in 
her redeseriphion of thts species from rock wallabies 
from South Australia. The ovejector was deserihed 
lor Wo ohendori by Durette-Desser & Bevericle 
(IYSO) who HfUstrated a short Yoshaped vestibule 
will) short sphincters und infundibula und sugvested 
that the morphology of the oyejector was 
Wtermedite between thar vound in the 
Trivbostrongyloidea and the Strongylotdea. 
Lachtenfels (PYs80) classified the oweyector of 
Woodwardosinnevties as lypieally stroneyloid and 
us her Yoshaped supertivially. but resembling 
relaed gener with J-shaped Gveyeciors im that the 
sphincters and tofindibyhe are short, The wvejeetors 
OF. pelrosale are sumilu to those of Wo ebendorss 
and confirm Pichtenfels’ interpreuiion. Beveridge 
(1Y87) ilustmted the uveyector ab We petragude 
(deseribed stmply as Weedlvuandosironelits sp.) and 
CONT Ted thal it foo ereed with the deseription and 
ullocalron suggested by Lichtentels (1980), The 
oveyeetor jn this genus Ǥ therefoce considered te he 
aw moditcd J-shaped ovejector uceording to. the 
delinitions of Lichtenfels (lOs80). wath the 
modification probably heme a alivect result of the 
slender clongaty body structure imposed upon this 
nematode genus by its cocalisation within epithelial 
tunnels OF the Bitstric Meesit, 

The owwsternatie position oof the genus 
Woodweardoyiongvii bas been the subject of some 
uneeruitinly. The type species was initially deseribed 
as Mioryitvostrae yas seared? indicating a 
close telanonship with the genus Phearwigostronyy- 
lay Yorke & Maplestome (926, Walnd (1904) 
Subilivided the genus Pharvigestrongyiias and 
erceted the new venus Woeeiinedistooerlis Toe We 
Wado) bul provided no explain of as 
possible pelathanships with Phaivigesreneytin. 
Mawson (197) ) redesceibed 2 woodward? front tock 
willities nd ereetod the mew venus Craviiecpay 
whieh she plaved in the trichostroneylon! family 
Aimidustomitidie (Travassos. 1929) based on the 
pipers al Inghs (i965. 1968) dealin with 
copvermence OF (he cephilie Icatures ot penitodes 
OkCUfide buted Hb the stom: linn OF herr hosts 
whe Tis view thal inany of the stroneylale nemidodes 
of Nistratian marsupials were members of the 
Tichostroney low hamily Armdostonmiutidie: 
Subsaqeatly. ab desenbinw Wi eben dari. Mawson 
()926) recognised the synensiny ob Crateceps with 
Woodiostrornevlis Hub placed Wewnltvared- 
oviesiies we the An dostomaticiw, Dhupete- 
Dessel & Beveridge (F980) ry contrast relerred: tte 
suri Co (he Stronsylolein amt Prchlentely (148d) 


pluzed the genus m the strongylord inthe 
Phuryogostrongylinea characterising the tribe 
primarily on the basis of a huveal capsule wath 
transverse strimions und therehy re-ussociuing 
Woodididosmoane\ tis wilh Planiicosmunertys. (he 
genus with which to owas irst linked by Weod 
(1931). Beveridye (1982), in a revision of whe 
Phiryogostrongylinea. omitted Waedweredestien 
fy/as on Ure basis of nneertiinities ds to tts affinities. 
The addition of a new species confirms. the 
characlers upon whieh the cents wits erected while 
providing some medification to the definition af the 
genus. prinevipally in relationship tthe morphology 
OF the oesophagus and Oveyeedor and the absence of a 
tlie euberaculun, Te assocnuion with mernbers of 
the Pharyngostrongyloided ts Supported on the buss 
of aw transversely strated buccal capsule. although 
this character beeurs alsin certain Benen OF the 
relted  (ribe Zoniolaiminea (Popewa, (952) 
(Beverndve 1984). This inerphologiew! character 
appears to be the only feutire upon which alfinilies 
evan be judped because other characteristies ol the 
genus are so highly moadihed ta acommuodite its 
tnusudl mode of existenee within the stomach wall 
What they are plylowenctcally unijformative. 
Therefore, in view of the luck of evidence to the 
contrary. and wilh the limited or even equivocal 
evidence OF assocnibons bused on the presence ol a 
siriited buecal cypsule, jt secs reasonable to 
consider Wordiwardestrong yas asa highly modified 
member of the tribe Pharyowostongylinea, 

The host and veographival distributions of 
menibers OF the gents dre not yet fully elucidated. 
On the basis ob current evidence, Woobendarff occurs 
ina variety of roek wallabies: scrub wallabies and 
hinguroos wong the eastern coast of Queenstand and 
New South Wales. Wo weedeard? is known from 
Kangaroos erie ning From the northwest of Wester 
Australie Galthourh based on ae ¢oo record: tron 
Brikua) ind from rock wallibies in South Austalia, 
while Wo permayale ts knewn tren nek walhibies 
IrontArmhem Land tn ihe Northern Territury. The 
feature common to all members of the genus ts thal 
they Pardsitise rock Wallabies But Hest cetatianshl ps 
Warrant ingore (herough tivestisatron bebop apy 
conelisivos cun he clawe Trond this Ghsenvalion, 


Acknowledgments 


‘Thats are dite fo De Jb. Netson for prosgsianal 
The Parasiles Hpor which thes deseriplion is biosee 
ain ty Dr DM. Spratt for readline a drat ob (he 
HOLSET ip, 


78 I. BEVERIDGE 


References 


BrveripGr, 1. (1982) A taxonomic revision of the 
Pharyngostrongylinea Popova (Nematoda : Strong- 
yloidea) from macropodid marsupials. Aust, J. Zool. 
Suppl. Ser, No, 83, 1-150, 

(1983) Taxonomic revision of the Zoni 
olaiminea (Popova) (Nematoda : Strongyloidea) from 
macropodid marsupials. /bid. No. 91, 1-88. 

(1987) The systematic status of Australian 
Strongyloidea (Nematoda). Bull, Mus. Nat. Hist. Nat., 
Paris, 4° sér. 9, 107-126, 


& Sprart, D, M. (1996) The helminth fauna of 


Australasian marsupials : origins and evolutionary 
biology. Adi, Paraitel. 37, 135-254. 

’ , Chosp, R. L.. BARKUR, S.C. & 
SHARMAN, G. B. (1989) Helminth parasites of rock 
wallabies (Perrogale spp.) from Queensland, Aust. Wild/. 
Res. 16, 273-287. 

Durerrr-Desser, M.-C. & BrveripGcr, |. (1980) Sur ta 
position systématique du genre Woodwardostrongy/is 
Wahid, 1964 (Nematoda : Strongyloidea). Bull. Mus. 
Nat. Hist. Nat., Paris, 4° sér. 2, 77-80. 

INcLis. W. G, (1965) The nematode parasites in the gizzards 
of birds: a study in morphological convergence, Proc. 
Zool. Sac. Loni, 135, 125-136. 


(1968) The geographical and evolutionary 
relationships of Australian trichostrongyloid parasites 
and their hosts. J. Linn. Soe, (Zool.) 47, 327-347. 

LICHTENFELS, J. R. (1980) Commonwealth Institute of 
Helminthology Keys to the Nematode Parasites of 
Vertebrates. No. 7, Keys to genera of the superfamily 
Strongyloidea. (Commonwealth Agricultural Bureaux, 
Farnham Royal). 

Mawson, P. M, (1971) Pearson Island Expedition 1969, 8, 
Helminths. Trans. R. Soc. S. Aust. 95, 169-183. 

(1976) Woodwardostrongylus obendorfi: new 
species (Nematoda : Amidostomatidae) from kangaroos, 
Ibid, LOO, 121-124. 

Sprarr, D. M., Beveripor, I. & WALTER, E. 1. (1991) A 
catalogue of Australasian monotremes and imarsupials 
and their recorded helminth parasites, Ree. S, Aust, Mus. 
Monogr. Ser. No. 1, 1-105. 

Wadip, S. (1964) A preliminary revision of the genus 
Pharyngostrongylus Yorke & Maplestone, 1926, J. 
Helminthol. 38, 181-190. 

Woop, W. A. (1931) Some new parasitic nematodes from 
Western Australia, Rep. ast, Anim, Path. Unive Canils. 1, 
209-219, 


MESORHABDITIS KINCHEGENSIS SP. NOV. 
(NEMATODA: RHABDITIDAE) FROM ARID SOIL IN 
KINCHEGA NATIONAL PARK 


WARWICK L. NICHOLAS* 


Summary 


Nicholas, W. L. (1998) Mesorhabditis kinchegensis sp. nov. (Nematoda: Rhabditidae) 
from arid soil in Kinchega National Park. Trans. R. Soc. S. Aust. 122(2), 79-84, 29 
May, 1998. 

Mesorhabditis kinchegensis sp. nov. was collected in an anhydrobiotic state in dry red 
sand under a bluebush. Maireana pyrimidata (Benth.) Wilson, 1975. This is not the 
usual habitat for Mesorhabditis which is commonly associated with rich organic 
matter. The same species was also found in agricultural soil. 

Key Words: Anhydrobiosis, Australia, Mesorhabditis, nematode, soil, taxonomy. 


Transactions of the Boval Sect ofS. Aust (L998), L222). T9BA 19 


MESORUABDITIS. KINCHEGENSIS SP. NOV. (NEMATODA; RHABDITIDAE) FROM 
ARID SOLL IN KINCHEGA NATIONAL PARK 


by Wakwiek L. NICHOLAS 


Summary 


SHOHOLAS, Web. 


(1998) Mesarhabeits kinchegensiy sp. noy, (Nematoda: Rhabdiidae) from aed soil in 


Kinchevsa National Park, Tras. WSS, Aust. 122 (2), 79-84, 29 Mity, 1998, 

Mesorhubdilis kinehevensiy sp. voy, was collected in an anhydrabiotie state in dry red sand under a hluebush 
Maireane pirinidare (Benth.) Wilson. 1975. This is not the usual habital lor Mesorkabdiiy which is dominantly 
fscice WHR rie organi rouner. The same species was also found in agricultural soil, 

Distinguishing features of this species ure that tn the made the fps ab the long. almost straight, distally lrsed 


spicules, are abrupily (irned ventrally, The formula for the aerapgement al the bursal papillae ts (2 


br 3) with 


none olthe pupillae (used at (hei bases. tithe female the Gil is long and pointed so thatthe distunee thai the 
posreribn Vulva to the anus is aboutone anda bull tines the tal length. 


Key Warns: Anhycrobtosis, Australia. Weserhadiditis, nematode, soil. laxononny, 


Introduction 


Most species of Meyerhabeitiy ive been reported 
from tieh deeaying organic matter such as humus, 
rolling wood or dung, Several speeies are usually 
found in close association with insects, Pew species 
hve been found in au habitats, The species 
deseribed herein was collected in an anhydrobiote 
stile from dry sandy sor with fiithe organic matter. 
One other-species of Mexorhaldilis, Mo spiculigera 
(Steiner. 1946) Osche, 1952 has been reported lo 
survive anhydrobiosis (Sudhuus 1978), 

Osche (1952) subdivided the very large genus 
Rhabditis into seven subseneri, One of which was 
Mesorhabditis, with the type species Rilwalylitiy 
ypiculigera Steiner, (936. The taxonomy of 
Rhabditidie has been extensively reviewed by 
Sudhaus (1974, 1976, 1978) Whe has retained 
Mesorhabditiy al subgeoerne rank. Phis view was not 
supported by Andrassy in his autboriGiive 
mongemiph on the suborder Rhwbditina (Andrassy 
1983) in whieh he considered Meserhahditiv to he a 
sepunile venus within the Mesorhabditinue, a rank 
accepted Hi this paper Sudhaus (LOOT) was not, 
however, persuaded by Andrussys acuments (Mut 
Mosorhobditis has aenerie canking, The difference i 
niniking rests an the Gasonomist’s inclination towards 
Tuinpimee’ or ‘split’, 

Within the suborder Rhabditina, the combination 
ol charieters that distinguish Mesorhabditis area 
Monodelphie femme with the villva: Well posterior to 
the Hd-poral ob Mie body wid a ponited conical til. 
The male has long, more or less straight spicules that 
are Jistally Tised, The male bursa is peloderian wath 


“Diwewton oak Motiv aid Zogloy Nawtnilign Nabeul Onieersary 
Content ACE C200, 


pared borsal papitlie arranged in three groups, 
lypivally bwo preecloucal, five peri-cloaeal and three 
closer lo the ip of the tail expressed by The bursal 
formula (2+54+3). 


Material and Methods 


Several samples of dry sandy soil were liken with 
aeylindrical metal corer, 12.5 cm lou. 5 cin miternal 
diameter, close to and below a bluebush on 4 
November [984, The siunples were phiked in plistic 
bags and returned to the laboratory in Canberra, Ten 
days later subsamples of 5 2 were placed on tissue 
paper in fap water in Baermanmn funiels. Atier 18h 
the funnels were drained and the nematodes 
collected. From one subsample. Giken tron directly 
heneath the bluebush, fifteen specimens of a new 
species Of Mesorhabditiy were found Clopether with 
many other nematodes). "This species was nat found 
inainy of the other samples 

The specimens ol Mevorhabeditiy were fixed i Sti 
formalin and transfered ta 5% aqueous glycerol, 
whieh wis concentrated by evapetanon at 4a Cc, 
then mounted on slides in anhydrous glycerol with 
cover Slips Supported by ghiss heads and ringed with 
Cilyceel (Gurr). Drawings and ricusurements were 
made will a ediera ducida wdachient on a Zeis 
Ulfraphot micrpscope, 

Type material hus been deposited in the Nations 
Nematode Collectu|l (ANIC) atthe CSIRO Division 
of Entomolosy. Cunberr ACT. 


Mesorhahditis kinchegensis sp. nov. 
(MIGS |S) 


Healowpes 9 Kinehega National Park. NSW. 4oxy 
1984, ANIC Nematode Collecuorn side QOOQ00S , 


RO 


speeimen QO000007, 

Paratypes: 6 34,5 29, Kinchesa National Park, 
Axi 194, ANIC Nematode Collection slides 
QOOO006-12, specimens QOQDQ008- 19. 


Measurements: Table 1, Measurements in im. 
Deseripiian of Halewpe male 

Body cylindrical, slightly tapered at head, rather 
bluntly truncated ut hind end (Fig. 1), tail short with 
peloderan bursa (Figs 1.3). Cuticle finely annulited, 
lateral field appears as Uiree parallel lines beginning 
in cervical region and extending as far as tail (Pig. 3), 
Six offset, rounded, clearly separated lips, each 
bearing a prominent labial papilla (Pig. 5). Buecal 
cuyily cylindrical, without pharyngeal collar. glottis 
possessing minute denticles, probably two (Fig, 5). 
Pharynx with strong muscular corpus. slightly 
expanded af metacorpus. narrow — isthmus, 
sutrounded by nerve ring, valved pharyngeal bulb 
lerminuling in very short trilohed cardia. surrounded 
hy intestinal tissue (Fig. 2). Secretory-excretory 
pore. ventral, level with base of isthmus (Fig. 2). 
Intestine, initially filling pseudocoel. becoming 
compressed about halfway along body by gonad, 
followed by rectum opening at cloaca (Fis. 1). Tail 
short. sharply pointed. Single testis reflexed dorsally, 
leading to short vesicula semipalis and long yas 
deferens, Clouca surrounded by a peloderan bursa 
wilh 10 pairs of papillae arranged (24+5+3) (Fig. 3). 


W, L, NICHOLAS 


Bursal papillae not fused it base, short posterior pair 
curled over, Two long narrow nearly straight 
spicules, capitulum distinet, distally fused, lips 
abruptly angled ventrally at about 25° to the main 
part, just beyond & slight constriction (noteh) (Fis, 
4). Gubernaculut a short straight rod, Posterior 
deirids at level of middle of spicules (Fig. 3), 


Paratypes and other meles 
Measurements: Table |, 

All the inale paratypes closely resemble the 
holotype. The level at whieh the spicules fuse, about 
50% of ther length, can only be clearly seen by 
squashing and rolling the specimen under a cover 
slip, which renders the specimen useless us a lype 
specimen 


Fomale paraiypes 
Measurements: Table (. 

Female paratypes vlosely resemble males (Pizs 
6,7) except for reproductive organs and lait (Fig. 8). 
Homodromous ovary reflexed dorsally i nid revion 
of body. Uterus extending to just beyond short 
Vagina and vulva. One paratype female (Figs 6-8) 
possesses sperm i) a short transitional region 
between ovary und uterus and six developing eges. 
about (5 pin in diameter and varying from 15 - 24 
fin in dength, Amphid fovea, a minnte oval slit at 
base of lateral lips, visible only in this. paratype 


Varin 1. Measerements of Mesorbabditis kinchevensis speci, 


---—————————— CO _ocn— (“—_— 


Sex/Type Male/Holo Male/Para n=6 Femuale/Pura n=5 
Mean SD Range Mean 5D Range 
ee" 

Length 524 AK? 49.30) 432-533 543 7196 407 -H62 
Max, width 2k 40 3.83 25-34 AQ) 24)4 JAAS 
Bueeul cavity 15 15 2347 I-16 (5 151 [416 
Corpus Ol 54 1,27 S137 ahi] (51 56-60 
Phavyix Pa) 126 37 [1-140 [22 6.9] }1R-128 
Heal to nerve rine 92 72 11,25 5A-KA 7 6,22 61-70 
Head to secretory /eseretory pore 116 92 142 77-100 Kd 2010 55-13 
Head to intestine |? 138 5.01 132141 135 K OY 1al-2 
Heid to gonad flexure 224 mit) 19.85 [88-245 242 56,08 205-284 
Heit to vulva - 439 OG ATS 455 
Head to anus 4a) 454 IO AMW)-SO7 493 57.2! 455-518 
Gonad length 2Nh 278 13,97 268-295 AI [34.14 THOS 45 
Rectum length 34 3| 108 23-3 a2 6) BU ER i} 
Yul 22 23 34 IR27 ait 14.77 32.70 
Vulva to anus - - - 05 7.89 37-10 
Spieule Is 4h 4.48 415 - 
Guberneulinn Ih a Mala) [8-24 
Do Man's a (8.7 7 1.6 12.7-184) au 242 S427 
De Man's b i 4 37 3A | 4.4) 47 AAT 
Do Mais 238 aI I4 17-95 0 5 24) 47-145 
De Mais Vu e - hI 732 77-82 


_ CC rr  C ——n— 


ANHYDROBLOTIC MESORHABDITIS 


rye 


Wuillad lad 
aL 


Wha dat 
IAIN 


1 ets 50um 

2 2a 50um 
346. 10um 

5 se 107 


Figs 1-5. Holotype male. 1, Entire male. 2, Ceryical region. 3, Cloacal region with spicules, gubernaculum, bursa and bursal 
papillae and lateral line in lateral yiew, +. Spicules orientated to show fusion. 5. Head and buccal cavity. 


82 W. L. NICHOLAS 


female (Fig. 7). Tail conical and sharply pomted 
(Fig. 8). Vulva posterior and distance from vulva to 
anus about 1.5 x tail length. Lateral line marked by 
three incisures extending from mid pharyngeal 
revion to caudal region (Fig. 6). 


Differential diagnosis. 

Mesorhahditis spiculigera (Steiner, 1936) Osche. 
1952, 1s the only other species reported to survive 
periods in anhydrobiosis (Sudhaus 1978). [t has a 


world-wide distribution and has been reported from 
New Zealand but not from Australia. It differs from 
M, kinchegensis sp. noy. in possessing a longer 
narrower buccal cavity, fusion of the bases of bursal 
rays 4, 5 and 6 and the tips of the spicules, though 
notched, are not angled ventrally. The ratio of length 
to width of the buccal cavity in M. spiculigera is 
about 10: 1 (illustrated by Sudhaus 1974 Pig. 7). 
whereas in M. kinchegensis it is about 4 : 5. Two 
other species, M. ssunvoghi Andrassy, 1961 and M. 


WIHT 


Chic habdadanbuandd 


" 


6 50um 
7, ee FS um 
Qo Se es 50um 


Figs 6-8, Paratype female, 6. Cervical region also showing the three incisures of the lateral line. 7, Head. §, Posterior body 


showing reproductive orgiuns, 


ASTYOROBIOTIC MESORMARDYTIS u4 


Inmeuspiemlosa (Sehuaurmans Stekhoven, 1905) 
Dougherty. 1955, abso have a notch close Lo the tip at 
the spicules, bul antike 7. kinchevensiy, heir spicule 
Ups are nol angled ventrally beyond the notel. 

Andrassy (1983) provides a useful key to the 17 
species he recognises. a summary Of diagnostic 
Chiricters and references to taxonomic deseriptions, 
Several species, WM, oscher Osdtner in Osche, 1952) 
Dousherly, 1955, MW megectilis (Sudhaus. 1978) 
Andrussy: 1983. A frreveheds UxGener a Osehe, 
(952) Dougherty, 1955. A vennwagli, WL 
Jiglandicola (Puehs, (937) Dougherty, 1955, M7, 
saliasy anid MO irariiensis (Meyl, 1953) 
Douvherly, 1955 ci be clearly distinguished by 
havingoa shorter female wil so that the distinee (rom 
YUL Ta Uns iS TCH preater Hain the tail length. Up 
the new spevies (he distance ts only about 1.5 x the 
hal length, A vanely ol features distinguishes other 
species From Mo Afmohewensis, Ui WL aiethi 
(Sudhuas. 197%) Andirissy, 1983 the spicules tire 
uch Shorter (29 36 pon compared With 41-5) ain in 
Mo hinchevensiy) The bueewl cavily of At 
cnivonterplea (Sudtius, (978) Andrassy, 19X83 is 
asymmetric. the phanyin of dL erigenmearenniy 
(Khem. 965) Andiassy. 1083 is unusually long, one 
third ol body lenwth, Mesarabedins africans 
Andrassy, HY82 bos labial papitlie curved: inwaruds, 
Mo uffime (ROrner in Osehe, 1952) Dougherty, 1955 
has ported tips. MM. renoipreulim OsGrner in Osche, 
1952) Dourherty, 1955, MW. belari (Niven, 1949) 
Dougherty, [O54 qn A. Gtritensiy possess only 
Wine borsal niys, the middle group faving four 
Tasted OF fhe nmere tise) five. di uM yerteriee 
Dassonville &  Heyis. P984. deseribed by 
Dussonville & Fleyis (hO84) afer the publicaion of 
Andrissy’s monograph, the laueral line has five 
jneisures rather (hin the more typienl three, as in A 
Kineheversiy, Sudhaus (97K) tas ohserved 
uberralons in fhe tadl length ane) besa 
Muvidil spectinens bab the characters used) to 
disthiviish MM, Rineheveusys ate consistent mall ihe 
ype specimens deserthed in (hts paper 


Htabilar 

Soil around plik too. The type specimens were 
onlleeted ie diy sind ie ai anhydrabiotie stare fren 
AVOUT The roots a blchtish, Medrecie pyciniidiite, 


rays at 


in Kinchega National Park, NSW, Three males of the 
Mune species Were collecled by M. Hodda from a 
field ol lupins on The Soil Conservation 
Experimental Farm at Cowra. NSW. These ure in the 
ANIC. collection, Nematode Colleetion slides 
H001290, 0001295 ail 0001286 butare bot ieluded 
vs pantlypes, as they come from uovery different 
habitat and ure mounted on slides with several other 
species of nematodes, 


Disteibunon 
AU present the species is known from only two 
lowawlities in New South Wales. 


Discussion 


The type-specimens ol Mesamiabditiy Machevensis 
sp. ney. come front an atypical habitat for 
Mesorhabditis, namely, arid soil with tithe orgame 
Maller TH Kinchega National Park. although this 
speeies bas also been collected front aeccultival 
Jund, Kinehega National Park lias a very irregulie 
annual rain, averiing 235 rim, and tin smnual 
evaporacion rite of 2000 din, Teniperitures reach 
49" Co suniier aie fall to 0) Coin-winler, Most of 
the previously described species of Mesenhabedtay 
hive been found i cdlecomposiig organic nutter sueh 
as modldy or rotting wood, or Iiinis (Andrassy 
W983). Severdl others hiwe been found in close 
ussecnition wil imseety such as searabid) heeile 
lurve. Mevorhatilits meveehiliy wits ussucnied 
with hymenopteran nests CAndtassy 183) 
Mesorhabditiy sudhetusi has been reported tony soi 
CMndrassy 1983) and AY srriadtes tron fresh walter 
(Dassonville & Tleyns [Ondi  Wesuelinbeitin 
spicaliverd, the olher species kaowe to survive in 
awhydrubioses, Has been Toundh i roti: wood: and 
horse due. Ths ducer larvae were associated with 
dune beetles (Sudhans (978), 


Acknowledements 


Jauneratclal lo Ms J. Sit for collecting samples 
Hom Kinchesa National Park ane) Dr ML boda. for 
Inmaking specimens fram CSIRO] Nematolopy 
collection wadhible lar stay. 


References 


Anmmasny lh (lO84) A Tesontimic Review of the Suborder 
Khaliliiina (Nemarike Sacerentid (Gditmens de 
VOlhee de ke Researches Selentiique ah lechmique 
Coulee Hers Peis). 

JJASSONMIEDD, AF ok HBV SL C1984) Proshwaiter 
nenitades treme South Adrien 7. New pind Kineawn 
species ooullewted ott Skirmmerepruiy  Mietoria, 
Phyrophvlaenien Obs a, 


OSCE OG. FINS I Systematik Und Phy lerie den Cartan 
Kivibdity (Nem ALO) Mae Tb Avs SMD, 190 80 

SUD MS Wo CPE) fur Systoriatik, Verbotene, Okeloure 
und Bralosie Heuer amd weg bekunoler hibuitlen 
(Nermiohad 1, Teal dba POE S. 174. 122 

(19700 Sonim h hilonisele Rerinerkiiwen uber 

Arenund Gattungen der Uiiterkumtie Rhabditinae vertu 
five (Rhahditidae, Neroatodiy, Vewnrafagien 22, 4976, 


84 W. L. NICHOLAS 


= (1978) Systematik, Phylogenie und Okologie (1991) Check list of species of Rhabditis 
der holzbewohnenden Nematoden-Gruppe Rhabditis sensu lato (Nematoda: Rhabditidae) discovered between 
(Mesorhabditis) und das Problem, “geschlechtsbezogener” 1976 and 1986. Nematologica 37, 229-236. 


Artdifferenzierung. Zool. Jb. Syst. 105, S 399-461. 


FIRST QUEENSLAND RECORD OF THE BURROWING FROG 
CYCLORANA CRYPTOTIS TYLER & MARTIN, 1977 
(ANURA: HYLIDAE) 


BRIEF COMMUNICATION 


Summary 


During fauna surveys conducted in Cape Melville National Park (150 km north-west 
of Cooktown) and adjacent areas, new species and new records of vertebrates and 
earthworms were obtained’’. Following a thunderstorm on 21.xi. 1995, large numbers 
of frogs were found in a localised area (14°34°45’" S, 144°29°50°° E) approximately 
7-9 km west by road of the Wakooka Outstation. A call unfamiliar to me was in the 
large chorus. Observation revealed a species of burrowing frog of Cyclorana not 
recorded in Queensland. Thirteen males and one female were collected. The call was 
recorded, tissues sampled and photographs taken. 


Trimesaetions of te Reval Sorte of SM (199%) 122(2) KS-RO 


BRIEF COMMUNICATION 


HIRST QUEENSLAND RECORD OF THE BURROWIN 


~~ 
vi 


7 FROG CYCLORANA 


CRYPTOTIS TYLER & MARTIN, 1977 (ANURA : HYLIDAE), 


Durie Faun surveys conduerd i Cupe Mehatle 
National Park C150) kan ooriewest af Cookdsva) aad 
AUJUCONL reds: Hew species an new cecords of vertebrates 
und Garth woes were obiined!: Following a thunderstorin 
on 2bsr 1995, latee pubes oF frows were found in a 
localised aren ChE ANAS S, 144" 20550" LY approximately 
79 kny west by roud of ihe Wakooku Qutsntion. A vall 
Unhunilin tome Wee oy the hire chorus; Observation 
fevenled aospeeies of burrowing trae ob Cyclaraier net 
recorded in Queenshind. ‘Thirteen males aid ane fenuile 
Wer Collected. The cull was recarded, issues sampled ane 
pholowrophs taken 

Qn the basis of norphotopy and oll | tentatively denis 
the Hop us Cvefarenn eryptatiy, a seal burrowing Hag 
previously Known from northern Wester Aust and the 
Northern herritary 

Hadividiibs conform in colour and appearunve to the 
Veseription af) ceyproris Tyler aod Martin, wath) a bility 
mottled dorsi of slute, erey uml saloon with a ulistinet 
silo phsteorbital bur (Fie Li The appearence is 
Stokingly siovlin to the phitayeriph in Tyler ened, (O82. 1 
preservadive, (he salmon colon ranol was lost The raptial 
pads Were salmon aad ladecl in proservative. The ventral 
sirtaee wars white ind intles had a shite thro, The 
LWP wits Gevered Will skin i all animals. Toes were 
hall webbed wilt no expanded terminal dieses 


Nip t, Male Cvilera eryprens Trem west of Wakoaka 
Outsuien, Queershinel, 


The general appeaninge is of idiimpy ain fobust frog 
(Pip. 1). Meastiremerts, following previous faemliods . fall 
Within the tange ol oreplaris! Sioul vent length ranges 
from 357-459 nim, with the only female measuriys 
38, )mim, Legs wee short (PL/S-¥ 033-0039) yn the eve i 
naris distince is wremer Or Jess than the intenarial spar (LE 
N/IN O.955-1.141) (Table 1), 

Frogs Were Found Hi temporary, rere fifledl pools alone a 
draiuge Whe ta lows open woodkind on chu soils, The 
verehtion wis duminited by Aveldleuee steneisray lever anid 
Enealypras leprophieba (vip, 2 Alvitude was 40+ 1 in 

Call dination was 503 tasce witha pulse repetition rite at 
[13 pulsesssee ! anda dominiune frequeney of SOU He The 
pulse reperiiow Mite is lawer thin those rages al oe 
cryplotiy described im the Tilerature and Lake Argyle area 
(P83-193 pulses see 2o hin Nie Like Argyle Tourist 
Villugets 145-160 pulses see! [Go b> Watson pors. anit. 
1997}. lake Argyle 1a8 pulses see for holorype, Daly 
Watters) CRible 2), Tenmpenaures at the callie site were 
2H.2 CO (wae) and 27.2" C (ane) 

Cv lorie ceoypons wos calling while Coating in walter, 
When the voeul sae inflided the anterior halt ot the ives 
Wath Hfled, The inflotig and deletion of the yoeul sae 
cured the body ta rock ia Toner similar to Nerden 
delaneseaphuy vbserved elsewhere im Cape York, Arotter 
localities in its tange Co ervgetatis usually calls whilst 
Hoorn a water (GE Watson pers, comm, 1997) although 
the hololype ovis auling from dand OA. AL Martin pers. 
cam, 1997), The snite fenile was collueied Mowting in 
the water in ayilhiry aniplexus. Two other species ot 
Cyelarana. Co brevipes and CL pevertalteendion. entled 
froin ihe banks Ob pomls whereas Linaeedyre ster crctes 
called with Co erypfediy in the pools 

The presence of ©. erypreriy near Witkooku  O)uistutigns 
represents worminge esrenston oF YOO ken east ror prey tous 
foculity records obtuined by Davies. Tyler und Watson at 
Borroloola, Northern Territory (SAMA R43702, 16°12 8 
13637) EF). The intervening area accoss The gull pkainis fas 
hot heen exteasively sampled and additional populations 
may be vapeeted, The location of all records far € 
ervyprotis is tit band across nerthers Acstitia (ham Berhy, 
Western Australia to Wakooke Qutstigen mur Cape 
Melville between 1430'S gd [2 20'S, 

Other frogs collected with ©) cevyplons were 2 €. 
brevipes, Co unvertinftandin.  literia albounitata, by 
coerulea, b. ribelli, Lined ynasies atuerity, Noveteler 
melanoscauplies and Clperaleie miunula. 

Voucher specimens of Cerys celleered ai the 
Wakooka site ie tn the Atherton office of Deparment al 
Lovironmenteatlection Sos SN 30000, SN 7I0b8 20, 72074- 
25, 7245-36, 7204044 


Taki) Morphofogical meannrenents of dd Cycloranu cry ptotis fem west af Wakooke Quisterion. Queensland 


Abbrevintions fallow Tyler & Martin (1975), 


SV] TL/SVL HW/SVL- HW/TL RD/HW L-N/IN 
Rare A745 9 (1339-0), 39 036-043 (1924-1, 169 (1.26-0.343 955-1141 
Meun 49.5110 356 O.386 1 URS (). 2M 1.65 


RO 


% J Morphologically these specimens conform closely wath 

Pda | th : i ft C. ervplotis. However in view of the size of its range 

; j extension as well as the differences in cull structure, the 

% Ate identification should be regarded as provisional until 
iS Ae age! b= Substantiated by biochemical analysis. 

‘ Me ae Dr M. Davies, University of Adelaide und Dr A. A. 

= giorceeand ES, raat Martin, Royal Melbourne Zoological Gardens reud un early 


version of the manuseript and provided helpful comments. 
Assouile Professor G. PF. Watson. University of Melbourne 
provided call analysis dati. Field assistance was provided 
by M. Blackman. Q. Hart and R. Worall. C, Prith supplied 
the C. cryptotis photograph. The referees, Assovrate 
Professor M. J, Tyler and Dr A, A. Martin. made significant 
fe contributions to the manuscript. Alf uf this assistance is 
eratelully ucknowledged., 


Fig 2. Habitat of Cyclorana eryptoriy west of Wakooka 
Outstation, Queensland. Froes were calling from the 
Lermnpordiry pool. 


Taunn 2. Call variation within Cyclorana eryplouis. 
Superscript numbers in source colurmm refer to references. W - Water lemperature, A - air temperature. at calling sites, 


Source & val Locality Dominent Call duration No.of pulses Pulse see! Calls min! ‘Temperature °C 
sumple frequency — (milliseconds) 

This paper n=] Wakooka, Qld S00 503 58 113.3 TA 28.2 (W) 
G.F Watson Lake Argyle area. WA 920 439-455 65-74 {45.8 83.9 


(pers, comm. 
1997) n=2 


Holotype! Daly Waters. NT 1060 530 - 158 24,1 (A) 


Tyler erals n=l Lake Areyle area, WA &S0-1 100 330-348 61-70 183-195 36.9 (A) 


‘Jamieson, B. G. M. (1997) Mem. Qld Mus, 42. 233-270. Tyler, M. J., Davies, M. & Martin, A. A. (1982) Copeiy 


MeDonald, K. R. (1997) /hid. 42, 307-309. 1982. 260-264. 
‘Tyler, M. J. & Martin, A. A. (1975) Trans. Ro Soc S ‘Tyler, M. J. & Martin, A. A. (1977), Ree. S. Aus. Mus. 
Aust, 99, 93-99, 17, 261-276. 


K, R. McDONALD, Conservation Strategy Branch Queensland Department of Enyironment PO Box 834 
Atherton Qld 4883. 


THE PREVALENCE AND DISTRIBUTION OF NEMATODES IN 
THE LARGE INTESTINES OF SHEEP IN SOUTH AUSTRALIA 


BRIEF COMMUNICATION 


Summary 


Nematodes from three genera (Trichuris Roederer, 1761, Oesophagostomum Molin, 
1861 and Chabertia Railliet & Henry, 1909) have been identified from the large 
intestine of sheep in Australia’. However, there is relatively little information on their 
distribution and prevalence. Oesophagostomum venulosum (Rudolphi, 1809) and 
Chabertia ovina (Fabricius, 1788) are believed to be widely distributed, particularly in 
winter rainfall areas** and Trichuris ovis (Abildgaard, 1795) and T. skrjabini 
Baskakov, 1924 are common species in sheep and goats’. 


Transactions of the Royal Soci at S. Aust (V99R), 122(2 87-88 


BRIEF COMMUNICATION 


THE PREVALE 


‘EK AND DISTRIBUTION OF NEMATODES IN THE 


LARGE INTESTINES OF SHEEP IN SOUTH AUSTRALIA 


Nennitodes from three gener (| 7richwis Roederer, 1701. 
Qesaphagesionin Molin, (S61 und Chabertin Ruillier & 
Henry, P09) have been identified from the large mlestine 
of sheep in Austr’, However, there is rebitively: [ttle 
information oon ther distibuiton aml prevalenee. 
Oevaphaygestonman venitoxiun (Rudolphi, 1800) and 
Chabert ovine (Fabricius, L788) are believed to he widely 
distributed, puuticulaely ta winter cotlall qurcas>! and 
Tricity ovis (Abildpaard, 1795) and 7) wkrjabini 
Bushakoy, (924 dre comiion species in sluep gad goats’. 

Beveridge and ord surveyed the distribution and 
prevalenee in South Australia of the ceanonncally 
Mportint Uichostoiwylod nenkdades of sheep, which 
occur iy the stonmuch and small intestine. and reported that 
some were betrer adapted to hot dry cnavironments (han 
others?) denmtodes. in the Taree intestine were identified 
only ineidemtally. 

Ty the present study the prevalence und regional 
distr iion of nematodes of the lhirge intestine of sheep in 
South Australi was determined by examining freshly 
collected caecu and colons of 313 sheep collected from 116 
Widely distributed localities. (Pig, 1) from [991-)993, 
Information On farm minagement, age. sex or breed of 
sheep Wats Unavailable and) provision Far the possibiliy of 
seasonal yarhilions in infection was not possible in the 
collvetion of material Intestines Were opened. emptied. 
washed in water andl the mitcenal suclice was inspected for 
nemilodes whieh occasionally adhere ty (he gir wall A iin 
subsample of the Tatestinal contents was examined for 
Tre HeTalodes UsiNg Un OLVTpus stereo-microseone, 
The remaining contents were washed through a 670 urn 
sieve Nematodes were collected. fixed) in LO bulfered 


formalin und cleared Th lacloephenol Tor tend feution, 
Male and female nematodes were identified hy comparison 
with published deseriptions of species! Four tematode 
species from Uiree gener were found) (heir prevalence in 
three rain kul zones is shown in Table b. Ritty-cieht percent 
of sheep were infected with nematodes. The mean numbens 
wi rie OF burdens of adult nematodes and the number of 
localities from which each species was 
shown i Tuble 2 

Trivhiwis ovisand 1, skejahind were conn and witlely 
Wisttibuted with the hitter species more prevalent, Both 
ferodes aeenrred i 140 OF animals and in 7% al these 
cuses 7 oskryabind owas the predaumaitt speeies. 
Sihificantly fewer 20 avivowere found in the low: paint! 
cone (<250 nin than mares with higher raimtall yg = 41 
ps O02), 

Onophagestomion venilosunr wos (he most commen 
nematode reeovercd and was most previlent lin sheep fran 
wrens with more than S00 mit of rainfall Nematodes of all 
three gener occurred concurrently in only four animals. all 
from the highest rainfull gone (S500 i. Pourth stage 
larvae of. veniesin were the galy Timature henatades 
foundk these wer prescot in 38 animals from 24 foculities, 
V7owith un annul rainfall greater than 300i, Weller areus 
of South Australia are upparently more favourahle for the 
Iransmission OF O. verufestnr (han drier areas, 

Banks! detected ©. evive in chotable” siimbers tn South 
Australia forty vears qe but did not vonsider it ta be 
Importint. The intlience of highly effective unthetniniies, 
sitce (heir itroduetion ta fhe P9608, njay Have contributed 
tothe current low prevalence of CL avid Prialiity spp, are 
generilly regarded as harmless! bur lave been assocned 


fecovered une 


Tabi l. Prevndlence (% ) according to rainfall of neniatodes inte large imestine ap steep in South Austratia. 


Rainfall zone 


*urusile Overall 200 ~ 349m 350 - 499mm =S00r0 
Triclieis aves att 1 ais 28 
Trichawis \krjabini 32 42 a4 28 
Oevoplagestonuil 43 30) 29 SY 
venulosin 

Chubertia ivina 5 6 i 5 


TABLE 2. The mean diner uid range of nennitode burdens and the number of localities (in 250-349 1m 
(et), 350-499nn Oh), >SO00m (ao) rainfall zones) from which cach spevies wus recovered, 


Nematode species Mean 


Trichucls avis 7 
Hrichwvis skrjabini 1 
Vesophagestamin venulosum 15 
Chabertia ovina 4 


Range No ol localities 
i by v 
0-63 I 13 \5 
(60 alt) a 7 
(261 22 att 20 
10 4 6 4 


The tohal Tocalities tn cach rainfall gone - Gi) = 23, (hy = 4h ic) = 49 


88 


with deaths of sheep in Australia during drought 
conditions®. Oesophagostomum venulosum is non- 
pathogenic’. lmmature C. ovina cause intestinal damage 
during their development but adults are less pathogenic!. 
Natural nematode tmfections in sheep offen consist of a 
mixture of genera and species, some of which appear to 


have hittke effect on their own but may contribute to disease 
caused by more pathogenic species!, Although the 
parasites found in this survey are not considered to be 
economically important, the data supplement previous 
records of gastro-intestinal nematodes of sheep in this 


region of Australia 


a 
oO 


gy? 
Yoo 


ADELAIDE 


O 


Fig. |. Distribution of samples examined from sheep in South Australia. One to 4 sheep were examined at each locality 
Negative for nematodes (9, one nematode species LU, two species A, three species @, four specics MH, 


‘Cole, V. G. (1986) “Animal Health in Australia. Vol. 8, 
Helminth parasites of sheep and cattle” (Austratian 
Government Publishing Service, Canberra). 

?Forsythe, B. A. (1953) Aust, vet. J. 29, 349-356. 
‘Banks, A. W. (1958) Ibid. 34, 20-26. 

iBeveridge, I. & Green, P. E. (1981) /bid. 37, 141-142 


‘Beveridge, 1. & Ford, G. E. (1982) /bid, 59, 177-179. 
Goldberg, A. (1951) Proc. Helminthol. Soc. Wash. 18, 36-47 
7Herd, R. P. (1971) Int. J. Parasitol. 1, 189-199, 
‘Farleigh, FE. A. (1966) Aust. vet. J, 42, 462-463, 
°Goldberg, A. (1952) J. Parasitol. 38, 35-47. 


M. G. OTCALLAGHAN, E, OCKLESHAW and J. ALLEN, South Australian Research and Development 


Institute, 33 Flemington Street Glenside S. Aust. 5065. 


VOL. 122, PARTS 3 & 4 
30 NOVEMBER, 1998 


Transactions of the 


Royal Society of South 
Australia 


Incorporated 


Contents 


Martin, H. A. Late Cretaceous-Cainozoic palynology of the Poonarunna No. 1 
well, central Australia - - - - - - 

Kolesik, P. Rhopalomyia lawrenciae, a new gall midge species (Diptera: 
Cecidomyiidae) deforming leaves of Lawrencia squamata 
(Malvaceae) in South Australia - - - - - 

Kolesik, P. Dasineura wahlenbergiae, a new species of gall midge (Diptera: 
Cecidomyiidae) damaging shoot tips of Lense Stricta 
(Campanulaceae) in South Australia - - - 

Davies, M. & Watson, G. F. Developmental biology of Uperoleia a “Tyler, 
Davies & Martin, 1981 (Anura:Myobatrachidae) - -  - 

Davies, M. & McDonald, K. R. A new species of frog (Anura: Seats 
from Cape Melville, Queensland - = - 

Davies, M. & McDonald, K. R. Developmental biology of Cnanise ei 
Davies, Watson, McDonald, Trenerry & Werren, 1993 
(Anura:Myobatrachidae) - oe M > 7 

Coleman, P. S. J. Changes in a Mangrove/Samphire Community, North Arm 
Creek, South Australia - — - - - = - - 

Smales, L. R. New species of Seurechina (Nematoda : Seuratidae) paras 
in dasyurid marsupials from Australia - - - 

Ferguson, M. A. & Smales, L. R. Spiroxys chelodinae Berry, 1985 (Nesumade: 
Spiruroidea) and Camallanus chelonius Baker, 1983 
(Nematoda: Camallanoidea) from freshwater turtles 
(Pleurodira: Chelidae) in Queensland, Australia - - - - 


PUBLISHED AND SOLD AT THE SOCIETY’S ROOMS 
SOUTH AUSTRALIAN MUSEUM, NORTH TERRACE, ADELAIDE, S.A. 5000 


89 


139 


185 


TRANSACTIONS OF THE 


ROYAL SOCIETY 
OF SOUTH AUSTRALIA 


INCORPORATED 


VOL. 122, PART 3 


LATE CRETACEOUS-CAINOZOIC PALYNOLOGY OF THE 
POONARUNNA NO. 1 WELL, CENTRAL AUSTRALIA 


By HELENE A. MARTIN® 


Summary 


Martin, H. A. (1998) Late Cretaceous-Cainozoic palynology of the Poonarunna No. | 
well, central Australia. Trans. R. Soc. S. Aust. 122(3), 89-138, 30 November, 1998. 
Palynomorphs found in Late Cretaceous-Cainozoic sediments are described. The 
Winton Formation yielded the Cenomanian Plicatella distocarinata Zone, but the 
uppermost part contained an equivalent of the late Paleocene Lygistepollenites balmei 
Zone, showing it should be reassigned to the Eyre Formation. The Eyre Formation 
also includes sediments that are an equivalent of the mid Eocene Lower 
Nothofagidites asperus Zone. An abundance of Asteraceae and 
Chenopodiaceae/Amaranthaceae pollen in an assemblage at shallow depth is thought 
to be Pliocene-Pleistocene in age. 

Key Words: Central Australia, Palynology, Late Cretaceous, Tertiary, 
Palaeovegetation. 


Transactions of (he Royal Secrery of S Aust (1998), 12203), 89-138. 


LATE CRETACEOUS-CAINOZOIC PALYNOLOGY OF THE POONARUNNA 
NO. | WELL, CENTRAL AUSTRALIA 


by HELENE A, MarTIN® 


Summary 


Marvin, HAL (1998) Lite Cretaveous-Cainosoie palynolowy of the Poonarunna No. | well. central Australia. 
Trans. R. See, 8. Aust. 1223), 89-138, 30 November, 1998, 

Palynomorphs found in Lite Cretaceous-Cainozole sediments are described. The Winton Formation yieldeal 
the Cenomianian Plieaella distocurinate Zone, but the uppermost part contained an equivalent ol the late 
Paleovene Lygivtepallenites babe’ Zone. showing it should be reassigned to the Eyre Formation, The Eyre 
Formation also includes sediments that are an equivalent of the mid Eovene Lower Nerhefiigidites ayperus Zone, 
An abundance of Asteraceae and Chenopodiaceac/Amuaranthaceae pollen in an assemblage at shallow depth ts 
thought fo be Plioacene-Pleistocene in age. 

The vegetation of the late Paleocene and auddle Rocene was mainly forests with minor herbaccous SALT 
communities, Gymnosperm pollen dominated the lute Paleocene palynoforis and proteaceous taxa were very 
diverse. Pollen of Cononiaceae/Elaeocarpaceae is moderately common and there is a wealth of a@iospern 
pollen, Inthe mid Rocene, pollen of Arauvariaceae, Casuarinaceae anda lithe Norhofages were dominant and 
there Was a great diversity of angiosperms. The Plivcene-Pleistocene palynofloras haye a limited diversity with 
pollen of the herbaceous/shrubby Cyperaveae. Poaceae, Asteraceae and Chenopodiuceae/Amaninthacesae 
Jominunt, Cusuuriniceae and Myrlaceae are the only likely trees and there is relatively lite pollen of these 
families, henee the vegetation was open shrublands similar to that found in the region today. There are, however, 
some dispare Gixa inthis Pliocene/Pleistocene palynoflora that are unknown in the wrid region today. 


Key Worps: Central Australia, Palynology. Late Cretaceous, Tertiury, Palaeovegctation. 


Introduction 
1dr an 


Taz . ey 4 @ Bore of this study 
This study of the Dethi-Santos-French Petroleum Boles hiportad BV POONARUNINA No. | 


Co, (Aust.) Poonarunna-| well was undertaken inthe Sluttet (1291) 
hope that ir would shed some light on the evolution # Macrotiossil site 
of the arid flora and vegetation. The location of the 
well, northeast of Lake Eyre. and within the first 
sand ridges of the Simpson Desert (Magnier'). is 
strategically placed for this purpose (Fig. 1). 

Finding preserved pollen in wrid/semi-arid regions 
isd major problem, Preservation requires anaerobic 
conditions in permanent likes. swamps, bogs ete, 


Once the climate becomes dry. these permanently- Soh \ Peachayarina 

wel sites disappear, Alternate wetting and drying in wes _. Branohe 
seasonal swamps and lakes destroys petlen. Rulon “3 

Moreover, when pollen ts deposited in permanently- : MW MAREE - L Callabora 


wel sites, 1 must he hunied deeply enough to escape 
the effects of a later fluctuating water table of 
future drier climate, HW iLis to reniin preserved. Deep 
weathering has undoubtedly destroyed much of the Vig 1 Locality map. 

pollen record, but where pollen has been preserved, 

Hood pulynofloras were recovered. This paper Cretaceous sequences intersected ii (he Poona 
documents (he pollen species recavered and reports runna-l well 

on the palynostratigraphy of the Tertitry and upper 


Geology 
‘ ) Wowie! Science. Universi of New S Wale , : 
“ae font WNcience, Mniversy OE Nery, Sault Wales Poonarunna-f was sunk to 1,696 im with the 
SVU ONY Moe. , * - . \ 
Manet 1 Heol) Well gumptetign report el! Poonmuner Ni | Intention of exploring the Palavozoic or Proterozoic 
SOU AT SLT Cupid basement strait beneath the Permian or Mesoaoie 


on) HA MARTIN 


cover beds. The well established a good reference for 
Mesoaore studies (Magnier), bul there was no 
jnterest in the Cuimozoie. The present study 
concentrates on the Tertiary palynostrutmsraphy tert 
includes Che uppermost of the Creticcous which 
vsuiblisbes the Tertiary/Cretaceous boundary, 

In ihe well completion report (Mugnier'), the 
following units ure defined from the lithology and 
correlation ol aaimmieriy lows with supplementary 
information fran sonie Jogs und resistiviry. The 
‘Tertiary and Quaternary were defined as consisting 
of Quaternary sand and alliviun (025.9 m) and white 
dolomite limestone interbedded with grey murky. 
sandy clay (5,.5-95.4 mi). The Lite Creticeaus 
iConamanian) (95,4-667,5 m) wis dedined as the 
Winton Pormution, consisting of alternating grey lo 
dark grey silt, fissile Shale und siltstone wih 
disseminuted pyrite. The present study records the 
palynology of the upper 146 m only, Le. the 
Calnozoie and the upper part of the Late Cretaceatts 
Sequence, 

The upped part of the Winton Formation examined 
in (his study is remarkibly Upiforn in appearance. 
From the palynolowy, the Cretiecous- Tertiary 
bonndivy occurs at about 110m. und the interval 
from 95-110 ti Should be reassigned to the Eyre 
Bormiation, since the Winton Formation elsewhere mW 
the Luke Eyre Basin js cortelated with the upper 
Phimepollenites, pannesus and Plieutelhe dixte- 
corinatis Zones. of Albin te Cemomaniin age 
(Krieg epal 1991), 

The Eyre Formation (Callen et af (995) is a 
widespread and distinctive fluvial to fluviokicustrine 
sund unit, but Jithologieal variations oveur i some 
chunnel facies. Plant fossils are characteristic. cmd 
Jocally abondint, Caurbonaceous horizons wat the 
Eyre Formation contain spores, pollen and a few 
dinoflagellates of late Paleocene to middle Bocene 
age. The Byre Formation was deposited in large 
menndering and braided streams (srieg era! T8997), 

The unit between 5.5 and 95.4 mis correlated with 
the Etadurina Mormation of probable law Oligocene 
(6 Pllagene ave. hut it ms most Tikely largely carly 
Minveng, Phe Bladen Formation was deposited i 
a evaporative food plan-hicnsting environment, 
vader ad clinute ner (han thator the Lyre Parmation 
(Knee eral WO]: Callen er al, T9YS), 


Matertals aod Methads 


Only curlines Were available for the present study. 
The possibility of conmbiminatiol is greater with 
cuuiags. bul avilh proper drilling and sanmpling 
provedures. reliable samples may be oblained. This 
topic as fully discussed by Martin (P98) In the 
present Sty, rhe Late Cretaceous assemblages 
conluincd véey few ‘Tertiary grains und barren 


samples in the Camnozoic section suggest thal 
coptimination is niinimnal anc these samples are 
reliable. 

The samples were freated with hydrochloric and 
hydrofluore acids to remave mineral matter. 
Controlled oxidation with cold Schulze solution and 
polussiuin carbonate was used to eloar the residues. 
which were then mounted jt elycerine jelly (Brown 
1960: Ciray 1965). 

The palynofloras were assigned to vores using (he 
ranges of diagnostic species. The Tertiary palyuo- 
floras were quantihed with counts.of about 150-200 
erains and percentages were bused on the total pallen 
count. The pollen spectra derived from the counts 
provide a hasis for interpretations af the palsen- 
verotillon, 

An assessment of the abundiiee ab micposcapie 
curbouted particles in the Tertiary ussemblases was 
mide. The formation of these particles t) conlro- 
versial, When they are found in mid-late Tertiary and 
Qualernury sediments, it is generally accepted thal 
they were formed from burning aid we ehareod 
particles (Luly era/, L980, Martin 1987; Kershitw ef 
a. W901). On the other hand. Sehopf (1975) has 
suvacsted that black curbonised particles huve 
formed by oxidation at the surtace OF swans 
However. there have heen numerous studies which 
shaw hit chareoal may be distinguished from other 
black varbonised materiul (Scouw L989; Cohen & 
Spach 1977, Sander & Gee 1990) and chureoul 
muy be found in sediments of any age, 


Palynostraligraphy 


hale Crehiweods paulynosteytigraphy is based on 
tharol Helby etal, (1987), the systematic paly nology 
is presented in Appendix |. the species identifedt ae 
given in Table band the sires of diagnostic speeres 
in Preute 2, 


1/0 146m, Pligatelly distocarinita Zane. 
Comonentan 

The assemblages in this zone lack the distinerin 
Phyllocludidies meaysani’ and Prafeuctlies spp. ol 
the Romans? Zone Plroytells distucertnade acl 
Trilobesporites irierereu/ans. whnse ranges gral 
with the 2 omewsenin Zone are presenr These 
assemiblawes thus fit the 2 disrecartuna Zune of 
Cenoommniun wwe (Fig, 2), Addsapallis enicifarns is 
usally present in the Cenomanint it the Luke lyre 
Basi (N, F Alley pers, comut 1995), bot it has for 
heen revarded tronr these assemblages, Burger 
(1993) reports (hit Foreriimiyprads daily, bas nel 
been recurded trom the Po diareeerinue Zone, bur i 
ispresent ip (he sample fram LLS-119 mob this study 
(Table |). 

Spores ol ferns, tycopods and bryophytes: are 


PALYNOLOGY OF THE POONARUNNA NO. | WELL 


‘TABLE 1. Late Cretaceous species identified from Poonarunna-1. 


+, present. ++, common. 


Depth (m) 


Spores 

Aequitriradites spinulosus 

A. Verrucosus 

Baculatisporites comaumensis 
Balmeisporites glenelgensis 

B. holodictyus 

B. tridictyus 
Camarozonosporites australiensis 
Ceratosporites equalis 
Cicatricosisportes sp, of Burger 
Cicatricosisportes spp. 
Clavifera triplex 
Crybelosporites punctatus 

C. striatus 

Cyathidites australis 

C. minor 

Dictyophyllidites sp. 
Foraminisporis dailyi 

F. wonthaggiensis 
Foveogleicheniidites confossus 
Gleichentidites circinidites 
Laevigatosporites ovatus 
Microfoveolatosporites canaliculatus 
Ornamentifera sp. cf. O. Sentosa 
Perotrilites jubatus 

Plicatella distocarinata 
Polycingulatisporites sp. 
Reticuloidosporites arcus 
Retitriletes austroclavatidites 
Ruffordiaspora australiensis 

R. ludbrookiae 

Sestrosporites pseudoalveolatus 
Stereisporites antiquasporites 

S. pocockii 

Stoverisporites Microverrucats 
Trilobosporites tribotrys 

T. trioreticulosus 

Triporoletes sp. ct. 1. simplex 


Gymnosperms 

Alisporites sp. ch. A. grandis 
Araucariacites australis 

Corollina sp. cf, C. classoides 
Ginkgoeycadophytus nitidus 
Microcachryidites antarcticus 
Podocarpidites ellipticus 

P. exiguus 

Podosporites sp. 
Trichotomonosulcites subgranulatus 


Angiosperms 

Cupuliferoidaepollenites cf. C. parvilus 
Dicopopollis sp. 

Foveotetradites fistulosus 


110- 115- 128- 134- 143- 
113 119 131 137 146 
+ + 

+ 
~ + + 
+ 
+ + 
+ + 
+ + 
+ + 
+ oo 
+ + 
+ + 
+ 
+ 
+ + + + 
++ ++ +e ++ ++ 
+ + 
a 
+ + + 
+ + 
ae + + ++ + 
++ - ++ 
+ 
+ 
+ + + 
+ + + + 
4 
+ + 7 
+ 
+ + + 
+ 
4 
+ + + 
+ 
+ + + + 
+ + 
oo 
+ + + 
+ + + + 
+ + + + + 
+f + + + 
+ 
+ + + + + 
+ + + + + 
+ 
++ 
++ + 
+ 
+ 
+ 


92 HLA. MARTIN 


Manne t continued... 
Depth (in) atte 115- 28 1ad- \43- 
13 119 al 137 lao 
Lihtucidites sp. et. (. Kattangutaensts + 
Lillavidites sp. + 
Phinopollenites augathellaensts ' + 1 
PL panninsas t + + 
Seroeloretrdi(es Varirenicilags t 
Tricalporites sp. ck To uparyexiiuts + 
Alpae 
Roryocncous briaait + 
Hovologinella sp. + 
Lecuniella sp. + 
Saeplodininim gravattensix + + + 
Sehizosporis relicnlatus + t 
Acritarch sp. inde. + 
ACE ZONE DIAGNOSTIC SPECIES Phimopollenites augathellaensis, Po pennosas and 
Toei: [asameeaen - Foveotetradites fistilosus being the most common 
MEDCENE | tae ct (Table 1). The angiosperm palynoflora may he 
swasrrcarn | (Tne placed in Suite IH, of Lite Apuan-Early Cenomanian 
‘ 4 age (Burger 1990), for it lacks the triporate forms of 
& Suite 1V which starts within the Cenomaniin (Burger 


Tavinrilinenites 


CAMPANIAN, 
Nevtrcohaypeditess 
SONIBLILE 
ur i 
= 
oO | 
4 Theoiporites t 
FE | pawtouan ones 


CONIACIAN | py jyettnios 


Hine ve 


sis 


TURONIAN 


TEM a wes 


3 
nae 3 
Ele 15 
CENOMANIAN in| ra 
By a 
E z 
2 
ALBIAN Phimopoienies s 
a 


PAGS 


Fig. 2. 


Ranges of diagnostic Cretuceous species, from 
ie & 


Helby ef al, (1987), 


particularly diverse and abundant in this zone, 
Cyathidiies miner is very common in all of the 
sumples and Kuf/ferdiaspera/Cicatricosisporites 
spp., Foruminisporis wenthagelensis, Gleiehe- 
niidites cireinidites, Laevigutesporites evetus and 
Srereisporites  untiqitasporites are sometimes 
abundant, Gymnosperm pollen is diverse and well 
represented with Podosporites spp. common in some 
sainples, Most of the trees in the palacovegetation 
would have been species of gymnosperms, There 
is a small angiosperm pollen content with 


1993), 

The five sequential samples ure essentially the 
sume age. but there are quantitative differences. The 
three oldest assemblages are diverse with a good 
representation of gymmnosperni pollen, but the 
angiosperm content is restricted in diversity and 
abundanee. The abundance of gymnosperm pollen 
suggests that forest would hive have been a major 
part of the vegetation. The assemblage at 115-119 m 
has fewer gymnosperms than the other assemblages 
and an unusual abundance of Srereixporites 
antiquasporiies, with affinities to Splegitien, and 
Foraminsporis wonthaggiensis, similar to the living 
hepatic Nothylas bruetelii (Dettmann 1963). This 
assemblage suggests that there were extensive bogs 
or Wetlands and fewer forest trees in the immedite 
vicinity. The uppermost assemblage has a diverse 
gymnosperm palynoflora, indicating forest similar to 
that of the oldest assemblages, and an inereased 
angiosperm palynoflora. The megusporangiate water 
ferns, Balmeisporites spp., are also well represented 
at this level, 

The microphinkton content of these assembliges 1s 
low with only a few forms represented, Lecaniella 
sp. and Sehizesparis reticulaiuy have probable 
affinities with the Zygnemetaceae (Grenfell 1995), a 
family of filamentous green algae usually found 1 
shallow, flowing fresh water. Berryececcus braunit 
may be found in fresh and brackish water (Pentecost 
1984) and Saeptodinium vravatiensiy is a tresh water 
dinoflagellate (Harris 1973), 


PALYNOLOGY OF THE POONARUNNA NO. | WELL 


TABLE 2. Jertiary species identified in Poonarunna-1. 
Percentages of total pollen count are given. +, present but not counted in the counts of 150 or more grains. 


Depth (m) 6- 73- 
9 76 

Spores 

Agalla sp. + 

Camarozonasporites amplus 

C. bullatus 

Camarozonosporites sp. 1.2 

Ceratosporites equalis 0.6 

Cyathidites australis 0.6 

C. paleospora 1.3 5.4 

C. splendens + 

Dictyophyllidites concavus 8.4 

Foveotriletes lacunosus 

Gleichenia circinidites 18 

Grapnelispora evansii 

Laevigatosporites ovats 1.2 

Polypodiacoisporites sp. ct. P. relirugatus + 

Poalypoeliidites spp. 0.6 

Retitriletes austroclavatidites 

Stereiyporites sp, 

Todisporites sp. 

Triletes sp. ct. 1) tuberculifarmis 1.8 

Triporoletes reticilanis 0.7 + 

Verrucosisporites Cristatus + 

Unidentified 0.7 

Gymnosperms 

Araucariaciates australis 1.3 14.4 

Cupressaceae/Taxodiaceac 0.6 

Dacrycarpites dustraliensis 

Dilwynites eranidatus 1.2 

D. tuberculanis 0.6 

Lygistepollenites florinii 8.4 

Microcachryidites antarcneus 

Phyllocladidites mawsonii 0.6 

P. retienlosaccatus 

Podacarpidites spp. 0.7 4.2 

Trichotomonosuleatus suiberanulatus 

Angiosperms 

Acaciapollenites inyriosporites 2.0 

Aglaaridia qualamis 

Amosopolliy dilwynites 

Arecipiles sp. cl. A. minutiscabratus 

Austtalopollis obscurus 

Beaupreaidites elegansifornus + 

Chenapodipollis chenopodiaceoides 19.2 

Cunoniaceae (tricolporate) 0.6 

Cunoniaceae (bicolpate) 

Cupaneidites orthothechus 0.6 

Cyperaceae 10.6 1.8 

ef, Dodonaea 0.7 

Rlaeocarpaccae 

Evicipites crassiexinus LS 


Gothanipallis bassensis 
Graminidites monoporites 73 


05 
94. 97- 1O1- 104- 107- 
97 101 104 107 110 


(16 


1.3 
0.6 
0.6 

1o 

1.3 
7.2 

i) 
99 


24.3 
1.9 


5.9 


34 


19 


1,9 
0.6 


0.6 


i=) 


=e NS Ke 
3 nt 


m= te be by 


S 


2 
2 


3.0 


0.6 


0.6 


“a 
rm 


0.6 


13 
0.6 


0.6 
0.6 


3.9 


0.6 


0.6 


94 
TABLE 2 continued... 


Depth (m) 


Haloragacidites haloragoides 
H. harrisii 
Hexpollenites austroclavatus 


Lewalanipollis cf. L. rectomarginis 


Lewalanipollis cf, Persoonia 
Liliacidites lanceolatus 
Malvacearumpollis sp. 
Malvacipollis diversus 

M. subtilis 

Milfordia homeopunctata 
M. hypolaeniodes 
Myrtaceidites eucalyptoides 
M. parvus 

M. verrucosus 

Myrtaceae unidentified 
Nothofagidites emarcidus 
N, deminutus 

N. faleatus 

N. vansteenisii 
Nuxipollenites kempii 
Polvyorificites oblatus 
Polyporina granulata 
Propylipollis ivanhoensis 

P. latrobensis 

P. ef, pseudomoides 

Pct. P. reticuloscabratus 
Propylipollis sp. 
Proteacidites adenanthoides 
P. adenanthoides/crassus 

P. angulatus 

P. annularis 

P. cooksoniae 

P. crassus 

P. fromensis 

P. grandis 

P. incurvatus 

P. sp. cl. P. ineurvatus 

P. ef. obseurus 

P. cf. slipplatis 
Proteacidites spp. 
Quintiniapollis psilatispora 
Rhopites alveolatus 
Rhopites sp. cf. R. alveolatus 
Santulumidites cainozoicus 
Sapotaceoidaepollenites rotundus 
Simplicepollis meridianus 
Simpsonipollis sp. 


0.7 


Sparganiaceaepollenites barungensis 0,7 
Tricolpites sp. cf. T. asperamarginatus 


T. sp. cf. T. confessus 
T. sp. ef. T. discus 

LT. phillipsti 

T. thomasti 
Tricolporites angurium 
T. lewray 


H. A. MARTIN 
7T3- 94- 
716 9O7 
12.0 3.9 
0.6 
+ 
0.6 0.6 
+ 
0.6 
1.8 
0.6 13 
+ 
L.8 
%Q6 
+ 
0.6 
0.6 
1.8 
5.3 
+ 
+ 
3.3 
+ 
13 
10 
+ 
0.6 0.6 
2.4 18 
19 
+ 
+ 
0.6 
0.6 1.3 
+ 
+ 
+ 
+ 
+ 


4.8 


0.6 
3.0 


0.6 


101- 104- 107- 
104 107 110 
3:2 2.0 3.9 
0.6 
4 
0.6 1.3 
0.6 0.6 13 
0.6 2.0 
1.9 0.6 
+ 
0.6 + 
0.6 0.6 
0.6 
13 2.0 La 
0.6 
+ + 
0.6 0.6 
13 1.3 1.9 
1.3 + + 
+ 
0.6 
70 3.3 19 
0.6 0.6 
+ 
+ 
0.6 
a 
1.9 2.6 0.6 
+ 


OG 20 


PALYNOLOGY OF THE POONARUNNA NO. | WELL 95 


TABLE 2 continued... 


_ ---—--- On —————— ——  SsSsSseSsSeSFSFSFSFFFFFssFFFSFFFsffsseF 


Depth (m) 6- 73- 
9 76 


Triculparites substriatus 41, paenestriatnus 
Tricolperopollenites endobaltreus 0.6 
Trierites minisculits 

Triporepallenites ambignis + 
Lubulifloridites untipodica/simplis 27.8 
Unidentified angiosperms 79 6.0) 


Alpae 

Borrvocwecus ++ 
Debarya + 
Morkallacysta pyramicdelis 

Pediastram + 


Summary of major pollen groups 


Spores 2.0 21.5 
Gymnosperms 14 29.9 
Casuirinaceae 5.6 12.0 
Myrtaceae 3.3 2.4 

Cunomaccae/Elaeoearpaceae 

Nothofagus 10.8 
*Proteacede” 3.0 

Asteraceau 27.8 

Cyperaceae 10.6 1.8 

Poaceae 7.3 

Reshonaceae 2.4 

Sparganiaceae 0.7 0.6 

Chenopod type 19.2 


94- Q7- LO1- HO 107- 
oT 101 1o4 107 110 
2.4 
+ 
2.5 9.1 13.6 15.0 143 
+ + + 7 
+ 
+ 
92 O7 Wd [1.8 3.7 
Si3 51.5 Cama) WS 42.5 
3:9 3.) 42 2.0 3.9 
La 0.6 20 
54 24 6.4 13.2 5,8 
12.4 144 12.8 9.2 8.5 
1.9 24 13 0.6 
1.2 13 2.0 13 
[.3 1.9 3.3 0.6 


TT 


These Cenomanian palynofloras are generally 
similar to those of Bathurst Island, northern 
Australia, described by Norvick & Burger (1975), 
except that the former contains  freshwiater 
microplankton whereas the latter has a very diverse 


marine dinoflagellate flora. Norvick & Burger 
oecurrences of 


(1975) illustrate the known 
Cenomanian palyoofloras and, except for three non- 
marine localities in the Eromanga Basin, all the 
others are around the northern, western and southern 
periphery of Australia. The stratigraphically 
important species of the Cenomanian have been 
studied. but other than this study, the report by 
Narvick and Burger (1975) is the only Australian 
report to document all the palynomorphs in Ceno- 
mantin assemblages. 

Tertiary palynostratigraphy follows Stover und 
Partridge (1973), Macphail (1996) and studies in 
Central Australia (Krieg ef af, 1991, Sluiter 1991; 
Alley ef al. 1996; N, F. Alley pers. comm. 1994). The 
systematic palynology is given in Appendix 2, the 
species identified in Table 2 and the pollen diagram 
in Figure 3. The definitions of the pollen groups are 
given in Table 3. 


Taser 3. Definition of the major pollen groups used in the 
pollen diagram, Mig. 3. 
A tut list of taxa is presented Appendix 2. 


Name on diagram Taxa included in the group 


Spores All spore taxa 
Arauicariaceae Arancariacties australis 
Podoearps Padocarpidites spp. 
Lagvarestrobes Phylloclaclidites mawsanti 


Dacryelisn 
Other gymnosperms 


Lygistepollenites flurinii 
All other taxa under 
gyninosperms 


Cunoniaceae Cononiaceae and Elaeoearpacede 

Casuarinaceae Haloragavidites harrisii 

Myrtaceae All species of Myrtaeeielites 

Nothofagus All species of Nothofagicites 

*Proteicene” ALL species of Beaypreaidires, 
Lewalanipollis, Proteacidites and 
Propylipallis 

‘Callitriche” Anstralapollis obscurus 

Cyperaceie Cyperaceae 

Poweoae Graminidites monoporites 

Asteracene Tuhuliflovidites spp. 


Chenopad type Chenopodipollis 


chenapodiaceatides 


ey HA, MARTIN 


Samples examined 


MAJOR POLLEN GROUPS 9? 4) & jotal pollen count 


ZONE AND 
AGE 


LAKE FROME SURFACE POLLEN SPECTRUM 


POONARUNNA No.1 WELL 


Acacia 


seen joses of s es 


5 


1 eeee 
Onher 
aCage 

a2 


ahacese 
Gymnosperns 


Pagocspus 
Daenydiun 
Cunonmacsaey 


Elagsocar 


Lagarosrrobios 
Casuarnnace: 


Fi. 3. Pollen dis 


Pilocene-Pleislocane 


Ss 
SS 
SS 
IS 


SS 
S 


NSS 


SSVOV 


SS 


SSS 
SSS 


Ys 
Y 
oy 
% 
wy 


Barrer 


Barren 
Upper L.baimer equiv 
Lala Paleaoene 


BeaA 


P aistacarintata 
Cenomanan 


Cyperaceae 
Astirat 


Crenapod type 


ucam. For definitions of the pollen groups, see Table 3. The surface pollen spectrum from Lake Frome 


(Shutter & Kershaw 1982) represents the present vegetation ined is added for comparison, 
J 2 


WTO om. Upper Lygistepollenites balmei Zee 
equivalent, later Paleocene 

Gymosperoy tax are diverse and their pollen may 
constitute mare thin half the count (Pig. 3. Appendix 
Ay. Podocarpus (Podocurpidites spp... ts the most 
common. and Lagerestrobos (Phyllaclaciidites 
mawseail) is well represented. Pollen) trom the 
unviosperm taxa Cunoniaceue/Elacocurpaceae. 
Casnurimuceae (Heloragacidites harrixit) and 
Myrlaceae (Myriacidites sp.) is present but that fron 
Norhafirei (Nothofigiditey) has not been detected in 
this study. although is is usually present bul 
extremely rare tn this zone (N, FL Alley. pers. eonm, 
N04) There is a profusion of the proteaccous lax 
Bedipreidiies, Lewelanipaltis, Prapilypelliy and 
Profeacidites spp, (see Appendix 2) with 
Protedidites cadenanthaides, Po cana PF 
crass, 2 frameusix, Po cookseniae. Po leduhtonit ad 
Poreflesus the most common, OF Cellfirfehe 
Wustalopollis obsciris), Restionaeaw GWilfontia 
spp.) and Cyperaceue (Cyperaceaepalliy sp.) are 
present alsa. There is i number al anidentuhed 
digiospermn pollen ivpes present. and while they mary 
not be miportane for swatieraphie Consideration, they 
Indivale wrieh and aibunckint aneiospeon element i 
the hue Paleocene, The content of curbortsed 
prirlicles ts bas 

These asembliges aee usstened to the Upper L 
baline! Zone equivalent. The oceurence wf 
Cihapirelspert evitsee ied Trinealpits eho- 
confessus, whose vanges end ab the top af the 


Maastrichtiun (Helby et al. 1Y87) ts anomalous. but 
they may be reworked. Other distinctive taxa usually 
found in the Muustrichtian are lacking and 
quantitatively, these assemblages are unlike those of 
the Maastrichtian. 


Comparisons 

When compared with the L. Meliner Zone of the 
Gippslind Basin, the stratigraphic ranges of some 
diagnostic und churucleristic speaies are not the same 
in the two regions. For example, Proteccidites 
erassus, Po lelehionii and Po reflevtis. present in 
Poonarunna hegin their ranges in the Hocene of the 
Gippstind Basin and PB framensis in Poonarinni his 
not been reported from the latter site (Stover & [yin 
19732 Stover & Partridge 1973). In contrast, 
Cunonidceac, Cyperuceae and Milfardia lypel- 
cqenrodes OF this study are not found in the Gippstind 
Basin. The rich and abundit profeaceots tori are a 
feature Of The assemblages i central Avstraha bat 
they are “heither particularly diverse nop partiectatly 
common im those of the Gippsland Basin (Stover & 
tvans 1973. p. 59). Phere ure similarities. however, 
Hat “vinnosperm Lisa aire diverse aad common in 
both localities, although the Hontinite species of the 
gone. Lo belmer, as not been found to this study. 
Phyllocladiditey spp. und Australopelles obscutiis dre 
characteristic of both regions, 

Che late Paleoeene L. bale’ Zone is reported trom 
the Southein, Mono ow the highhinds of southeast 
Australi (fayloc ead M00) Gyomosperi poten 


PALYNOLOGY OF THE POONARTINN ANOLE WEL n 


Jominates some of the ussemblayes and Neviofayns 
may be abundant in others, Angiosperm taxi are well 
represented anid the proteaveous clement may be 
relatively abundant in sone: Austredopotliy-vbycuruy 
wid spewies of Mvriceeidifes have not been recurded 
from the Monaro, unlike the other tare Paleocene 
lovalitivs, Cunoniatede, Cyperaceue and Resti- 
onideac are lacking, in keeping With other localities 
in Southeast Australia. but in contrast to thal of 
central Australia 

In the Otway Basin, the Gembierina edwerdsii 
Zouule (Harris 1971) i lite Paleacene in age also 
(Stover & Partridge 1973), Gyimnosperm pollen is 
abundant and A. abscess usually present The 
protetceaus HAA are Well Pepresented in tie Otway 
Basin. but Cunonincene, Cyperaceae andl Resti- 
onaceae have not heen recorded, 

Sluiter (1991) records date Paleocene palynotioras 
from the BMR Mulvorina 2 bore. southeast of Lake 
Pyre. The assemblages are dominated by an 
excepuonall abundanee (up to 07% ) of Cunontacene. 
Gymnespenn pollen is combo aid Netiofages 
mire. similar lo those of Poonarunna-!. but the 
proteuceous taxa ure net particularly abundant, The 
late Puleoeene-carly Hovene suite is similar but with 
wil mcrease in Myrtaeeae. the proteaceous taxa and 
Ansralopillis obscurys, With an overall decrease in 
evanospern pollen (Stuiter 1991), The palynofloras 
ob these lwo localities near Lake Tiyre are thos 
very similar bur Pounarunnd-) has much less 
Cunonmicede 


PO-U4 ot 

Pollen is extremely sparse in dis interval and is 
insufficient for stiuly. Berrvececens, however, is 
sometimes abundant, indicus a fresh-beackish 
Water lake environment, Carbonised particles ure 
present throughout in small amounts bur are 
particukuly abundant at tbe 79-85 m level. 


73-70 nt Lower Nothofacuires wsperus “evr 
eauevatent, und kocene 

The eymnosperm pollen cantenct ts less Una teat 
of the 2. botnet Zone equivalent but os soll 
comsidenible, Attuciriieene — LArdiderridcies 
aistrlty, Dilwwnites -arenatardin) and Detereliinn 
(Lariepallentios foraniy ire the ost abundant in 
(his group, The anwsiospeyin Hora is paruculathy eet) 
With Carstiarijaccae (Haleragectites hurisyit) wid 
Netholasus (Nutialagtdites spp.) moderately 
abundant The spore vontent (ferns. bryophytes. 
Iyeopods) is mocerate. The proteaceaus content 1s 
restricted in diversity and vihundanee. The 
sarhonised particle coment ts low, 

This assembluve is similar to that of the mid 
Bovene unitot Peachawarinna-2 of Lake Eyre Basin 
(Sliiter (294) As well es (he general chaructertstics 


expressed ubove. Cunonfuecac or Myrtaceae 
(Myrraceidites spp.) may be common and the 
proleaccous conten! may be more diverse in the 
usseinblages of Peachwwariana-2. [also contains 
species which Hirst appear ut the late Eovene Middle 
Nothofugidites asperus Zooe of the Gippsland Basin, 
View Auluoridia qualmix, Proleacnlites ch PB, 
wipplatus and Tricolpites themisii (Stover & 
Partridge 1973. 1982). However Slumer (1991) 
rewards all of these species fron the mid Bovene unit 
of Peachawarinna-2, 

Assemblages al Nelly Creck, southern Lake Eyre 
Basin (Alley eral. 1996) are dominated by pollen of 
Casuaripaceae (Aelerugacilites Aeisyiiy anil 
gyninosperm pollen is relatively common with 
Araucatiaceae (Araneurtacites australis, Dilyyiites 
granwaiuy) the most abundant Angiosperm pollen is 
well represented and there is a swwealth of protcaveous 
species. Myriuceae (Myrracesdites) and Nathafagius 
(Nothofiiyidies) are consistently present, Tieolpites 
thamasirand Proleccidiies stipplaiis, with ranges of 
Lower and Middle V. asperus Zone equivalents in 
central Nustralia. are present alsa (Alley ev ef, 1996), 
The Poonurunni-| assemblage thus compares well 
with those from Nelly Creek, with the exception Unt 
the proleacequs componenl is more diverse and 
abundant in the latter, The Poonirunna-! assemblage 
is therefore assigned to the Lower NO dsperny Zone 
equivalent of mid hocene age. 


Commperiveny 

Mid Eocene macrofossil assemblages at Nelly 
Creek. Poole Creck and in Some of the Silerete floras 
have a number of taxa with botanical affinities i 
common with the Pooparunna-b  microfossil 
assemblage, and the weneral characteristics are 
similar Four proteaceous teal faxu. Ageriis 
(Ariucariicede). Podovarpaceae.  Civinaastonia 
(Casuarinaceae) and Myrceipln thon (Myrtacese ). ire 
listed (Christophel af al 1992), More detailed 
comparisons OF macro- and microfossil assemblages 
ure pat possible beeause of the dillerence in 
provenance, transport and preservation of the plant 
parts Fouad br (he owe types of assemblages. leat 
physidgnomy yields invaluable evidence about the 
veseraition and (his topic (§ discussed baler, 

Tie Pooburunna-l assemblage bus less Nothofiguus 
und more Casuaringceat when compared qwith the 
Lower Norlofieidites ayperus Zone of the Gippshuid 
Basin (Stover and Partridge J973. 1982. Partidye 
1976) Aswell. the minges of muny Species are not the 
same in the tyr localities. Phe nen-woody swamp 
taxa of Cyperacede. Restionacedeé and Sparcaimnicear 
have not heen repented fra the Lower 
Nothofagidites oyperus Zone of the Gippsland Basin. 

This Poonarunna-| palynoflora is probably time 
equivileal to the Prateacidtites pachypolas Zone af 


OM HLA. MARTIN 


the Ouwuy Basin in Victoria (Harris 1971) whteh os 
rich In Nothefavas and proteaceous taxu, Mid 
Eocene floras are also found ifthe St Vincent Basin, 
South Australia, where the palynofloras have wy 
abundance of fern spores but frequencies of 
symnosperm, Casuarinacene und Notiofagis pollen 
are Jow, There is a diversity of proteaceous and other 
angiosperm pollen. will no single wroup dominant 
(Alley 19872 Alley & Broadbridge 1992). 

When the mid Eocene palynoflora ab Poonarunna 
is compared With those of Anglesea (Christophel e7 
a, 1987), the latter has more Nothofevus and fewer 
gymnosperms, Palynofloras at Bungonia (Trussell 
& Owen 1988) also lave few pymnosperms when 
compared with those in Poonurunma. 


61-73 m 

Pollen was not recovered from this interval bul the 
alea, Momyveceecus, is present and very abundant 
from 67-73 m. The 70-73 ny level has Peers 
also. These algal species indicate a fresh-hrackish 
water lake cnvironment. The carbonised particle 
eontent is low through (his interval, except for te 
67-70. m level where tt is high, 


YO] i 

Pollen was. not recavered from this interval. 
Curbonised particles are present ihroughoul, 
increasing Up the section, except for the 16-20 in 
level where (hey decrease. 


f-Goin. Plineene-Pleisiecene 

Asteraveae (1 7ithulifloridites spp). Poaceae 
(Cadmmuriites monoporites) and the chengpou type 
(Chenepeadipullis chenopodiacevidess make tp 
the bulk ol the pollen coun, Casuarinacene 
(Halaniuaciudites — harristiy antl Myrlacene 
(Mvrfaciddes spp) are present. but pyiibospern 
pollen ind spares gre miniial, Curbonised particles 
dre extremely common, 

This type of assemblage has mob heer feconted 
previously from central Australia and there is ne 
direer means of dating 7h The general quantitative 
uspeers of the ussemblage suggest oy Pleistaeene ae. 
bused upon experienee in southeastern Australia 
(Marti) 1987) ohut i licks the distinctive Pleistocene 
Hihuhflavidies plewstecenteus (Macphinl 1996). 
Palvporing aranidara of late Miodene Pleistocene 
(Mawphi) (86) os present. Preqhertcies similir to 
(hose shown in Fig, 3 may be Polit i shieibliids 
and regions lonay (Stuiter & Kershaw TY82), When 
the Pouonarugia-T assemblave ts compared with 
surface sainples lront shroblands vround Lake 
brome. ii is romnurkubly similar, except Cora greater 
CY periceae pollen content. suggestiiy hat there was 
yore swam vesetation in the Pliseene Pleistocene 
Hin there sat Lake brome today 


Tubaliflaridies spp. Yurse appear in the rad 
Miocene of Southeastero Australia (Stover & 
Partridge 1973), bat there are wuimerous reports: ol 
these species from the early Miocene aid atew fray 
the Oligocene (Muller l98}io My experience bis 
shown that most pollen types are not common inthe 
carly part OF their ranges. but become more abundant 
faite, AL Poodaruinna, Tidiiliflertdites is the mast 
abundant group, Suggesting a relatively your ape. 
ie. Miovene if no Pleistocene. 


1-61 
No pollen sas recovered, The charcoal) particle: 
content was very high. 


Palacoveyetation and palaeoenvironment 


The Cenomiuniin palynoloris are donmimiued by 
bryophytes, lycapods. pleridephytes und wymno 
sperms but the angiosperm content is relatively small 
and this Suggests that the vegetation was mainly Forest 
with gymposperms forming the (major part ol the 
canopy, if not the whole of it One palynoflory 
sigwests more extensive bogs or wellands, its 
discussed previously, These palynofloras predate the 
#ppearance of proteaceous and Netlofaguy pollen 
types Which become a distinctive clement in-yourper 
palynofloras, 

The hte Paleocene palynofloras of Poonsrinn-| 
indicate what the region was iainly forest. The 
eymnosperms Padecarpuy (Padacdnpidites spp. 
Lawaraytrobos, the Huon pine Ce aaledidites 
maser) and Daervdiuin (Lypistepollenites 
Hlermnin) were common. ind there was a diversity of 
aiher Yyminosperm tind, 2. Decrvearpis (Dace 
rarpites), Microcachyrs (Microcachiryidites). Cusuia- 
rinucewe (Heloregacntites herrisii), Nethafasus 
(Nathofasidties spp), Myrtaceae (Myrracerdifes 
spp.), Caunomaceae/Elaeoctrpicede were pact of the 
forest Canopy, though probably a relatively (ior 
part. There is a wealth of profeadeous pollen ty pus 
bul mest of them cannot be identified with living 
tina. AL leust some of ther are Jikely to have been 
lprest trees, Similar to the protwacenus (rees foand iy 
rainforests toray. Proteuceous species have iW very 
luw pollen representation (Kershaw 97, O71 
1976, Murti 197%) hence. Wese eXciner praleneeniis 
haku Were probably fur mere ubiidont i the 
Maleocene vevetation than is suggested hy the pallens 
frequencice, Swamp communities were Revie i 
extent aad conlayned me on-wwoorly fina ef. 
‘Cullivicvhe® (Avatralepoltis, ebscuruy) (Macphal 
990). Cyperaveae ane Restionuceie, Other 
wigiospenms were diverse (Table 2) and if they were 
law pollen producers. ds is tie case with most insect 
pollinated species. they were probably mote 
abundant than is Sugwestéd by the pollen canits 


PALYNOLOGY OF THE POONARUNNA NO. | WELL 99 


A lresh water Jake environment with copious 
Botryococcus and some Pediastrum succeeded the 
late Paleocene floras, through most of the presumed 
Eocene, There is. however, insufficient pollen for 
study through most of the sequence. Within this lake 
sequence, there is a layer of sediment with a goud 
pollen content, suggesting that the lake had receded 
from this area and that it had become vegetated. The 
id Bocene palynoflora in this layer suggests mainly 
forests with little non-woody swamp vegetation. 
Arwucariiceac, Dacrvdiam, Casuarinaceae and 
Nothofagus were prominent, the proteaeeous content 
Was low at this particular location and there was a 
greal diversity of other angiosperms, 

The Pliocene-Pieistovene vegetation was probably 
open woodland or shrubland, generally similar to that 
of today in arid: abd) semiarid regions, ‘Trees, if 
present, were probably Cusuarinaceae and 
Myrtaceae. bul both of these families contain shrubby 


Tarr +. Register vf strated specimens. 


taxa as well as trees. Acacia (Acaciapollenites 
miyriosporites) is present, and as Acacia has a very 
low pollen representation (Sluiter & Kershaw 1982), 
it may have been common in the vegetation. similar 
to that of today. The very Jow frequencies of 
Araucariaceae and Podecarpus may have resulted 
from long distance dispersal. or there may have been 
small, rare stands in the landscape. IL is remarkable 
that a Plio-Pleistocene palynoflora has been 
preserved at all, A wetter climate and some 
condition(s) which allowed a more rapid burial of 
sediment must have prevailed for a brief interval, 

IP itis accepted that most of the carbonised particles 
result from burning, then the Tertiary recerd (Fig. 3) 
suygests that burning increased during the presumed 
Neogene, reaching a maximum in the Pliocene/ 
Pleistocene. There was, however, episodic burning in 
the Eocene. Very lew carbonised particles are found in 
the late Paleocene and Cenomunian palynofloras. 


CRETACEOUS SPECIES 
Acriuirch sp. indet. In 


Nequiniradites splinieasius 4A,B 
Aeainriradites verracisus 4b F 
Afisporires spect. As aranedts ON 
Arainaridedtes duatralts OB 
Bacilalisporttes con ancusts AG 
Baliieisporites wlenelwensis SA 
Balmeispovites hialodicivus 4H 
Balmeiwporiles ftaludictus Al 
Bilmeisporites tridictyus 4B. C 
Balineisporites tridietyuy 40 
CONTE TOS IMOSPONTIOS USI OHNTS Ay 
CONN ONPOVUON UUSIELTOHNTS 4K 
Ceratosporiios equatis SLL FP 
Creatricasinportes sp, of Burer 6C 
Chewiricasixportes sp, of Burwer 6D, F 
Claviferi triples 6] 
Clavifera triples 6k 
Corellia sp. ch Oo clesseidos eo 
Cryhelosporues puanerdtin OL. M 
Crybelosporttes [ACHAHIN tC 
Cryhelaspurites sirhaus TALK 
Crpulifereidecpoullenties el, Co prarvuliey oI 
Cyaridites ustealis ol 
Cyathulites minor 6H 
Picopapalliy sp. OM. N 
Dictvophyttidites sp. GN 
Forwninisports daily ra8) 
Peel ispeeis won ae edenyis it) 
hoveauhliheniidites confossus Th 
Poveatetradies fistilasus OA, B 
CM HOLES Cire dadidites oP 


Horalegiedht sp. HON, 


Slide No. England Pinder 


coordinates 
3060-3 Wa3-0 
BOGS A O35) 
3064-1 163-0 
A060-3 K4l-2 
A064-| p55.4 
3064-| 135-| 
3060-1 035-1) 
3UO0-3 J51-1 
A0O4-| 560-0) 
3060-| M34-1 
3062-| 056-3 
3063-1 P47-2 
3060-3 V53-\ 
3062-3 Cid4-4 
3004-3 S40-4 
3004.3 R44) 
3065-2 MS5O-( 
3003-1 R29.) 
3002-3 Vaud 
SOOO 4 O40) 
AMY Var 
AOO2-] R440) 
OH4- | W322 
$64 3 S5N-] 
WAL 3 Md5-| 
3000-1 Jas-| 
3060-3 S454 
3060-4 Pls.) 
S004. | W53-1 
A -3 33-0) 
A003 P31-2 
WH 3 S40-2 
Mi 42 L603 


LOO H. A. MARTIN 


TABLE 4 continued... 


Species Fig. Slide No. England Finder 
coordinates 

Laevigatosporites ovatus 7G 3060-3 Y42-4 
Levaniella sp. 10C 3063-2 M54-0) 
Lecaniella sp. 10D, E 3063-3 O#0)-2 
Levaniella sp. 1OF,G 3063-3 F50-0 
Lecaniella sp. 101 3061-1 H36-3 
Lifiacidites sp. cl. L. kattangatdensis oP 3060-1 $33-0 
Liliacidites sp. 9S 3060-1 N29-3 
Microcachryidites antarcticus 9D 3060-3 V42-0 
Microfoveolatosporites canaliculatus 8K, L 3063-1 QO43-0 
Ornamentifera sp. cf. Q. sentosa 7TH, 1 3062-1 O53-0 
Perotrilites jubatus 71..M 3064-2 Feo-4 
Perotrilites jubatus 8C 3064-2 N49-2 
Phimopollenites augathellaensts 90 3060-3 O36) 
Phimopollenites augathellaensis 9Q,R 3064-2 Q52-0 
Phimopollenites pannosus 9U 3060-3 O36-3 
Phimopollenites pannosus OV 3060-3 GS0-0 
Plicatella distocarinata 4c. D 3060-3 N37-4 
Podocarpidites ellipricus OF 3960-3 X43-2 
Podocarpidites exiguus OF 3060- | M294 
Podosporiies sp. OH. 1 3060-3 X39-4 
Podosporties sp, OI IK 3060-3 Y44.2 
Polycingulatisporites sp- SA. B 3060-3 F54-3 
Reticuloidesporites arcus 8G 3063-3 039-0 
Retiriletes austroclavatidites 8D 3060-3 Y45-0 
Retitriletes austroclavatidites 8E, F 3060.2 P40-0 
Ruffordiaspora australiensty 6A 3003-3 J54-3 
Ruffordiaspora ludbrookiae 6B 3060-3 M36-0 
Saeprodinium gravatiensis 1OH 3060-3 Y37-0 
Schizosporis reticulatus 1OK 3064-3 O40-4 
Senectotetradites varireticulatus oT 3060-1 R40-3 
Sestrosporites preudvalveolatus 60 3062-3 K41-3 
Stereisporites antiquayporites SN 3061-3 O50-0 
Stereisporites antiquasporites 80 3061-3 H46-3 
Slereisporites pocockit 6F, 3063-1 Q47-0 
Stoverisporites microverrucatus 8M 3061-2 O59-0 
Trichoromonosuleites subgranulatus 9G 3060-2 U65-0 
Tricolporites sp. ch. 1) apoxyvexinus OXY 3060-1 M6l|-3 
Trilohosperites tribotrys of 3064-3 H45-0 
Trilobosporites trioreticulosus 8H 3060-3 R48-4 
Triparaletes sp. ef. T. simplex &J 3060-3 X4A0-4 


TERTIARY SPECIES 


Acaciapollenites iyriosporites 13C 2983-1 N34-4 
Aglaoridia qualamis 7G 3000-3 P4]-2 
Amosopollis difwynittes ls 3057-2 $40-3 
Amasapoallis dihvynites 13F 3057-1 N36-0 
Amosapollis dilwynites 13G 3006-3 D31-0 
Araucariaciales australis 121 3000- | O40-() 
Arecipites sp. ct. A, miniuiseabratius 13D 3006-1 X42-1 
Australopallis obseurts I3N.0 3059-2 TS3-0 
Acolld sp. HW 3000-2 J45-0 
Beaupreaidites clegansiformis I3I,K 3000-2 LAS-3 
Bairyocoecus braunit I8B 3000-1 V42-3 
Barryecoccus braunis 1sD S000- | V42-] 


Cumarozonosporires amplus WA 3006. | W38-4 


PALYNOLOGY OF THE POONARUNNA NO, 1 WELL 101 


TABLE 4 continued... 


Species Fig. Slide No, England Finder 
coordinates 
Camarazonesporites amplus 11B 3000-3 042-1 
Camarozonosporites bullans 11D 3059-2 O40-3 
Camarozonosporites 8p. H1E,F 3000-2 E47-0 
Chenopodipollis chenopodiaceoides 13L 2983-1 Y53-0 
Cunoniaceae (tricolporate) 13S, T 3058-3 O41-0 
Cunoniaceae (bicolpate) I3L.V 3057-2 T37-1 
Cupressaceae/Taxodiaceae 12D 3058-2 $55.3 
Cyathidites paleospora UW 3059-1 O60-2 
Cyathidites splendens HG 3059-1 V45-3 
Cyathidites splendens UH 3059-1 V52-2 
Cyperacenepollis 13M 3000-3 D39-0 
Dacrycarpites austratiensis 12L 3059-1 $50-0 
Dicopopollis sp. 18k 3000-1 29-0 
Dilwynites eranulatus 12) 3000-1 R42-0 
Dilwynites gramtlatus 12k 3006-2 W39-3 
Dilwynites eranilatus IBA 3006-2 V58-0) 
Elaeocarpaceae 13Z, AA 3005-1 K39-0 
‘Ephedra’ notensiy 13B 3006-2 133-4 
Ericipites crassiexinuy 13P.Q 3000-! R30-0 
Gleichenia circmnidues 11c 3059-3 LS9-2 
Gothanipollis basxensis 144A, B 3006-3 W57-1! 
Grapnelispora evansit 124 3059-2 Q43-3 
Halavagacidites haleragoides 13R 2983-1 030-2 
Huloragacidites harrisit 1G 3000-1 V4l-4 
Hexpollenites austroclavatns 14D, E 3059-2 49-4 
Lewalanipollis tf L. rectoniareinis L6G 3006-1 F43-4 
Lewalanipollis of. Persoonie ISI. K 3000-3 C55-1 
Liliacidites lancealatus L4P 3058-3 V50-0 
Lygistepollenttes florinti 12G 3059-2 P4)-1 
Malvacipollis subtilis 14h 3000-1 Q28-0 
Micracachrvidites antarcticus 12N 3059-2 L34-1 
Milfordia homeopunctate 14c 3000-1 228-0 
Milfordia homeapunetata lay 3000-1 PAT-3 
Milfordia hypolaeniodes 14k 3006-3 049-0) 
Myrtaceudites eucalyptoides 140 2983-1 Q33-1 
Morkallacysta pyrantidalis 1kC 3059-2 Q28-1 
Myrtaceidites eucalyptoides 14R 2983-1 V41-2 
Myrtaceidites verriicosus 148 3000-1 S40-1 
Nathofagtdites emarcidus 14L 3000-1 K58-2 
fothofagidites demitutiis 14N 3000-3 T48-0) 
Nothofagidites faleatus 14M 3000-2 P36-4 
Nuxipollenites kempti 140, P 2983-1 J46-1 
Pancolpate sp. [SE 2983-1 139-0 
Panporate sp, 18G 2983-1 Q40-1 
Pediastrum sp. tSA 3000-1 R53-2 
Phyllocladidites mawsonii 12E 3059-3 R43-0 
Phyllocladidites reticulosaccams 12H 3059-2 Q42-4 
Podocarpidites exigiuus 12M 3059-3 K39-1 
Polyorificites oblatus I4BB.CC 3000-1 O59-1 
Polypadiacoisporites sp. ct. P. retiregatus 11L,M 2983-1 J43-2 
Polyporina granulata I7A,B 2983-1 L46-0 
Propylipollis ivanhoensis 147 3006-3 Q47-3 
Propylipollis ivanhoensis 14W 3058-2 Q58-2 
Propylipollis latrobensis 14X 3005-1 U40-0 


Propylipollis sp. cf. P. pseudamoides 16B 3006-2 V64-0) 


102 H. A. MARTIN 
TABLE 4 continued... 


i 
Species Fig. Slide No. England Finder 


coordinates 
Propylipollis sp. ef. P. pseudomoides 16C 3058-1 $56-4 
Propylipollis sp. cf. P. pseudomoides 16D 3006-3 Q47-3 
Prapylipollis sp. cf. P. reticuloscabratus 14U 3006-3 $433 
Propylipollis sp. cl. P. retieuloscabratus 14V 3058-1 $56-2 
Prapylipollis sp- 14y, 7 3006-1 X39-1 
Propylipollis sp. l4AA 3005-1 X43-4 
Proteacidites adenanthoides ISA. B 3058-3 O49-2 
Po angulatus 15G 3057-1 X57-0 
Po angulatus 15H 3059-3 I38-0 
P. cooksoniae 16A 3006-2 MS50-0 
P crassus Isc, D 3059-3 159-2 
P. fromensis 1365.7 3059-1 049-2 
P. grandis IS] 3059-1 R60-4 
P meurvatus ISN 3000-1 $40-0 
Psp. ef, Po ineurvetis 16N, O 3057-1 K34-1 
P. ef, obscurus 16d. K 3006-1 K46-1 
Pel, stipplatus lol 3006-2 $58-0 
Proteacidites sp. | 16E. F 3059-3 O41-0 
Proteacidites sp. 2 16L,M 3006-| MS1-0 
Proteacidites sp. 3 16H 3059-3 Q51-2 
Quantiniapollis psilatispora I7L. 3000-3 C53-4 
Retitriletes austroclavatidites 12B,C 3059-3 N59-2 
Rhopites alveolatus I7E 3000-1 $40-2 
Rhapites sp. cf. R, alveolatus 17P,Q 3059-2 U65-2 
Santuldmidites cainozoteus 17F 3000-1 X44-2 
Sapotaceoidaepollenites rotundus IIR 3059-2 T62-1 
Simplicepollis meridianus 17C, D 3000-1 KS4-2 
Simpsonipollix sp. 13H, 1 3059-2 $27-0 
Tricolpites sp. cf. T. asperamarginatus 17H, 1 3000-1! P50-4 
Tricolpites sp. cf. T. confessus 170 3059-2 R41-0 
Tricolpites sp. cf. T. discus 17M, N 3000-1 Q40-0 
Tricalpites phillipsii 177 3005-1 W434 
Tricolpites thomasii 17U,V 3000-1 W39-1 
Tricolpites sp. 180, P 3058-2 Q5}-2 
Tricolporites angurium WW4 3000- | P29-2 
Tricolporites leures I7TW, X 3005-1 Q46-0 
Tricolporites sp. | 17€C 3058-2 N41-0 
Tricolporites sp. 2 I7AA, BB 3057-1 T33-4 
Tricalporites sp. 3 18] 3058-1 P55-2 
Tricolporites sp. 4 [8s K 3058-1 W530) 
Tricalporites sp. 4 I8L 3058-2 N42-4 
Trivolporites sp. 5 18M, N 3057-1 W33-3 
Tricolporopollenites endobalteus I7R,S 3000-1 062-0 
Triletes sp, ct. Ty utbereuliformis LIK 3000-1 N54-3 
Trierttes minisculus I8Q 3058-2 $55-3 
Triorites sp. 18H 3059-3 T33-3 
Triporoletes reticulatus 12F 3059-2 S57-3 
Triporopolenites ambiguus I7Y 3000-1 L43-4 


Tubulifloridites antipodica/simplis I8R 2983-1 §39-2 


PALYNOLOGY OF THE POONARLINNA NOL | WELI 104 


The development of the arid Mora and vegetation 


The aim of this study was te find fossil evidence 
about the development of the arid flora and 
veweuition, The late Paleocenesmid Bocene 
palynotloras deseribed here. with a@ substantial 
tainforest element. are clearly not arid adapted. The 
vegetation, however. grew ol the floodplains und 
depositional basin, and such habitats would not be 
the first in the hindscape to register aridity. Middle 
Rocene macrofossil assemblages from other 
lowalities in the Lake Eyre Basin have some large- 
leuved Land. Consistent with inforest. and ip variety! 
oF smaller leaved taxq, suggesting selerophytlous 
yegetulian (Christophel e/ af, 1992; Christophel 
1904). The veeetation is interpreted as being gallery 
nudtorest along the watercourses und selerophy Tous 
vegelutiin, adapted both 1 low fertility and 
scusonally dry conditions. in the intertives. These 
maveofossil assemblages are unique to cenrral 
Australi (Greenwood ef a! 19980: Greenwood 
1904). 

The diterval betWeer the imid BKocene and the 
Plincene-Plerstocene cid not yield pollen, The 
Phiocene- Pleistocene assemblage is penerally similar 
to thitt produced by the extint arid Shrublanus ot 
Lake Vrome (Fig. 3) but contains some disparate 
clements, Dodomeiea Iriqguetra (Nuxipollentions 


kempii) ts tound in this assemblage, bul the modern 
species is restricted to wet seleraphyll forests of the 
southern half of the cast coust of Australia (Martin 
1997). Dodenaed tiquetre 15 also present in the mid 
Bocene of the Lake Eyre Basin (Sluiter 1991) and 
probably became extinct in this region at some time 
fier the Pliocene-Pleistacene. 

There huve been miuny studies On the Mora of the 
arid zone that have generated Various hypotheses 
ubout it} orgins, As many of tbe tisit to the arid one 
show affinities with related taxa in adjacent regions. 
miost of the hypotheses involve reeruiiment from the 
floras surrounding the arid yone (Barlow 1981). 
Such studies, however. do not reveal taxa which once 
grew in the arid zone and have beeome extinet there. 
such as Dedonuea triquetra, ‘Vhe fossil record 
sueesis thatthe vegetatn developed by continneus 
adaptation of some of the taxa already in the fegion 
(by evolution of new species) to a drying environ: 
ment. Those taxa that could nor adapt te the arid 
environment disappeared from the region. 


Acknowledyments 


Tam indehted ta N-P Afley of the Department of 
Mines and Energy Resourees (formerly the South 
Australian Depariment of Mines and Energy) for 
Invaluable assistance with this project which seas 
supported by wn Austrilinn Research Grant, 


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106 L.A. MARTIN 
Appendix | 


Late Cretaceous Systematic Palynology, For the distribution of the species in the bore, see Table 1. Por the 


register of illustrated specimens. see Table 4, 


Spores 


Genus Aegultriradites Delcourt and Sprumont emend, 
Cookson & Dettmann 19Ot 


lype species: Aeyutriradires dubius Delcourt & 
Spromont emend. Delvourt, Dettman & Hughes 1963 


Aequitritudites spiuilasus (Cookson & Dettmann) 
Cookson & Deltmunn 1961 
FIG. 4A, B 


Comments. The spinulose clements over the distil sut- 
face ure about | tim in diameter and the exime | um thick, 
Compare with A. verrivesis. Spore body, 45 pm. overall, 
55 um, 


Stratigvaphic Range, Marly and Middle Cretaceats 
(Detunann 1963), Aequitriradites avcus/spinilasas, Tron 
the Muroypara florida Zone. late Jummssie, into 
Phyllocludidites mansani Zone, Tironiin-Contacion 
(Helby et et, 1987), 


Aequitrirudites verrucasins (Cookson & Dermiunn) 
Cookson & Deltriunn 196) 
FIG. 4E. F 


Comments. The verrucate elements over the distil sut- 
face ure 2-3 pint in diameter and the exine 2-36 yun thick, 
Compare with A, spiudeasits. Spore body, 48-57 pm, over- 
all FO-RS pom. 


Stratigeaphic Runge. Widely distributed in south-eastern 
Australia in the Upper Mesovorw (Detain 1963). 


Cienus Bevalatisporites Thomson & Pflug 19534 


IWpe species: Keculerisparites pennarins (Wolff) 
Thomson X& Pilg P54 


Bacutadsporties cameiunensis (Cookson) 
Potonie L956 
MiG, 4G 


Stratigraphic Range, Prom the late Jurassie-larky 
Cretaceous. it is common throughout the Upper Mesocote 
(Detimann 1963), 


Genus Balmeisporites Cookson & Dettmann L958 


Type species: Balineisperites halodicyins Cookson & 
Derrmann [958 


Balueisporiter gleneleensis 
Cookson & Dettmann 1958 
FIG, 5A 


Comments. This species is similar to Bo falevtictviey bul 
the spore body is larger and the exine thicker: The inner 
homogenous layer is 5 um thick on these specimens. conte 
pared with 12 pin on A. fralpdiecivus, Spore body, 162 4 112 
um, 


Stratigraphic Range, Within the Plicdsela distocertretis 
Zone. Cenomanian, to within Tricelpires puchvextrens 
Zone, Santoman, of south-eastern Australia (Detimann & 
Playford 1969), Cenormanian, possibly Turaniiun of nerth- 
west Australia (Norvick & Burger 1975), 


Balmeisporites Ralodtcwus 
Cookson & Dettmann 195k 
FIG. 4H, 1 


Comments. Most of these lurge megaspores are broken, 
The spore exine Consists Of un toner, homogenous layer b2 
po thick and an outer granular aver about 5 yigr thick, the fat 
ter supporting the muri of the reticulum. Spore body, #7- 110) 
pm equatorial diameter, overall, 137-166 pin xs 170-235 pny 


Stratigraphic Range, Cyhelasperites sirtaiusy Subsone, 
Late Aptian-Early Albian, to the lower part of the 
Appendivispories distocarmans Zone. Cenomaniao 
(Dettmann & Playford 1969), 


Balmetspovites tridiciyuy Cookson & Dettmann |YS% 
FIG, 5R-D 


Comments. The absence of a reticulum, a thick. ier 
homogenous layer (5 pin), an outer granular layer. 1-2 pm 
and the large membranous winglike oulgrewths distin- 
guish this species (Cookson & Detiinann [Y5K). These 
specimens show sinuous ridges 4 pot high (arrow), whieh 
MAY anastomose, Spore body, $2-85 8 42-110 pin. 


Stratigraphic Range, Aptiin-Albiun (Cookson & 
Detimunn 1958; Detonann 1963), Cenomunian. this study, 


Genus Cumarezenesporires Pant 1954 ex Potonie 
1956 cinend, Klaus 1960 


Type species: Refasperiies eretaceauy Weyland & 
Krieger 1953 


Camarosonosporites austrutiensis Norvick & Burger 1975 


FIG. ALK 


Comments, The distal surface has coarse rugulae about 3 
um wide, Separated by grooves | yim wide, On the proxinnuil 
surface, the pattern ts finer und the contact surfaces ure 
almost smooth, This species is smatier than CL aanplies, 2%- 
S7 um compared with 57-109 pin respectively (Norvick & 
Burger 1975; Dettmann & Playford 1968). Equatariel 
diameter, 40-44 pum. 


Stratigraphic Range. Albian of the Great Artestan Basin 
und Albian into Turoniaia of Northern Australia (Norvick & 
Burger 1975). 


Genus Ceratosperites Cookson & Detumanin LOSS 


Type Species: 
Dettmann 1958 


Ceratasporites equilis, Cookson & 


PALY NOLOGY OF THE POONARUNNA NO. | WELL 407 
E 


Figs 4. Cretaceous species. A, B. Aeguitriradites spinutosus. C.D. Plicatella distocarinatis. Eb. Aeguitriradites verruce- 
sus. G. Buculatisporites comaumensis. UW, 1, Balineisporues holodieytus. J. K. Canimoconosporites australicnsis. Scale 
bars = LO pm. 


108 HA, MARTIN 


Cerafosparites equalis Cookson & Detain 1958 
FIG. SE, FE 


Stratigraphic Range, From the Revitrileses wetheroensis 
Zone, latest lurassie (Helby en uf 1987) trough the 
Paleocene (Hurris L965; this study), 


Clomiricusisperties sp. oF Norvick & Burger 1975 
FIG, 6C-E 


Deseription. Spore tilete, ainb tiangulin, sides slightly 
convex lo deeply concave, Most spores hive deeply con- 
cuve sides, appear 3 lobed and present in equatorial view, 
Lasiree ex tench almost To equator, membranaus fps 4-5 put 
high, Three sets of parallel muri dinastomose in radial region 
OF distal surface. Edges OF muri irregular or wavy with 
small Fovule Within muri, especkily where sets of mur 
anustomose. Four muri and intervening grooves, &- 10 ym, 
Equatorial diamerer 43-53 ain, polar diameter 35-40 pin, 


Comments. This species is distinguished trom: Auifferd- 
laspord dustralaiis by the Tegular ature et the muri 
Norvick & Burger ()975) fuure an undeseribed species, pl, 
20 Tig. 4. similar to this one. 


Strativraphie Range. Norvick & Burger (1975) note that 
their undescribed species in the Cenomanian iy ehuraenristic 
ofthe Upper Albian in Queensland. Cenomanin, this study. 


Genus Chavifera Bolkhovitena YOo 


‘Type species: Clavierd frinfex (Bolkhovilinwy) Balkhoy- 
Hind YOO 


Clayifera iiples (Bolkhoyitinad) Bolkhovitina 1966 
FIG. Ot, K 


Comments. The distal surface is strongly arched and the 
proximal pytaniidal, The nifterhadial crassitudes ure 4-5 200 
thick. features Which distinguish it from Gleietenidites. Us 
alfinities are with the Gleicheniaceue, 


Stratigraphic Range. From within the Caproyporna par 
deve Zone, Albitn. ii northern Australia und rare in the 
Micatela distocainde Zone, Cenomaniin, southeastern 
Austiihia, through the Forcipites Jamu Zone. Maasti- 
chlian (Helby ef cal L987). 


Geous Crybelosporires Detinann 19634 


Type species: Crybelesporites sirtatuy Cookson & 
Detimann L958 


Crybelasporifes punctatis Detimann 1963 
FIGS 61, M, 7 


Overall diamerer, 36-55 4 28-48 pon. 


Stratigraphic Range. Lower Creticcous (Dettinann 1963), 
Cenomanian, this study, 


Crybeloyporites strats 
(Cookson & Dettman) Dettmann 964 
PIG. 7A, B 


Comments, The sclerine is 4-5 pind thick with a henge. 
neous ioner layer | pin thick and a ruffled outer layer which 
is irrewularly Strigte on the proximal side and ceticulite on 
the distal surface. The mur are thin and sinuous, and the 
lumina 4-4 pm wide, All these feattires are a guad fit with 
Co sirfeeties. 


Stratigraphic Range, Crvbelesporites steletis Zone, inte 
Clavifera Triples Zone. Late Aptian into Turenian 
(Dettmann & Playford 1969), ©. sérietius Zane through 
Phimopollenites pannosus Zone. latest Aptian through 
Albian oF gorther Australia und continuing inte the 
Tubulifloridites litter Zone, early Maastrichtian of southern 
Australia (Helby ef ef 19ST). Cendimaniin of northern 
Austrailia (Norvick & Burner 1975) this study), 


Genus Cwathidites Couper |953 
Type species: Cyathidites australis Couper 1954 
Cvarhidites australis Couper [953 
FIG. 61 


Stratigraphic Range, Common throughout the upper 
Mesozoic in southeastern Australian (Det 1963), Prom 
the Permian (Poster $979) into the Tertiary (Hitrris 1965). 


Cyaidites miuar Couper 1953 
FIC), 6H 


Comments. Very conten to this study 


— Strtigriphie Runge, Prom the Jurassic (Dettinane 1963) 
info the Tertiary (larris 1963), 


Genus Dicnophyflidites Couper emend. Detlimann 1965 
Type species: Dictvophvlidites harris’ Couper (58 


Dictyvaplryllicites sp. 
FIG. 6N 


Comments. There is considerable variation inthe popilie 
tion assigned to this genus, 


Genus Feruininisporis Keutgsch 1959 


Porenmninisports wort egien sis 
(Cookson & Dettmann) Detimann (63 
FIGS 60. 7E, F 


Stratigraphic Range, Prom the Rufferdiaspora austrulient 
viN Zone, cartiest Cretaceous. 10 the Phyllectudidites niin 
yet Zone, Turoniin-Coniieiin (Helby ef el. LOST) 


Forannisporis dear 
(Cookson & Detiminn) Detmiunn 1903 
FIG 7D 


Stratigraphie Range. Widely dispersed in the Upper 
Mesozore of southeustert, Australia (Dettmann 1963). 


Gettus Poveduleichentidites Norvick & Burger |Y75 


Type species: Fovewaleichentidites (al. Glejehentidiies 3 
comassiy (Hedlund) Norvick & Burger 1975 


Foveauleichentidites Comfossus 
(Hedlund) Norvick & Burger 1975 
FIG. TILK 


Description, Amb toiangular with rounded upives, (rilete 
lusurae thin and straight. reaching to apices, Exine | yin 
thick with interradial crassitudes up to 6 pm wide, Fovulae. 
< | pm in diwmeter. spaced up te | pin apart accur on bork 
surfaces, Equatorial diameter, 30-32 yim. 


PALYNOLOGY OF THE POONARUNNA NO. | WELL LOY 


B 


Vig. 5. Cretaceous species continued. A. Balmetyparites glenelgensix. B-D_ Balmeisparites iridietvus, EE. Cerataspurites 
eglalis, Seale bars = 10 um, 


110 L.A, MARTIN 


Stratigaphie Range. Cenoniunian of northern Australia 
and sporadically in the Albian of the Great Artesian Basity 
(Norvick & Burger 1975). 


Genus Gleicheniditey Ross ex Delcourt & Spruniont. 
emend. Dettmann 1963 


Type species! Cleiehentidites senonivus Russ 1949 


Gleicheniidires crecinidites (Cookson) Dettmann [963 
PIG. OP 


Strutizvaphie Range, Gleieheniidites spp. first appear in 
the Ewly Jurassic CHelby ef ak PORT Ch cireduiedites ts 
vommon in Upper Mesozoic sediments af southeastern 
Austalia (Detimann 1963), tis comparable to Gleielienin 
ond ranges through the Tertiary tothe present diy, 


Genus Loevigatosperites Vruhan (Y33 


hype Species: Leerivafesporites vileerds brim) 
brat 1933 


Linvigaresporites ovalis Wilson & Webster 1946 
hIG. 7G 


Comments. A conmmnion und widely distributed species in 
the Upper Mesovene (Dettinanh 1963. Norvick & Iurger 
1975) and (hrough the Tertiary, This very common in some 
af the samples al this study, 


Genus Microforeolaiasporitey Kriteseh L959 


Type species: Micrafoveolatosporites conalicilatis 
Dettmann 1963 


Micrafoveatuiasporiies canulicniaus Detunann 1963 
HIG, aK, L 


Stratigraphic Range, Albian ob the Great Artesian Basin 
(Detlniann 1963; Norvick & Burger 1975), Cenomanian of 
northern Australia (Norviek & Burger (975) und central 
Australie (this Study), 


Genus Orawnentiferd Bolkhovitina [966 


Type species: Ornamentiferd echima (Bolkhoviting) 
Bolkhovitind [Go 


Ornamentferd sp cl QL ventavet 
Detionn & Phivtord 19o8 
Pie, WL | 


Deseniplion, Ah wiinetlay with romnded angles, illete 
sear will elevaled pnrembranous lps. Inferridial crass itudes. 
Ho 20 tin long, 5-6 pio wide, bear simious ragulae <1 yr 
tinh aod tpn wide, Rugulae extend over distal surtace, 
Prosonel surlace palteried witht low verrucae, Diamerer, 32 


a) 


Comments. The rugulite pattern ayer the eriussitudes di 
fers Hom OO) seitesa whieh has erassitides with serrate 
fein. Phe pater atthe distil sures Covers the entire 
supface On TES form whercis ibis restricted to a reiieular 
area. With the pices io the inlerradial region on GQ) sents 
(Detmano & Plinford 1968). This specimen is similir fo 
the ane Hanred by Norviek & Burger (1975, pl, 23, fig. 3), 
WiHhOUL deseripnien, 


Stutighiphie Range. Por OQ) ventas. within the 
Ticalporites apeyvetinus  Zome ti within the 


Nothafavidties Microtlora, Conmeiin to Canpamin 
(Deumann & Playford 1969), From the Trreedperites 
apoxvextins Zone through Parcipites lorigus Zone. 
Santonian through Maastrichtian (Aelby ef al 18) 
Ornamenifera ch, OO. setosa, Conemanian (Norvick & 
Burger 1975: this study), 


Genus Perotrilites Ledtiman ex Couper 1953, emend. 
Evans L970 


Type species: (designated by Couper 1953) Mororeifites 
vranulatus Couper 1953 emend, Evans 1970) 


Perotrilitey jubatus (Dettmann & Playlond) Evans 1970 
FIGS 7L.M.8C 


Comments. This species is distinehve with two math 
Vidges bering spinose crests, running mere or less parallel 
to the trifere lisurae on the distal surlice (Cpseudomuri ol 
Norvick & Burser 1975). Spore body. 45-58 pin diameter, 
Zona, 25-30 pun wide, 


Stratigraphic Range. PAinepollenites penmoasis Zone, 
Lite Albian through Claviyere ripley Zone, Early Turoniu 
(Detimann & Playford 1969). 


Genus Plicatela Maljavkina Tt 


Type species: Pleaella trichacanthe Matjay kina 110. 
hy subsequent designation of Potonie [G0 


Plicatella dixtocerinita 
(Dettmann & Plavtord) Dayies }O8S 
HG, 4c, D 


Comments, Parallel muri occur on both distal und proai- 
mal surlyee. “The three sets of mut run parallel to the equa 
tor ind on the distal surface, coalesee ta form a fin-like pre 
jection ii the radiil region, height 5S pind and projecting 5-8 
yn beyond the equator. The muri are 2-4 pin Wide dnd the 
grooves, 13 pm, Equatorial diameter, 54-62 kum, 


Stratigraphic Range. Fron within the Captesycre pave 
doxa Zone do within the Plivllocladidites nidiwsenil Zone, 
Albiun through “Purontan, stirling cartier fi narthern 
Australia thar in the southeast (Helby ef g/, 1987), 


Geos Molvementarisporites Simonesics & Kedeves 
; ! . 
enend, Playford & De ina 1965 


lype species: Molveingulattsperites corculiy Simmonestes 
& Kecleves 1961 


Palveingulatisparnes sp. 
FIG, BAL B 


Descoiphon, Amb sabelreulir, tilete lasunie have shiek 
ened nmureins, 2-3 pin wie, wah rewular stviations about 2 
ben apant, Leuatarkal thickening 293 pin wide, distal supkice 
hears (Wo circular concentric ridges. Surface is psi 
Dinneter, 48 pn. 


Comments. A rare species ih this study. Species oF 
Palvciigilatispariies are more Cypieal Of the Jurassic, but 
some May be found i the Cretaceous (Phivtard & 
Detinain 1965. Hetby ef al CLOT). 


Genus Merenvidusperites Plug 1953 


Type species! Reticulaldasporitey deutdtiis (PAlag) Llue 
ast 


PALYNOLOGY OF THE POONARUNNA NO. | WELL 111 


Fig. 6. Cretaceous species continued. A. Ruffordiaspora australiensiy. B. Ruffordiaspora ludbrookiae. C-K. 
Cicatricosisporites sp. of Norvick & Burger, PL G, Sfercisporites pocockii. H. Cyathidites minor. 1. Cyathidites australis, 
J. K, Clavifera triplex. L. M. Crybelosporites punctanis. N. Dictyoplyllidites sp. O. Sestrosporites psuedoalveolatns. 
P. Gleicheniiditey cercinidites, Q. Foraminisporis wonthaggiensis. Scale bars = 10 pan. 


12 H. A. MARTIN 


Reflenloidasporites wenus (Balme) Dettinann 1963 
FIG. 8G 


Stratigraphic Range, Jurassic and Lower Cretaceous sed- 
dents (Delton 963) Cenomanian, this stidy, 


Genus Retiietliies van der Hammen ex Pierce emend. 
Doring. Krutzsch, Mat & Schultz in Krutzsch 1963 


Type species: Revitrileres globasuy Pierce 1961 


Remarks. Lyeopediumsporites has been restricted to 
forms with loveo-reticulie sculpture formed by pits closely 
spaced to form a reticulum. Reritileres aecommodiates a 
positive reliculite sculpture formed of raised muri (sec the 
discussion in Backhouse 1978). 


Reditriletes austreclavertidites (Cookson) Doring, 
Krutvsch. Mai & Schultz in Krutasch 1963 
FIG, SD-F 


Stratigraphic Range, Widely distributed in Jurassie and 
Creluevaus sediments. 


Cienus Kufferdiaspera Dettmann & Cliltord 1992 


Type species: Raffordiaxpora Gil Madirigisparites) ais- 
Haliensis (Cooksont Dettmann & Clifford 1992, by subse- 
quent desmnation af Dettmann & Chitford 192 


Kuffordiaspora australiensts 
(Cookson) Dettmann & Clifford 1992 
FIG, 6A 


Comments, The narrower muri distinguish this species 
trom &. fdbreakiae, The muri have straight edges, thus tt 
is distinctive from Cleatricosisparites sp. of Norvick & 
Burger (1975). 


Stitigeaphic Range. Prom the Nuffordiaspora australien- 
viy Zone, earliest Cretaceous (Helby er af TY87) te 
Clavifera triples Zone. early Coniacian (Detumann & 
Playford 1969). 


Ruffordeispora tidbrookiae 
(Dettmann) Dettnnann & Clifford 1992 
PIG, 6B 


Comments. The wider muri distinguish this species from 
© pustraliensin. 


Swratigraphic Range. From the Crvbelosporites stylasis 
Zone. earliest Cretaceous, lo the base of the Coptespure 
juiradoxa Zone, latest Aption-earliest Albin (Dettmann & 
Phiylord 1969), This species is rare in Lis study and sone: 
whit corroded: henee Wo may be re-worked, 

Genus Sesraspertties Dettmann 1963 


Type species: Cinewlatiyporites prendoeleulatas Couper 
1958 


Seatrapuriies predidodiwalainiy (Couper) Detar 1963 
FIG. 60 


Djaumeler, 35 pir, 


Stratigraphic Range, Upper Mesozoie of southeast 
Australie (Denman 1963). 


Canis Srereispariios PV (99 


Type species: Srereisporiies srereaides (Pole & 
Venité) Pilug 1953 


Sieveisporites aniiqueasporiies (Wilson & Webster) 
Detomann 1963 
FIG AN, O 


Comments, Pullers of low verrucae, <1 pt henht, 2-5 
um wide. creates negative reticulum on distal surliee, 
Pattera varies [rom barely perceptible (Mig, $O) on small 
specimens to conspicuous (Pig. SN). usually on lurger spee- 
imens. Equatorial diameter, 26-46 jm, 


Stratigraphic Range. Mesozoic and Tertiary strata, 


Sreisparties pacodckit Burger L980 
FIG. OF, G 


Stratigraphic Range, Vneommon in the Barly Cretaceous, 
Similar forms bave been found in the Cenomanian (Burger 
JO80), 


Genus Staveriyperiies Norvick & Burger 1975 


Type species: Sloveriyparttes aferaverrucaniy Norvick & 
Burger 1975 


Stoverisparites micraverrucatuy Norvick & Burzer 1975 
FIG, SM 


Comments. Creseentic shaped elevations which delinnt 
or partially enclose circular or elliptical shallow depres: 
sions are characteristic of the genus, This species dilfers 
from Staverisporites tintariy (Cookson & Dettmann) 
Novick & Burger 1975 in that So mierwerricatiy his 
Mmicroverrucate ornamentation. Diameter, 32-35 jim. 


Stratigraphic Range. Cenomanian of Bathurst Ishind 
(Norvick & Burger 1975), probubly the Albian of the 
Carpentaria Basin (Burger 1973). Cenomianian of Central 
Australia (thts study). 


Genus Trilehayporites Pant ex Potonié 1956 


Type species: Trilehosperites hanwenious (Delcourt & 
Spumont) Patonie 1956 


Trilebosporites tribotrys Dewmann 1903 
FIG, Bt 


Striligeupice Runge. Lower Cretaceous ta the Olwiay and 
Great Artesian Basins (Detimunn L963), Cenaniiinan. this 
stuicly 


Hilohosparites Woreticulosis Cookson & Detimiann 1958 
PIG. SH 


Statigeaphic Range. Fran the Coplasperit pandas 
Zone too within the Plicatella distocutinaia Zone, latest 
Aptian into eurly Cenomuntiia (Dettmann andl Play lord 
1969), Cenomanian of northern Australie (Noryick  & 
Burger (W753), 


Genus Friperaletes Michedlishvilt 1960 emend. 
Playlord 1971 


Type species: Jriporeleies simentarty Miehedlishyilh ih 
Mtchedlishyili & Samotavieh [960 


PALYNOLOGY OF THE POONARUNNA NO. | WELL 113 


B 


Fig. 7, Cretaceous species continued. A, B. Crybelosporites striatus. C. Crybelosporites punctatus. D, Foraminisporis 
dailvi, B, F. Foraminisporis wonthaggiensis, G. Laevigatosporites ovatus. H, L, Ornamentifera sp. cf. O. sentosa. J, K. 
Foveogleicheniidites conjossus. L, M. Perotriletes jubatus. Scale bars = 10 pm. 


V4 HOA MARTIN 


Triporoletes spel, To smmplex 
(Cookson & Dettmann) Playford 1971 
FIG 8] 


Comments. The specimens of this study. when compared 
with 7. simplex, have a thinner mner layer of the selerine, 
</ pm compared with 1.5-2.5 wm and the muroid ridges are 
nore Variable when compared with three radislly oriented 
rilges on 7) yiraplex. Diameter, 44-56 pm. 


Stratigraphic Range. For 72 sonplex. Albian and Aptian 
(Detimann 1963). Cenomanian, this sudy. 


Gymnospernis 
Genus Alisporites Daugherty 194] 
Type species: Afisporites api Dougherty 19d] 


Alisparites spect A. grandis (Cookson) Dertiminn 1963 
FIG OA 


Ciomments. The size tinge Of these Specimens is rather 
small when conpured with 78-136 x 36-70 wn for AL erei- 
iis, Overall size, 50-70 x 27-33 ym. 


Stratigraphic Range. Por A. grendis, Upper Jurasste and 
Lower Cretieeous (Dettinann 1963) inta Paleocene (Harris 
1965). 


Cienus Araweneacites Cookson ex Couper (953 


Type species: Araueariacies anytralix Cookson [O47 
designated by Couper 1953 


Araucariacites australis Cookson 1047 
fic. 9B 


Stratigraphic Range. Widely distributed m the Upper 
Mesozoic (Dettmann 1963) and in the Tertiary, to the pre- 
sent duy us species of Arevearia and Agartiis (Cookson & 
Duigin M951) See also the Tertiary specimens in Pies 120, 
J 


Genus Coralling Malyavkini emend, 
Coriet & Traverse bO75 


Selected Synonprny 
1953 Clhassapadiix Prag. 
Mor full svnonamy, see Cornet and Traverse (1975). 


Remarks. Classopaltis was originally deseribed as tricul- 
ponte and the original deseription of Careline wars vague 
sul adequate, With an emended deseription of Ceralfina. 
Classopalflis becomes a [UMOr Sy NOAY TN, 


‘Type species: Cyralling compacta Malyavkina }Ot9o 
Coralting spel OC. oldsseudes Pllug emend. Pocoek and 
Jiinsantis P96) conib. ney, 


PIG. 9C 


Comments, Rare al Poonarunna. 


Stuiigmiphie Range. Widely dispersed in Upper 
Mesovoie sediments (Dettmann 1863) Currlline 
(=Chassepollis) simples ind Corcllina (=Clayvapaltis) spp. 
are recorded from the Cenomani of northern Australis 
iNorviek & Burger 1975), Corgiing spp. aire lound from 
the Upper Tilassic-transidional to Lower Jurassic into the 
Maastrichtie, Upper Crtiecots (Melby et al. 1987). 


Genus Microcauchrvidites Cookson ex Couper 1953 


Type species: Microcuchryidites antarcticuy Cookson 
(947 
Mierocachevidites antarcticuy Cookson |Qb7 
HIG, YD 


Stratigraphic Range, Appears first po the Mipraypoea 
florida Zone of the Late Jurassic. [tis common through the 
Eurly Cretaceous and continues throuphout the Creticeous 
(Helby ef al. 1987) and the Tertiary to the present day iis 
the Tasmanian endemic. Mieroeacheyvs  lerragena. 
(Cookson & Pike L954a). See also the Tertiary specimen 
Fig. [2N. 


Genus Podocurpidires Cookson ex Couper 1954 
Type species, Podacarpidites ellipicus Cookson [O47 


Podocarpidites elliprieus Cookson 1947 
HIG, 9E 


Stratigraphic Range. Usually abunditt in the Jurassic andl 
Creiceous (Detunain 1965) and continues through the 
Tertiaey (Phutis 1965; Martin [973a) lo the present as 
Podeeurpus (vensi fet). 


Podocarpldiies exigius Harris 1965 
FIC), OF 


Stratigraphic Range. Cenvmanian, this study, Paleocene 
(Harris 1965) e0 F eafwii at Bungonia NSW, mid Bocene 
(Truswell & Owen JOS8) fate Eocene inthe bare Basin 
(Milne 1988). 


Genus Podesporites Ria (43a 
‘Type species: Podosportns oipakyii Raa 19d 


Podosparites sp. 
VIG. YITK 


Description, Grins Uisaccate. outline lenticular ta eineu- 
lin Cappie 2 pivthick. has fine, uniform reticulin, Cappula 
psikute, Subtriingular, Sacer hroully eresventic wry dis- 
Hiner radially arringed muri within saved, Muri reticulate at 
extronties OF savor Overall sive, 22-43 8 3045 yin, con 
pus, 25-35 wot sacet length 20625 jan, height and wach B 
10 jim, 


Comparisons, The fori is scolar tor Pecasporiios Obstes 
Phaskell M968. whieh. however bas a seabrate cuppa, con 
pared with the distinetly reticulate putter OF the specnnens 
my Unis study. Teis aso siinikie do Padespeeies teedivies 
Haskell 968 which also has a seahale eappu and irregular 
ly reticulate sieel. when Compared with the tadial disposi 
Linol the murtol the specimens of this study 

Comments. Superticially, this species resembles 
Lyyistepollentios florins in the pattern within (ie murh of 
Ihe sauce, but the dimensions af the sacer are much smaller 
than those of 4, flew ini, 


Genus Hriehetmonasiieies Couper 1953 emend, 
Dettmann Loss 


Type species: Hictanonasulcites subyruniianiy Couper 
1954 


Trichmentonosilopes subaranitaiis Couper 1953 
WG, OG) 


PALYNOLOGY OF THE POONARUNNA NO, | WELL 115 


Cc 


hig od. Cremeeous species caninued. A. B. Polycingulatispurites sp, C. Peratrileles jubatus. D-F. Retinrileres austaclava- 
fides, OL Rericiloidosporites ares. VL. Trilobosponites mioreniculosus. 1 Tilobosparites trinhotvs. J. Triparateres el 2 
simplex, IK. La. Microfeveolatasporines: eanetieulana, M. Sroverisparites aucroverricanis, Ny O. Stereisparites aariguans- 
porttes. Scale bars = 10 yan. 


116 H, A, MARTIN 


Stratigraphic Range. Barly Cretaceous (Dettmann 1963) 
(hrough the Miocene (Macphail 1996). [is morphological- 
ly similar to some modern species of Phyflodiadus, ound in 
Tasmania, New Zealand und elsewhere (Cookson & Pike 
19S4a). 


Angiasperms 
Cupuliferoidaepollenites Thomson & Thiergart 1950 


Type species: Cupuliferoideepallenites  liflaretsts 
Thomsen i Potonié, Thomson & Thiergart 150 


Cupuliferoidaepollenites sp. el, C. parvulis 
(Groot & Penny) Dettmann L974 
FIG, OL. 


Comments, The grains are prolate, tricalpate with slit- 
like colpi, extne < 1 unm thick, not clearly differentiated into 
nexine und sexine, with a scabrate surface. Size, 12% 10 
tun, 


Stratigraphie Range. Late Albian, Cenomanian, Turonian 
in dorthern Austtaha (Norvick & Barger 1975; Dettmann 
[973). 


Dicvalpapaltis Pllumel, 1956, emend. Potonié 1966 
lype species: Dicalpopollis kackeli Phlanal 1956 


Dicolpopellis sp. 
FIG. 9M, N 


Description. Amb more or less circular, hwo broad, rela 
lively short colpi, Exine 2 um thick with thin nesxine, robust 
columellae supporting a reticulum. In polar region, murus 
plus lumen fovether measure | yim. Luminat larger, up lo 2 
Lim in infercolpal region. Heads of the columeltlae distinc! 
under reticuluin, Size. 41% 26 um. 


Comments, Norvick & Rurger (1975) desenbe one dicol- 
pate type and figure two additional forms fron Bathurst 
Island. The type desertbed here is different from any otf 
(hose. 

Foveaeiradites Singh LORS 


‘Lype species: Foveorerradites fismlasas (Dettmann) 
Singh LOR 


Poveatetradites fiviniosus (Dettmann) Singh LORS 
PIG. LOA, B 


Side of tetrad, 42-58 pm. 

Stratigniphic Runge, Cenormanian of northern Australia 
(Detman 174) Norvick & Burger 1975) and central 
Austrabin this study, 

Liliieidites Couper 1953 
‘Type species: Liideleites ketitangareensis Couper 1953 


Liffacidites sp. cl. L. Ketteniyatecayis Couper 1Y53 
FIG. 9P 


Size, 42x 37 um, 
Struligeaphic Runge. Cenumanian of Bathurst and 


Melville Islands (Dettminn 1974, Norvick & Burger 1975) 
and of central Australia (this study), 


Liliacidites sp. 
FIG. 9S 


Description, Shape oval, monosuleate, sulcus extending 
length of grain. Exine <1 wm, nexine (hin, baculate/clavite 
columellae slender. Surface pattern scabrate, Size, 24 x 17 
tim. 

Distribution, Rare in the Cenomanian of this study. 


Phimopottenites Detemann l273 


Type species: Phimopollenites poamesus (Delimanit & 
Playlord) Dettmann L973 


Phimapollenites aaginhallaensiy (Burger) Detumuam 1974 
FIG. 90, Q. R 


Sive. 27% 40-33 po. 


Stratigraphic Range. Albian to Cenomunian (Burger 
1970: Detain 1973; Narvick & Burger 1975) 


Phimopollenites painosus 
(Detmunn & Playlord) Dettmann 1973 
FIG. 9U, W 

Sive. 12-15 x 15-20 pm. 

Stratigraphic Runge. Phinopallenties punnosuy “one, 
Late Albian. through Tricelperttes apexvexinous Zone, 
Santonian (Helby ey el! LYN7), 

Senectourradites Dettmann 1973 


Type species: Seneciolerradies varireticulatis Delhi 


1973 


Senecimevadiey varireneulaius Detimann 1973 
FIG, YT 


Size of tetrad, 30 x 55 pm, 

Straligruphic Range. Cenomanian of qorthern Australi 
(Dettmann 1973; Norviek & Burger 1975) iind: probably 
Cenomanian of the Otway Bustin (Dethuann 1974) 
Cenomanian of this study, 


Trivalporites Cookson ex Stover & Evans 1973 


Type species: Vriealperites sphaerica Cookson LV, 
designated by Stover & Evans (1974) 


Tricalperites sp. cl. To apexvexinis Partridge (987 
FIG. YX, ¥ 


Description. Shape almost spherical. long colpi real 
almost to poles. Colpi with well defined borders, pares with 
invegulay edges, exine | opin thick. with two layers af 
upproxtmately equal thickness. Surface smooth and faintly 
seabrate. Size. 14 pm polar diameter x [5 pm equatorial 
diameter. 


Comparisons, The grain of this stuady is similar to 7. 
dpexvecimies butit has a thinner exine and the nexine ts rel 
atively thinner than that ob 7) epaxve vines. 


Stratigraphic Range. 7. apowveninits, from the 
Tricolporites apexvexinus Zone, Santonwn, ime 


Nothafagidties senectus Zone, Campanian (Helby et af 
1987), 


PALYNOLOGY OF THE POONARUNNA NO, | WELL 7 


Fig. 9. Cretaceous species continued, A. Alisporiies sp. cf AL grandis, B. Araucariacites australis. C. Corolling spel © 
chasseides, Do Microcachrvadites antareticus, U. Pedocarpidites cllipticus, Po Padocarpiditey exigius. GC. 
Trichotomonosulenes subgranulatus. H-K. Podosporites sp. L. Cipdiferoidacpollenites sp. et. CL parvatus, M,N. 
Dicalpopollis sp. Arrows indieate colpi. OL Q. R. Phimopallenites augarhellaensis, Po Litiacidites sp. ct L. kaitani- 
sataensis, S. Liliacidites sp. T, Senectotetradites varireiteulatus, U-W. Phimapellenites pannosus, X,Y. Tricolperites sp. 
el 7 apoxvextnens, Seale bars = 10 pum, 


Hs H. A, MARTIN 


Microplankton 
Genus Horeloginella Cookson & Eisenack 1962 


Type species: Hordloginella lineata Cookson & Eisenack 
1962 


Harodloginella sp, 
FIG. LOL 

Description. Body rectangular with folds on the surface. 
cormers rounded, Long splits (20-22 kim) extend inwards 
from concave depressions on shorter sides of rectingle. One 
pore. 15 x Sum. with thickened cdge, in central position of 
one of Jonger sides, Wall < To um with granular 
striate Surface pallern, Size, 94d x 82 um. 


Conmnents. The body does not show any sign of tabula- 
tion, but the splits extending from the indentations are char 
acteristic of Horologinella, The specimen figured shows 
openings al the corners of the rectangle, but these are 
thought to be the result of damage. 


Genus Leciniella Cookson & Fisenack 1962 


‘Tvpe species: Lecaniella margostriata Cookson & 
Fisenack 1962 


Leeauniella sp. 
FIG. 10C-G, | 


Description, Cyst spherical, splitting mta two halves, 
halves fattening out {o saucer shape. Whole cysts (Fig. LOF, 
G) rarely seen. Each half bas outer marginal zone 5-7 pr 
Wide with radial striations and central rugulite-reticulale 
area. muri about | am wide, lumina 2 pm. Whole cyst, 39 
pin, halves, 28-62 pm. 


Comments, The unsplit cyst (Pigs 1OP, G) and one con- 
trucled hall (Fig. LOL) show a more distinet saiated murgin- 
al zone than the other specimens, but this is probably due to 
the state of contraction. The specimens here differ from 
Lecaniella margosiviaty which has a much coarser reucu- 
late pattern. Leceniella dteryora Cookson & Bisenack 1962 
has a finer reticulate pattern and the radial striations wre 
much courser than on these specimens. 


Lecaniella has probable affinities with the Zyenematacae 
(Grenfell 1995), This tamily of filamentous algae ts found 
mainly in shallow, flowing fresh water. 


Genus Saeprodiitm Uarris LOTS 
Type species: Sueplodiniuin gravattensiy Wartis 1973 


Sueptodinium eravatiensis Harris 1973 
FIG. (OH. 


Comments. Uncommon in Poonaruuna-1, 


Straugruphic Range Paleocene (Harris 1973), 


Cenomainian. this study. 
Genus Schizasporis Cookson & Dettmann 1959 


Type species: Schisesparis veticulatus Cookson & 
Dettmann 1973 


Schizosporis reticularis Cookson & Dettmann 1973 
TG, 10K 


Stratigraphic Range. Widely distributed in the Upper 
Mesozoic of eastern) Australia (Dettmann 1963). 
Cenomanian of northern Australia (Norvieck & Burger 
1975) and of this study. 


Comment, Seftisosporty refieriarus bas probable affini 
ties with the Zygnemataceae (Grenfell 1994), 


Acritarch. sp. indet. 
FIG, 10) 


Description, Quiline broadly elliptical, pylome oval, 15s 
10 um, near one end, Border of pylome psilale, 1.5 pm 
wide. Wall two layered. | pin thick. with patches of pitted. 
grooved. vertucate and broad linear thickenings. Size, 105 
x 75 um, 


Appendix 2 


Tertiary Systematic Palynology, For the distribution of the species in the bore, see Table 2, For the register of 


illustrated specimens, sce Tuble 4. 


Spores 


Genus Acedia Lam. 
Type species: Azella filiculoides Lam. 


Acalle sp. 
FIG. WI 


Comments. Massulae that have lost all the microspores 
have been found, 


Straugraphie Range. Probably from the beginniny of the 
Campanian (Hall 1974) to the present. Mid Eocene, this study: 


Genus Canirogonesporitey Pant ex Potonié cmend. 
Klaus 1960 


Type species: Camearezonosporites cretaceus (Weyland 
& Krieger) Potonié 1956 


PALY NOLOGY OF THE POONARUNNA NO. | WELL. 119 


Fig. 10. Cretaceous species continued. A, B, Foveotetradites fistulosus. C-G, I. Lecaniella sp. C-E, 1 are half cysts and 
F-G, a whole cyst with part of the filament attached. H. Saeptodinium gravattensis. J. Acritarch sp. indet. Arrow indicates 
pore. K. Schizosporis reticulatus. L. Horologinella sp. Scale bars = 10 um. 


120 ILA. MARTIN 


Camaraconeasporites anpls 
(Stanley) Detimann & Playford 1968 
FIG_HALB 


Stidivrphic Range. Upper Cretaceous and Paleocene 
(Detimiinn & Phiyfacd 968). 


Crmmtnconaspotiies bullatis Harris 1965 


FIG, ID 


Comments, The specimen illustrated his rirreawer erunsi- 
lndes and is slightly smaller than those deseribed by I ltrris 
(1965) and Detuvinin & Playford (1968). Size, spore becly 
28 um. overall. 43 pam, 


Stratigraphic Range. Late Cretaceous (Dettmann & 
Playford 1968) through Paleocene (Harris 1965), 


Ceneroconosparites sp. 


FIGS IE, P 


Description. Spore trilete, amb cireutar, lasurae straight, 
extending about) of radius, Exine 1-2 pn thick with inter 
ridigh erassitudes, 4-5 pn thick. Surface partern oF iiter 
locking rugalite } pm wide, with lumina T-2 pny wide and 
</ jm high, Equatorial diameter, 35 pm, 


Compariveus, The spore is smaller und the patteen much 
foer than That seen on Cameurmcenosporitos abulensls. 


Distribution, Lower A. asperis Zone Equivalent, mid 
Eocene (this stualy ). 


Genus Cvathidites Couper 1953 
Type species; Cyatiidifes qustralis Couper P53 
Cvathidites pulevspore 
(Martin) Alley & Broadbridge 1992 
FIG, Ill 


Suratigraphie Range, Throughout the Tertiary, Pound in 


the lame Paleocene, mid Eoeene and Pliocene-Pleistocene of 


this study 
Cyuthidites splendens Harris 1965 
PIG. WG, A 


Comments, The scubrite surhiee pattern is distingive and 
heeomes etched our on corroded specimens (Fig 111), 


Stuitigraphic Rane, Tate Paleocene and eauly Eocene 
(Hanis 1965, 1971). 


Genus Gleicheniidites Ross ex Deleourt & Sprumoni 
emend, Dettmann [963 


Type species; Gleiehenfidites senoniouy Rass (949 


Gleivhentidites eiremidites (Cookson) Demmann 163 
PIG. UC 


Siratigraphic Range. Jurassie (Helby er ad, 1987) be the 
presentoas the fern Gleielesite. 


Genus Grapnelispara Stover & Purtridue 1984 


Type species; Grapnelispora evansir Stover & Partridge 
{9X4 


Grapnelispare evansii Stover & Partridue 184 
FIG, 12A 


Comments, This beautifully preserved specunen has / 
appendages, One with a récurved-haok tip and the others 
with 3-4 short-hranched. recurved tips. Endospore SO yim. 
overall spare body 70 yin, length of appendages 90-100 jan) 
Tis found in dominantly non marine to marginal marine 
covinmments (Stover & Partridge 1984), 


Stratigraphic Rauge. Upper part of the fercipires 
(Tricolpitey) lengus Zone, middle to late Muastrichtian, 
with fost occurrences found in the latest Maastrichtian 
(Stover & Partridge 1984), late Paleocene (his study) pos 
sibly reworked, 


Genus Polypodiveoisparites Powe 195 | 
Type species: Polypodidenisporties speciosus Potonid [934 


Palypodincoisporites sp. et. Foretirivvettis Muller 1968 
FIGS IIL-M 


Comments, This specimen is) very similar ty 
Polypadinevisporites sp. ek Po retirngatis os described by 
Truswell eral. (1985). It is similar to Preriy uirhresune anil 
Preris tenia of the Preridaceae (Martin & MeMinn 1993), 
Sive, 45 pum, 


Stratigraphic Range, Late Oligoeene lo eurly-mic 
Miocene (Truswell er al. 1985). late Miocene into 
Pleistocene (Martin ak MeMinn (993). Phoecene. 
Pleistocene (his study}, 


Genus Refiirdees van der Mammen ex Pioree emenil 
Doring, Kratyseh, Mai and Sehulty 1963 


‘Type species; Kerirtleres globosus Pieree 1961 


Retitriletes ausipaclavatidites 
(Cookson) Potonié 1956 comb. noy, 
PIG. 128.0 


Description. Lasurae wah psilate berder upto 4 am wide, 
extend (fy) spore radius. Proximal surface psikie or with 
faint, raditting ridges, Distal surface reticulum bes muel | 
Hor high, 0.5 po wide aid linia 3-7 pnt in diameter, xine 
J pin thick, excluding reticulum, Dianieter 30-35 jin 


Stralivraphic Ringe. Species of Revitrifeles are signit 
cant in the Cretaceous but a few iy be found through 
much of the Tertiary, 


Genus /ifetey Erdtman ex Couperomend. Detain 193 


Type species: Triletes tubereniifarnos Cookson |Y47 
emend, Dettmann 163 


Triletey sp. ck To aibereutifuris 
Cookson 1947 emend, Dettmann 164 
FIG, Nk 


Desenpion, Outline triangular, Wilete lasurae extendiny 
for mostol the nulius, Bxine |p thick, bearing lurce ver 
rucie/rugulae, 3 pin high inter-radially, up to G pum high al 
upives, Distal surface pattern coarse rugulite-reticulum, 
muri 3-5 pm high, laming abeut Sun diameter, Proximal 
surfice pattern simihee but with lower more widely spaced 
wements. Size range, 58-64 pn, 


Distribution, f. balmed Zone Equivalent, lite Paleoeene 
and Lower Vo aspen Zone Equivalent, mid Eocene (this 
study), 


PALYNOLOGY OF THE POONARUNNA NO, | WELL 121 
Cc 


Fig. VI. Tertiary species. A, B, Camarozenesporites amplus. C. Gleicheniidiies circinidites. D. Camarozonosporites 
bullatus, E, F Camarezonosporites sp. WH, G. Cyathidites splendens. 1. Cyathidites paleospora. J. Azolla sp, K. Triletes 
spc To tubereuliformis. L. M. Polypodiacoisporites sp. ct Po reriraganis, Seale bars = 10 ym. 


122 HA, MARTIN 


Genus Triperoleres Mtchedlishvili emend. Playford 1971 


Type species: Triperoletes sinaulariy Michedlishyili in 
Mtvhedlishvili & Samoilovich 1960, 


Triporoletes reticulatas (Pocock) Playtord 1971 
FIG, 12F 


Strazraphic Range. Early Cretaceous (Dettmann 1963), 
Cenomanian (Norvick & Burger 1975). Tertiary (this 
study). 

Gymnosperm pollen 


Genus Araucuriacites Cookson ex Couper 1953 


Type species: Arauedrecites anstralis Cookson 947 ex 
Couper 1953 


Araucariacites australis Cookson (947 
FIG, 121 


Stratigraphic Ranwe, See this species in Appendis |, 


Cupressaceac/Taxodiaceae 
FIG. 12D 
Stratigraphic Range, Late Paleocene, (this study) to late 
Tertiary (Martin 1973a) and the present day. Macrofossil 
studies (Peters & Christophe! 1978) show that both of these 
families predate the Paleocene, 


Genus Daeryvearpites Cookson & Pike 1953 


Type species! Daervearpites australiensiy Cookson & 
Pike 1953 


Decrvearpites austratiensis Cooksan & Pike L953 
PIG. 121 


Stratigraphic Range. Late Paleocene (this study) to late 
Tertiary (Martin [973a), 


Genus Dilwyaites Marris 1965 
‘Type species; Dilwyriter eranalatis Harris 1965 


Dilwyntes granuleatas Harris (965 
FIGS 123. KR. 13A 


Suraligraphic Range. Maasirichtian through Miocene 
(Stover & Partridge 1973; Partridge 1976). 


Genus Ephedra 1. 1753 
Type species: Aphedrea distachyva 1. 1753 


‘hphedra’ notensis Cookson 1956 
FIG. LB 


Comments, ‘The fossil is similar to Ephedra (Cookson 
1956), which, however. is much largen A very similar 
pollen morphology is found in the Araceae (Martin 19734), 


Stratigraphic Range. Karly Cretaceous to Eocene 
(Cookson 1956). carly Miocene (Martin 1973a, LO84b) in 
Australia, Miocene in New Zealand (Mildenhall & 
Pocknall 1989). 


Genus Lygisrepollenites Stover & Evans 1973 


‘Type species: Lyvistepolleniles florinié (Cookson & Pike} 
Stover & Evans 1973 


Lygistepollenites florinii 
(Cookson & Pike) Stover & Evans 1973 
FIG. 12G 


Stratigraphic Range. From within the Phyvllocleadidites 
mawsontii Zone, the basal portion of Santonian (Melby ef al. 
1987, Fig. 2). widespread throughout the Tertiary, to 
Pleistocene, about 26,000) years ago on the Atherton 
Tableland (Kershaw 1985). 


Genus Microcachevidires Cookson es Couper 1953 


_ Type species: Micracachrvidites antares Cookson ex 
Couper 1953 


Micrecachryidites amarcticus Cookson ex Couper 1953 
FIG. 12N 


Comments, See this species in Appendix |. 


Genus Piyllocladidites Cooksun ex Couper emend, 
Stover & Eyans 1973 


Type species: Phvllocladidites mawsonti Cookson 1947 
ex Couper 1953 


Phyllocladidites mawyenti Cookson ex Couper 1953 
HIG, 12k 


Stratigraphic Range. The base of the Proreacidires 
Superzone, Late Cretaceous (Helby ef al. 187, big, 2). 
through most of the Tertiary on maintand Australia, to the 
present day in Tasmania as Lagerastroboy franklinil, the 
Huon Pine (Playford & Detrmann 1974). 


Phyllacladidites rericilasaccinis Uartis L965 
MIG. 12H 


Stratigraphie Range. Paleocene (farris 1965; Stover & 
Partridge [973) 


Genus Podocarpidites Cookson ex Couper 1953 
‘Type species: Padoecarpidites ellipticus Cookson |947 


Podacarpidites exiguus Harris 1965 
FIG. 12M 


Stratigraphic Range. Cenomantin (this study), Paleocene 
(Harris 1965), mid Gocene (Truswell & Owen 198K: this 
study), late Eocene (Milne 1988). 


Angiosperm pollen 


Genus Acuciupolleniies (Cookson) Mildentiall emend 
Mildenhall & Pocknall [O80 


Type species: Acaciapollenites myrlosparites (Cookson) 
Mildenhall 1972 


Acactapollenites myrtosporites 
(Cookson) Mildenhall 1972 
PIG, 13C 


Stratigriphic Range, Late Oligocene (Truswell er al, 
1985), but usually early Miovene (Stoyer & Partridge 1973) 
to present day, as Acacia, Found only in the Pliocene 
Pleistocene of this study, 


PALYNOLOGY OF THE POONARUNNA NO. 1 WELL 123 


Fig. 12. Tertiary species continued, A. Graprelixpora evansii. B.C. Retitriletes anstroctavatatus. DD. 
Cupressiceue/Taxodiaccae. E, Phylloctadidites mawsonii. PF. Triporoletes reticnlatny, G. Lygistepollenites florinii, Hi. 
Phyllocladidites reticulosaccatus. 1. Araucariacites australis, J, K. Dilwynites granulatus. L. Dacrycarpites austratien 
giv. M. Podovarpidites exiguus. N. Microcachryidites antarcticus. Scale bars = 10 yim. 


Ia HA. MARTIN 


Genus Agluoridia Erdman 1960 
Type species: Aglveridia cyclops Erdman 1960 


Aglaoridie qnalianiy Partridge in Stover & Partridge 1973 
FIG, 17G 


Stratigraphic Range, Southeast Australia, mid Boeene ty 
varly Oligocene (Stover & Partridge 1973). mid’ Eocene ta 
carly Pliovene (Macphail 1996), Lake Eyre Basin, early- 
Wid Bocene (Sluiter 1991), 


Genus Amasopallis Cookson & Balme |962 


Type species: diavopellis eruciarmis Cookson & 
Bale 1962 


Amosopollis difwynensis Harris [972 
FIG, [38-G 


Deseription, “Tetrads rhombiodal, asually crumpled 6 
some degree, Lach grain has distal sitlets extending most ol 
the diameter of the gun. Exine < | un. psilale with patel 
of granules <1 pm diameter aver aren around sulcus, Some 
oralbotthe granules may be missing an less well preserved 
specimens. Size range, 57-75 Ln diameter of tetrad, 


Stratighiphie Runge. Paleacene (Hartis 1972, this study). 
Genus Areeipites Wodehouse emend, Anderson 1960 


Type species: Arecipitey pumretaiiy Wodehouse ex 
Potonie 1958 


Arecipites spo cl Ay nitutiscahraltes 
(Melntyre) Milne 198% 
FIG. 13D 


Description, Grain elliptical, monosulente, suleus extend 
ing moslal the length of the grain, ends rounded, Exine | 
Him, levtite with minute perforations. thickness cndesine. 
velexine upproximately equal Surfaee finely scabrate, 
sparse graniles [0.5 pot on distal surfiice, Size, 25 8 17 pn, 


Comments. The sparse vrunules on the distal surfiee are 
nob seen on A. minifiveubracs. 


Stratigraphic Runge. Por A. winnitveabratiy. Paleooene 
(MeIntyre (968), lates Rovene (Milne 1988), 


Genus Austratapelfiy Krutesch 1966 


Type species: daatrdopattty abseurus (huris) Krutzseh 
1966 


Australopotlis vbsenras (Harris) Krutzsh eniend. 
Stover & Partridge 1973 
FIG. 13N,Q 


Stiitigraphic Range. Cenomunian through Paleocene 
(Helby ef af }O87, Stover & Partridge 1973), 


Genus Beaupreaidies Cookson ex Couper 1953 


Type species: Beanpreaidites elegunsiformis Cookson 
1950) 


Beaupreaiditey elegansifornis Cookson L950 
biG. IAs, Ke 


Stratigraphic Range. Infrequent in the early lo late 
Maustrichtiin ar earliest Danian in the Otway, Bight and 


Duntroon Basins (Dettmann & Jarzen 1996) through 
Miocene (Stover & Partridge 1973), Fora full aecount at its 
distribution, see Dettmann & Jarzen (1996), 


Genus Chenopodipoalliy Keutesch 1966 


Type species: Chenopodipaltiy nudtiplex (Weyland & 
Pflug) Krutzsch L966 


Chantopodipollis chenupridinceoides 
(Martin) Truswell er af. 1985 
WIG, 13L 


Stratigraphic Bange, arly Oliocene (Muephail & 
Vruswell 1989) to the present in the families 
Chenopodiieewe and Amaranthiceae, Pound only i the 
Pllovene-Pleistocene OF this study. 


Cunoniiecae 


FIG, 1aS-V 


Description, Grains bie or (ri-colpale or colporate, finely 
reliculide, very sroall. dbout (0 pm. Triealporaie type (Pig. 
138, T) mast common, bicolpate grains (Pig, 13UL Vi few. 
Three fossil types attributed ta the family (Luly er af, 1980) 
Size range, & TL um x 7-9 pin, 


Comments. Mildenhall & Poeknall (1989) deseribe 
Tricalpiles inconspicuous and alribute it (oO Cunoniacene, 
as well as possibly several other families. lhe size range, 
however, is Girger, 17-26 pi x 1TO-TS pine compared! wath 
That wbove: hence this form species is inuppropriate for 
These specunens, Modern species Callicomed servatitelid 
(Fig. TAY, bicolpate) and Cermophvtian virchowil (Pig, 
JAW. X. (ricolpitted are given for comparison, 


Stitigraphic Range, Late Paleocene (Sluiter Y9 1) to the 
presear in the eustern coastal fardforest, North Queeastan 
to Tasman, 


Genus Cyperuevaepolliy Krutasch (970 


Type speeies: Cyperaceaepallis, neagenicus Krutyseh 
1970 


Cyperaceaepolliy sp. 


FIG. 13M 


Comments, Phe grain ts (nanwzular in shape with ane pore 
und a fine seabrate pattern. For descriptions of song mad- 
er Cypernmeae pollen types and the vurivlion found in the 
finily. see Tseny-Chieng (1972) and Heusser (1971), Most 
specimens ure folded or crumpled. Size range, 30-37 pam. 


Straligraphie Runge, Lite Paleovene (Sluiter 199}; Alley 
ecu, 1996) to the present as the lamily Cyperaecac. 


Elteocurpuceae 
KG, 134, AA 


Comments. The grains are small, triculporate with thin 
walls. < 1 yim, with @ psilite surtace (Luly ef af 19s) 
Because of their small size, hey may be dilficull (oO separate 
from Cunontaceae. Fig. | ABB is modern Elaeocarpus ren: 
wlaius, Size, equatorial view, 9 x S pm. polar view, 11 pm. 


Stratigraphic Range. The sume as that ol Cunoniucede. 


Genus Ericipites Wolehouse 19233 


PALYNOLOGY OF THE POONARUNNA NO, | WELL 125 


Vig. 13. Tertiary species continued, A. Diliwynites granulaius. B. Ephedra’ notensis. C, Acactapoallenites myriosporites. D. 
Atecipites sp, ch A. ainitiscabratus. EB. FG, Amasespollis difwyuites, UW, 1. Stupsanipolis sp. J, K. Beaupreaidiies 
elevansifarmis. L. Chenopodipollis chenopodiaceuides. M. Cyperaceaepallis sp. N. O. Australopalliy obsenruy. P,Q. 
Lriviptes scubratus, R, Haloragacidites halorageides. So T. Cunoniaceac. ticolporate form, U,V. Cunoniicese, bigolpate 
form, WLOX, Modern Cunoniaceae, Cereteperaliin virchow?i, a tricolporate form, ¥. Modern Cunoniaceae, Callleener ser- 
ratifolia, a biculpate form. Z, AA. Elacocarpaceac. BB. Modern Elaeecarpua reticular. Scale bars = 10 um, 


126 HA, MARTIN 


‘Type species: Erieipites Jongisileaus Wodehouse 1933 


Frieipites scabrams Harns 1965 
HIG. 13P,Q 


Stratigraphic Range. Prom the late Paleocene i south- 
custern Australia (Harris 1965), late Paleocene tn central 
Australia (Sluiter 199] ), 


Genus Goianipallis Krulasch [So 
[ype species: Gothanipollis gethid Krutzsch 1959 


Goluipollis bassenvis Stover in Stayer & Partridge |973 
PIG. I4A,B 


Stratwvaphic Range, Middle Locene to middle Miocene 
(Stover & Partridge 19734, Macphail & ‘Traswell 1989), hate 
Paleovene (his study). 


Genus Haloragacidites Couper 1954 
Type species: Helaredgacidites iriavatus Couper 1953 


Halorigacidites haveist (Couper) blarris 1971 
IG. 1G 


Comments, The Paleocene specimens ure smaller, with 
poves Hat hardly protide whea compared With those of the 
Lovene. bul they fit the species well, 


Stroligraphic Range. Paleocene (Stover &  Partridue 
1973) (othe present ay Cusuarinicese, 


Haloragaecidites Haloragaides Cooksan & Pike 1954 
FIG, JAR 


Stratigraphic Range, Lite Miovene (Stover & Partridee 
l97F to the present as Ganeepus/Halondgiy (Cookson & 
Pike [954h), This sometiines recorded i carly Miocene and 
possibly older sediments, eg. Tulip ee al (982) and 
Hriswell ef af (1985). Pound only in the Pliocene- 
Pleistocene of (his study, 


Crenus Hevpollenites Uhiergart 1937 


Type species: expatlenitey ecus (Potonig) Thiergart 
1947 


Hexpotlenies aneiloctavatus Metatyre (968 
TG, 4D, 6 


Comments. The size, shape and density of the seulpturing 
are highly variable (Stover & Partridge 1974). This spec 
Wen has relitively somlbscalpriring Size, 25 6 19 jin 


Stratigriphic Ronge. Maastrichtin to Oligocene (belly 
et adh JOT: Stover & Partridge LOA) i southeastern 
Austin, Phe diving geri Mev produces His pollen type 
yndis Founh in porthern Austailia today (Murtin Lo77) 


Genus Lenvetaupotlis Dethnann & Rurzen 196 


Type species: Lewelaiipollis fvcteros Dettomnn a 
Jaren L996 


Leiididiipallis spoch bk. tectaiiigiaiy (oouksait) 
etm de Nirven 1996 
HIG. baa 


Description, Sides staight toslightly cancaye, pures 9 || 


wou diamicter, Exine 5S pm thick, thing considerably 
towards pores. Nexine 3 um thick, becoming thinner in 
vome around pore. with faint channeling. Fine columelhic 
support irregular verrucae up to 5 um diameter, Surface pal- 
tern around pores granular, Size. 55 pm, 


Comments. The pattern is similar to that of L. rectani- 
ginis, but it lacks the distinctive disaggregation of the nex- 
ine around pore of the latter. 


Stratigraphic Range. For 2. reefomarginix, Campanian (0 
Maastichtian or earliest Danian in the Onway Bast 
(Detumunn & Jurzen 1996), middle Eocene ita hate 
Miovene (Stover & Partridge 1973). ef. 2. reermierginity, 
late Paleoeene (this study). 


Lewalanipallis speek Persons 
FIG. I5d_K 


Descnphon. Grain irresularly square wilh) + pores 5 pin 
diameter, Exine 1 pum thick with three layers of approxi- 
mately equal thickness. Middle layer has very fine, hardly 
distinguishable columellae, Surface patierm very finely 
seubriite with larger more widely spaced ‘granules’. 
Granules not visible in opligal section, hence they may be 
tiny perlorations through twetum. Middle layer of caine 
thins towards pores. ‘Granules’ more conspicuous on thine 
nerexing around the pores, Size. 38 pm. 


Conments, This ype of grain is found in Perseanii Wig, 
ISL. M), which however, is triangular with 3 pores. Four- 
Porcd LANs ure SOMETAGS sven in species Ol Protencce 
Conospermiun also fas an exing thinning towards the pores 
bur iasmnel lager and thicker-salled than Persoonti. 


Distribution, Mid Eovene (this study), 


: Genus Lilacsdites Couper 1953 
Type species: Lileedlires Kaimineataensis Couper [983 


Lilaeiditey hinceolans Stover in Stover & Partridge 1973 
MiG. 14h 


Strittigraphie Binge, Latest Paleocene through Miocene 
(Stover & Partridge 1974: Muruidge 1970), Late Paleoeene 
(this stucly) 

Cienus Malwiripollis Urns 1865 

Type specigs: Mafvacipolliy diversus tharris [05 


Mualyacipalis sabilliy Stover & Paciridee [973 
Pia. dH 


Comments. The species is desenhed as 4oni- ar slephnee 
porate (hertris 63; Stover & Partridge 1973, respectively) 
but same Of these specimens are panperite, 


Stratigraphic Range. Late Paleocene through Miocene 
(Harris 1905; Stover & Purtridpe 1973). 


Genus Milferdia Bedimian (960) 
Type species: Miffondio hypalaenaddes Ercinio [eth 
Milfardia homeaprinetate 
(Metityre) Parties i Stover & Parnidtge [973 
biG. 14. I 


Contents, This species has the small anoncliute Revira 
(ype of pore, Pig, 14 hs a thickened annulus 2 pun wie, 


PALYNOLOGY OF THE POONARUNNA NO. | WELL 127 


Fig. 14. Tertiary species continued, A, B. Gothanipollis bassensis, C, J. Milfordia homeopunctata. Arrow indicates pore. D, 
E. Hexpollenites anguloclavatus. F. Liliacidites lanceolatus. G. Haloragacidites harrisii. H. 1. Malvacipollis diversus. K. 
Milfordia hypolaenoides, L. Nothofagidites emarcidus. M. Nothofagidites falcatus. N. Nothofagidites deminutus. O, P. 
Nuxipollenites kempii. Q, R. Myrtaceidites eucalyptoides. S. Myrtaceidites verrucosus. T, W. Propylipollis ivanhoensis, 
U,V. Propylipollis sp. ct. PB. reticuloscabratus. X. Propylipollis latrobensis. Y, Z, AA. Propylipollis sp. BB. CC. 
Polyorificites oblatus. Scale bars = 10 tum. 


128 HA. MARTIN 


Whereas the pore on Fig. 14d Carraw) is not thickened. Size, 
34-47 ui. 


Strulivraphic Ruaowe Early Bovene through Middle 


Miocene (Stover & Partridge 1973), late Paleocene (Stuiter 


199]). 


Millontio liypolienoides Kedtman 1960 
PIG, 14k 


Comments. This species Nas the distinclive scalwate (pute 
(erp wilh Jurger pits, typical of the fantily Restionaceac, and 
a hinge Aypalaena type pore with granules aligned alone 
the tang, Size, 30-40) pin 


Stratigraphic Runge; Lule Paleocene (his study) to the 
present, 


Genus Mvrtaedidiy Cookson & Pike amend, 
Potonie 1960 


‘Type species! Myrtaceidites mesanesay Cookson & Pike 
1954 


Myrraceuiteys enealyproides Cookson & Pike [954 
FIG, 14Q.R 


Comments, this pollen type is found in the 
Ancophorafblondwood wroup of Rivelyrea. There are, 
however five other groups within the genus (Chialson & 
Martin 1995), Size. 17-19 jim, 


Sttigruphie Rage. From the carly Eoeene (Alley eal 
199) 10 present (Martin 1094), Round oily in the Pliocene- 
Pleistovene of this study. Sluiter (MOE) records 
Mvyilocemdiloys spp. titoueh the late Paleocene, curly and 
nid Bocene of the Lake Byre Basin, bot the species are ot 
differentiated, 


Muyrtaretaites Veriicisus 
Partridee ty Stover & Partridue (973 
tiG, b4S 


Stufigraphic Range Barly boeene through Miovene 
(Stover & Variridge 1973; Macphail }96), carly Eocene 
(Suter 1991}, mid Eocene (his stuily), 


Cents Nerhofueuirey (Bedtmun) Potonié [You 
Nowoluadicn: emmen (Couper) Potonie 


lype species: 
[You 


Noalaeniiics ennwetdiy (ooksorny bhirtis 1465 
Pen bb 


Statiaphic Range. Marky Eocene through Mincere 
(Stayer & Parriduy TOT 4). mid Moeene. Chis stucly ). 


NVethofeuidites faleains (Cookson) Stover k Bvans 1979 
PIG, 14M 


Stulizraphic Range, Mid Boeeoe through mid Miacene 
(Stover & Parinidie “\O73), 


Nathofauitiies deaianiis (Coakson) Stover & Evans b973 
FIG, LAN 


Comments, The U-shaped colph witht miurgins narrowly 
rivivied by inrotled oxime anck all Other feitures of the mor- 
pliohony fie hose of Novomdinras CDenniol & Pockiall 
(990), Sive, 22-27 pine 


Stratigraphic Range. Farly Rocene (Stover & Evans 
1973) ite carly Miocene (Cookson 1959), 


Genus Niavipolleiies Elsik emend. Frederiksan 1980 
Type species: Nivipollenizey eluirhoinensig Elsie L924 


Naxipallenites kempit sp.nov, 
FIG. 140, P 


1976 Trivolporttes sp. all, Diplopeltiy Kemp, po 114, Tig. 
di Wy 

JO8T Tricalparites sp. ull, Diplopeltis Bint, p. 286, figs 
34, $5. 

(ORS Nivipellenites sp, Truswell ef al. p. 286, fig, be, | 

1989 Nixipattemtos sp. all) Diplapeliis Mildenhall & 
Poeknall, p. 47. ph ll figs 1, 3 

DORKS Mavipallevtites sp. Macphail & Troswell p. 327. lig. 
Ob Ib, 

(994 Dadonied tice pollen type Martin, p. 28. tig. 
FAV (ajetk). 


Molowpe: Slide AMP }03724, Lnadlind binder coord 
mutes O/T (Pablo 4d), Grain in equittorial view, 29 4 20 pur, 
Fig. 140, P 


Etymology: Named in honour of B. M, Truswell (née 
Kemp) who first documented this species, 


Diagnosis, Grains prolate. (ricolporate with protruding 
pores. Distinctive thickened titereolpal bands meet at the 
poles, Exineg in outline thinner at equator, thicker towards 
poles. 2 layered with thin columethite hiyer in between, col 
tinellac barely visible: Oily the outer layer thickened to 
produce intereal pal Beads, 


Description, Girdins pwolate, poles broadly rounded. tr 
colporate with long colpi extending alaist to poles, Pores 
3.44 pn protrude, transverse margins thickened, Lxine [-2 
Heh at equator, bas two cistinel kiyers with thin, line. 
columellite layer bebween them, Columellie not always 
visible. Poles 3 um thick Where distinehve thickened jer 
colpal bunds meet Only outer hier of exine becomes 
thickened, inner and columeliae layer remaining same 
thickness over whole grain, Surfuce pattern seabrate. Sie 
minge, 29-40 4 15-30 ui. polur x equatorial dhumetur, 
respectively (O specimens), 


Comment Kenip (1976) uliibuted this pollen iype to 
Diplopeltis Badl, (Cieore & Erdtian (969) Unlike WV 
Aempii. the esine over the poles of 4 at ihe S spevies al 
Diplopeltis is either the sume thickness over the whole of 
the grain or itis thinner than over the rest of the grain, The 
filth species has pormed poles where the extie is Mieker 
than the rest ol the grain. whieh otherwise bas a unilorn 
ihickness. The columetlae are well defined in Diplopetin 
aod in soine species, extend throughour the thekened part 
ol the exine. unlike V. hempil. Dodanued jmiguatha bas Vary 
similar thickened intercolpal bands meeting al (he poles and 
‘fine. thin columethite layer dike No kempii The fossil is 
thus very similar ta 2, reiguerre and Jess like Diplapettis. 
Daeleniaent Phjuetra ts the only species ia the genus with 
this pollen morphology. and is found in cucalypt forests oF 
daimp sites und in gullies along the southern hall at the cast 
ern Coashil stvip of Austratia (Marlin loud (UOT), 


Swratigeaphie Ringe. Mid Eovene of central Australia 
(Kemp 1976; Stuiter 1941) to rhe present day, on fhe south 
em part ab the cast cowl OMbartin (99). Found only in the 
Pliocene-Pleistocene of this study, 


PALYNOLOGY OF THE POONARUNNA NO, | WELL 129 


Nig. (5, Tertiary species continued. A, B. Proteacidties adenunthoides, C.D. Proteacidites crassus, E, B Proteacidites 
framensix. GUL Proteactedites augulaius, 1 Proteacidites wrandis. J. K. Lewalanipotlis sp, ef. Persoonia. tL, M. Modern 
Porsoonis lamina. N. Proteacieites tnenrveaius. Seale burs = 10 urn. 


1a) HA, MARTIN 


Genus Polvermfirites Martin (Y730 
Type species: Polverificites oblanis Martin 19730 


Polverificitey oblatus Martin 1973 
VIG. I4BB. CC 


January L9?Sa Polvarifienes ables Martin, p. 45, ties 
1OQ. TON, 

May LOTS. Heliciperites usirus Partridge in Staver & 
Partridge, p. 27) pl. 26, figs 3-5, 


Stratigraphic Range. Early to late Eocene (Stover & 
Partridge 1973), late Focene to mid Miocene [Martin 
IUX7), wd Rocvene inte Pleistocene (Macphail 1996). 


Geos Polvparine Naumova es Potonie 1960 


Type species: Palvporine mtidtivtivnosea Naumov ex 
Potorme P60 


Polyporing sranulara Martin (97a 
KIG. ITACB 


Description. Grain spherical, panporate with about 24 
pores, 2-3 yin diameter. Exine 2 pm thick. with sexine, col- 
urmellate kiyer anc tectum of approximately equal thick- 
hess, ‘Teetum hats numerous perforatious 0,5 Lum wide, 
spaced about | pm apart Surkice has scattered weanitles, 
Siatlur to same Amuranthuceae, Size. 37 pum. 


Stiiivraphic Rungwe, Date Miocene tito Pleistocene 


(Macphail 1996), Pound only inthe Pliocene Pleistoeenyg of 


this stucly, 


Genus Propylipedis Martin & Parris 1974 


Type species: Propyiipalfis refieuloseabrajuy (Llareis) 
Martar cd bharris (924 


Prapylipollis ivanhoensis (Martin) Milne Loss 
FIG. 141, W 


Description. Sides siraight, apives truncate, pores 4-7 yn, 
Histhily about 5 um, neaiie and sexine approx unmarely eyial 
thickness, wide very Tinely volumellite liver Sunfuce pat 
tern finely scabrate with scattered granules and/or tovalte. 
Pores have well-defined moditiert zone where surtace puit- 
tern is tpere grunulir and nexine is moditied. Sive, 23-30 
yin, 


Comments. These specimens are more variible, espectu- 
yin size. aolin the thickening. than these of the oreinal 
deseriprion (Vbarii 197 $i), can Pie. PAW witli catich hare. 
wr opore “Phis petien type is simian te that al species of 
Heliila, Groey and Macoeemic Tound iy east cost ein 
Lorests, 


Shatitruphig Range bate Pogene (Milne POssi, kite 
ovene through Phocene (Martin $987). late Paleouene 
({thissdughy ), 


fropylipedix datebonsis (Harris) Main & Flows PO74 
BIG TS 


Strtiriphre Kame. Larly to hite Bocene tstover & 
Partnidge P9771 lane Paleocene CHnurIy TYOS. His study). 


Pray tipallis sp ch Po pyereomreateden 
(Stover) Delumann & Kuegen 1996 
Hii. 1OB-l) 


Comments. Some specunens, ee, Pig. LOC. fit the diag- 
nosis, with the exception that tiny columellae mity just be 
visible on the coarsest part of the reticulum and the size, 
ninge here is a little larger than the 27-36 um of Staver & 
Partridge (1973), Others, e.g. Fig. 16B, D. have a pattern 
thal ds punctorreticulate. with the dimensions af the lumina 
and imuri less than OLS pi. the lower limit for Po pyeueda- 
Moides, Size, 30-44 pm. 


Stratigraphic Range. For 2 pyerndinmoides, Campunian- 
Maustrichtian or earliest Danian al the Otway Basin 
(Dettmann & Jarzen 1996), elsewhere in southeastern 
Austedtia, late Paleocene into Oljgocenc (Stover & 
Partridge 1973). 


Proplipollis sp. cl. Po reticntosesshratins 
(Haerist Martin & Parris 1974 
FIG. IHU V 


Comments. This form is a delicate version al rere 
loscuibranis where the exineg ts thickened uronnd the pore 
but notas heavily as those of Haeris (1965), aad dhe rele. 
Jum is finer. Size, 25-35: un, 


Strubgraphic Range Por Po fetinloseabreties. 
Campiniine Maastrichdan of the Owe Basin (Datta 
Jarzen 1996). Southeastern Australi, lite Paleouene tity 
Tate Kocene (Hartis 1965: Stover & Partridee 1974) ch 2 
reticulascabratis, lule Paleocene (his stds), 


Prapvlipallis sp. 
PIG, 14Y, 4, AA 


Deseription, Shape triangular with protruding. domed 
pores. Exine T-2 pm, sesame and nexine apprexaenately 
equal thickness. [rrevular, broken reticulate pattern. muri 
< J pint, lumina 1-2 pm. Small columetlae mity just he ota 
ble. Pores 2-3 pm diameter, protrude 2-8 pun, Nexinae thins 
lowards pore, hase al pore may be marked by a toteh or 
inward protrusion, Size, 20-30 pun, 


Conimernts. ‘The distoetive pore is seen in present day 
species Of Bankiin and Grevillea, The sive. however is 
much smaller that these living taxa fh is untrke 
Proteacidiies hakenides Couper which ts much hirger than 
these specimens and has a codrse-grinular pater GABE 
Martin 1973), 


Distribution, Lake Paleocene (this study) 


Cienus Proteaeidites Cookson ex Couper emend: 
Machin and Harris 1974 


Type species; Profegetdiles adenuniinidey Cookson 
J950 


Protedcidiies spel. 1 deferiitiinides Comkscnllt eras ysis 
Cookson coniples 
FiGi 1SA-D 


Comments Cooksan (1950) dillereudais tMese (wa 
species on (1) the shape (Fo adendiniieidey has straight ba 
slightly cencive sides wheres H rredasiis hits very Concave 
sides}, (2) the pattern (Peeves Nos Hine reheuhiin 
whercas Fo ooreavsus lis a codrse reticulum) and (4p size, (7 
tideniutihoides ws stnaller 32-48% poe equatorial) dhuneter 
wheres 7 ocrosses js lurger, 38 um). There wre seme other 
ininor differences as well. Stover & Partridee (1973) 
deserthe ai Iecrorype of Po adenirthutdes whieh is oie 
larger (73 Lm) Than the sive fiige quoted by Cookson 


PALYNOLOGY OF THE POONARUNNA NO. 1 WELL 31 


oe. 


tre eg 


Fig, 16. Teruary species continued. A, Proreaciditey cookseniae. B.C. D. Propylipalliy cl Po pseudeamuides. E. F. 
Proveavidites sp. 1G, Lewalanipollty sp. et Lo rectomarginis. A. Proreacidites sp. 3. lL. Proteaeiiites cl, P stippleites, J, 
K. Proteaciditey sp. cl obyciurus, L. M. Proteactdites sp, 2 N, O. Profteaeidites sp, et) Po ineqeviaties. Scule bars = 10 un, 


132 MHoA MARTIN 


(1950), Moreover, Stover & Partridge stile That both layers 
of the ex ie thin towards the pores whereas Choksen stares 
(thatthe nexine does not thin towards the pores, 


The two species as deseribed by Cookson are reeownis- 
uble in his stidy, bur there are Uitermedintes Mat ure citi 
cHLlo phike either species Thinning al the exine lowsrds 
(he pores is rarely seen ib is very slight when presen, Size 
tune forthe complex, 28-45 yun, 


Stratieraphic Runge, Por Bo adenuiifeides, (rom early 
Campanian to Bocene in the Onyay Basin (Dettmann & 
liven (996). Par soutien Australia. lute Paleoeene int 
Oligocene (Stover & Purwilge 1973: Milne 19ss), Fors” 
Chassis, Widdle Campaniin- Maastichtialt ae carliest 


Dank in the Otway Basin (Deftruarin & Jorgen L996) Bar 


southern Australia. early into fate Eocene (Stover & 
Partidve 1973: Milne TONS) Bor the complex, late 
Palcovene (this study), 


Proteavidires anguhius Stover in Stover & Partridge 1973 
RIG, 1SG. Tt 


Comments, These specimens fit the deseription of /t 
uneudeius WIN the exception thar the eine is usually drick- 
erauround the pores, Whereas ia the original description, i 
nity be thinner (Stover & Partridge 1973). The precise 
alnuwALot thickening. however is Variable. Sive rinwe, 2K- 
35 un, 


Stratigniphie Runge, Maastrichtian and Paleocene 
(atover & Partid@e 1973), late Paleocene (this study )_ 


Praeactdiies cooksonide Dettnann & Jurzen 1996 
FIG. 164 


Comments, The smatler size differentiies (his species Fron) 
Foran (Delimann & Jarzen 1990). Size range, 3746 pin, 


Stratiaphic Bunge. Cimparniin-Muasnichrian oe eel) 
est Danian in the Otway Bisin and possibly Late 
Cretuccaus/Puleeeene in the Bight/Duntroon Basin 
(Dethoiino & Jurgen 1996). Late Paleocene (iis stiely) 


Proteacidites fromensis Maris 1972 
HIG, 15k, Fe 


Comments. These speeimens fit te original desenption 
well excepl thal the surface pattern is finely punctosretien 
Joly, grading Lo grarular around the pores, in Contrast la the 
evenly weninulor to seubrate pattern af the original deserip- 
thon (Harris 1972), Size, SO-71 yann, 


Stratigraphic Range Mileoeene (Hurry 19727, late 
Voleqeene to inich Eocene (Stuiter P99) Tite Paleouene 
his study), 


Protwacidites erandis Cookson Mat 
FIG. 15t 


Stetivmiphic Runwe. bate Palepoene Ty tid Eeoene 
Stover Xe Proririve 19732 Purch LO 70d late: Paleiwene 
Hihis stily 1, 


Proteqeiditey ficurvcdis Cookson (YS 
rIG, [SN 


Deseniplon, Giri anisopolar, sles Conc ye, aplees IMI 
vate Pores stadirly depressed Splint dnimeron seme pn 


Hhieh (Wining to? panel Corvin Gnarls Lowirds: pore 


Sesing, vexing upprosimaely equal thickness, Columetlae 
support iegularly-shapedl pemmmae/vercucie, 2-1 pm dia 
eter, becommig smaller near pore und prodicing. granular 
partern hear pore. Sive, 3-50 jan 


Comments. The merphology Why 2 feirveatis buat ties 
specimen is slightly smaller and thier walled than the 
ringe yuoled by Slover & Partridge (1973), 


Strigriphie Runge, Late Paleocene through mid Eocene 
(Stover & Partridge (1973), oid Eocene (this study), 


Praeaviliies sp. ct, Fo lerrvaities Conkaon (50 
FIG. TON, O 


Comments. The morphology is very suntan do that a 
joueveitis except that the exing dogs not thin towards the 
pore and the gemmae/verricae are the sume size over the 
grain, wilh a ring of slightly larger verrucae around (be 
pore. Sive, 40 pin, 


Distribution. Lite Paleoeene (his study), 


Hroteavidires sp. ch Ey obsenrits Cookson [950 
FIG. 16d Kk 


Deseription, Sides straight to eonves, pores prouudiny, 
byine 2 yim thick, sexing and nexine more oF less equal 
Width, columelie indistine Surtiee pattern verrucate, ver 
Micue 1-2 pn jo umametern bul several tay be fused tile 
lurger sheets, especiilly over the poles. Nexine broken ilo 
course granules a done 2-3 wi around pure, Size, 32 pn 


Comments, The sexine differs from 2 vbscurua whiell ts 
Widely hacuhite andl semewhat reticuloid (Cooksan 1950) 


Stniligraphic Range. Por 2 abscurus. carly Bovene to lite 
Mineene (Stover & Partridge 1973), Por Protedcidites sp. 
Ch PF obsetiries, late Paleocene (his study). 


Promeachdines sp cl. Bo srippletts 
Portridwe in Stover & Partridge 1973 
FIC. tol 


Comments, This form fits the description of 2 snpplania 
with the exceplion thar the nesine is sliehily Hicker aroun 
the pore than in the miteradial region te is less like 
Protedcndites rectiy Pocknall & Mildenhall (984, the differ 
eneées being (1) the grain is much Girger, the size aiven belaw 
compared with 32-49 urn for 2 reels, (2) the nexine is equal 
faror slightly thicker than the sexine, whereas itis (ree lines 
thicker on Preeti, (4) the columellae are very clear spaced 
O5-10 pm apart and (he surface uniform cand granule, 
whereas the columetive are Tiintand the surface scubrate on 
PM rects (Pockoall & Mildenhall M984). Size, 60 pin, 


Stratigraphic Range. Por 7 atipplariy, iid) Goce inte 
Oligocene USrover & Partridge 1974) and mid Loeene Hite 
mid Mincene (Miephail 1996) th scauthenstent Austratie, 
Porch Poytipaletis, late Paleocene (this study) gad nite 
Bocene (Slutter 1981) in the Like Byre Risin, 


Protects sp 1 
WICK. 16H. 


Deseripron, Sides straight te slielitly come, esi 4 
py tluek an interned region. Uhiinigg towards pears 
Nexing, sexing uipprowimately equal tiekniass, lair 
incomplete, Columellie praduec strane granidhir sume 
matierd. pieees OF TECH Forn rugulile-reticukite pattern, 
rugulie abont 2 pi owide. supported hy we mwa al eel 
Hiielhie Aira. O6-84 pty 


PALYNOLOGY OF THE POONARL NWA NO. | WELL 153 


Comments. This species differs fom 2 frmiecyis in that 
(he sides are Jess concave und it has the rugulite-reticulate 
surfice paren, 


Distribution, Late Paleocene (his study). 


Prateacidites sp, 2 


MiG, loL.M 


Deseriplion, Sides siaieht to concave, exine 2 pn 
thick, distinct sexing and nexing approximately equal 
thickness. CXTNe NOL LinainyE towards pore. Pores 2-4 pinyin 
Wiuuneler, surface pattern finely reticuhue. Size, 19-27 pon. 


Distribution, Late Paleocene (his stady). 


Prive idiles sp. 3 


FIG, Lol 


Deseriplion. Cai Wetiulae with stramht of slightly 
concave sidys, apices tranedte, pores 4-4 pin wide, Exine | 


2 pi thick. with the HexTne, columellate layer and sexine of 


upproximulely equal widths, The exine may (hin stiehthy 
towards (he pores. Surfiee pattern finely verrucae in the 
Titereolpal abel polar resions and seabrace around the pares, 
Size. 33 pn, 


Distribution, Late Paleoowne (nis sticty ). 
Conus GCoiuinupolliy Mildenhall & Poeknall lose 


Type species: Quiniiupaliiy pyitativpera (Marin 
Mildenhall & Pocknall Loso 


Oniatinigpaltiy priladspara 
iMartind Mildenhall & Pockoll 1oso 
BIG 7, 

Comments, These specimens are slightly kiger those of 
the orainal deseription, (Ss anol this study compured. weal 
WHAROSTMO OF TS pin (Martin 19730 and they hick of the 
Stil seme circular expinsion of the end of the colpi seen 
on (he original specimens. These differences ure cehitively 
minor and Mildenhall & Poeknall (1989) note similar yvari- 
HLio, 


Straligraphie Range, In southeastern Australia, kite 
bocene tothe present in east coast rainforests. curly Eocene 
inthe Gippsland Basin wae mid Bocene ta Pleistocene in 
the inland Murray Basin (Macphail 1996), In the Lake Eyre 
Basin, fd bovene (Sluiter 1Y9L) and late Puleoeene (this 
study). 


Genus Ahopilies Wodehouse 1933 
‘Type species: Rhopites bradley Wodehouse (Ya 


Khopites aiwokuus (Couper) Pocknall & Crosbie [982 
PIG, WE 


Struligmiphie Runge. Mid Eocene to hile Plineene in New 
Zewhind (Poeknall & Crosbie 1952). Oligovene-Mioeene in 
Queensland (Hekel 1972). late Rovene to Plioeene- 
Plemtoceie in southeastern Austtalia (Macphail & Truswell 
O80: Martin 1974a) In the Luke Eyre Basin, lute 
Faleosany-early Bocene (Slutter (991), mit Bocene (his 
Stocly), 


Khopites spock R ulvectains (Coupert 
Pockiall& Croshie 1087 
NG LPO 


Description. Grain oblate. ticolporate with long colpi, 
very small apocolpia. Colpi with well defined margins and 
granular colpal membrane up to 4-5 pm wide at equator 
Endopore with thickened transverse margins ynd capped 
With raised plug bearing granules, Hyine [1-5 yim thick 
with thin nexine and perforate tectum. well defined col- 
umellae and reticulate surface pallern. with mat = 0.5 pny, 
Jummina 0.5-1.0 pm. Size, 30 um polar view, 26-30 pin 4 22 
hin equatorial view, 


Comments. The wide colpal menmbrabe and thickened 
endopore resemble KR. aleeolatis Which, however, has psb 
late volpi, 


Distribution, Late Palueeene (this study), 


Genus Satalinidites Cookson & Pike emend. 
Potonié 1960 


Type species! Samelimudiios cumocaicus Cookson & 
Pike 1954 


Seortalunddites cataozeious Cookson & Pike S54 


BIG. U7F 


Stultgraphic Range. Southeastem Australun early 
Eovene into lite Rocene (Stover & Partridge 1973. |982: 
Partridge 176), Lake Eyre Basin, iid Bovene (Sluiter 
191; Alley ef ah L996, (his study) 


Cenus Superacevidaepullenites Potonié Piriysor & 
Thiergart [O50 

Type species: Supeiaceaidacpatlonites Gal Pallenites) 

imantiestts Polopie 193] 


depativendacpollenites onailus Waris (972 
VIG. 17, K 


Strtigniphic Range: Southeast Atistralia. early Racer 
(hrough Miocene (Stover & Partridge 1973), mid Eocene to 
mid ‘Tertiry (Harris (972), Lake vre Basin, lite Paleocene 
(this stad), and mid) Bowene (Sloiter 199]. Alley et el 
1996), 


Genus Simplicepollis Prurris [965 
Type species: Sraplivepallis meridians (65 


Simplicepaltis meridicants Harris 1965 
HG, 17, 2. 


Comments, This pln tetrad is rare when compared 
with the usual tetrahedral retried, 


Straligraphie Ramee, Southeast Australia, bate 
Cretaccous into dale Bocene (Stover & Partridge 1974), 
Lake Eyre Basin. late Paleacene-early Eoeene (Sluiter 
1991), mid Locene (this study). 


Genus Sirmpsenipellis S. Ry Srivastava [975 


Type species: Sinparcnipods mudlonsis SIG. Srivastava 
lOTS 


Sumpsonipollis sp. 
FIG, |3H, 1 


Comments Phe renulur strate ridges on top of u perto- 
nited tectum place this pollen (pe in Sinppsenipaltis, The 
specimen illustrated is 42 pm polar diumeter x 26 pm equa- 


IAd HLA, MARTIN 


torial ditunetir, much larger than Sinpyenipediiy mullert 
Kemp in Kemp & Tlirris (277 with dimensions of 4-22 
Hn polur diameters POe 2) pn equatorial duneter 


Diswibution, Late Paleocene, 
Genus Sricelpiies Cookson ex Couper (954 


Type species: Tricolintes reticulatis Cookson 1947 (sab- 
sequent desigaation hy Couper 1953) 


Trivalpites sp. cl, T. aspercnarsiniy Melntyre L908 
FIG ITH I 


Deseripdion, Grain obhite, triculpate with wide. gaping 
open colpi. LXue 15-2 wim in intercolpal region. thinning 
nhirkedly towards colpi. Nexine thin, collumethite bayer 
thin with distinet columellie, tectum = 1 ain in intereo)pal 
fevwon. surface scabrate. Size, 25-29 pm. 


Comments. The exine thinning towards |he colpi is not 
seen on 7. usperunerginis. The very thick tecuimn and the 
ane thinning lawards the colpi wre not seen on Trice/pires 
neroblins, 


Stratigraphic Runge, Vor 7 daperananminy. Paleorene im 
New Zealand (Metntyre L968), lower ‘Tertiary to late 
Miovene in Queenshimd (Hekel 1972). Tricalpires sp. et 0 
dsperemurginis Wil Rovene (this stacy) 


Tricolpiles spel, To cenfessis 
Stover in Stover and Partridwe [974 
FIG. 170 


Commerits. These specimens are very simihir to those 
ipinully desorbed except that (hey are slightly laren, the 
size given below compared with a maximunpal 25 pm, The 
exme is 1 15 ui, Owe layers are distinguishable und: the 
surfice is psiliite/scabrue whereas the original description 
stiles that the Livers in the exine are not clearly different 
ated Sive, 3-31 um, 


Strtigruphic Range Por 2 caresses, Santoniarn to lates| 
Mititstrichtitn-cartiest Paleovene (Helby endl. 1987), ef 7 
confess, late Paleooene (his study), 


fricuipiies spel 7. divens Harris in Kemp & Harris 1977 
EG, 17M, 


Comments, Lhis specimen is larger thin those described 
by Haris (in Kemp & Harris 1977), 30 aim equatorial diam 
cher compured with 17-23 pn respectively, Phe morpholog- 
ial Hoatures, however, are very similar. 


Strmgtuphic Runge, Por T discs, Paleocene, Ninetyeast 
Ridve Tidiin OQecan Ukemp & Harris 1977) and hate 
Eocene, Luela Basin (Milne 1988). For Tricvelpites spel E. 
dives, Wid Toeene (this study). 


Hrivalpites phillipsi: Stover in Stover & Partridge 1974 
FIG, (7T 


Slutereplic Range. Southeast Australia, Paleocene into 
lute Eocene (Stoyer & Purtridge 1973, 1982; Partridge 
1976). Lake Eyre Basin, early Eocene (S)uiter 1991) and 
mid Bucene (this stuely). 


Ticolpites Hiamnasii Cookson & Pike 1954 
FIG, 17ULV 


Stratigraphic Range. Mid ahd the lower part ot the late 
Eocene (Stover & Partniuee 1973, 1982), Lake Byre Basi, 
carly and mid Bocene (Sluiter 991), witd and dite Eocene 
(Alley eral, 1996), mid Bocene (this study), 


Cithis Triealporites Cookson ex Stoyer& Eyagns JOT 


Twpe species! Triedlporites spheericn Cookson (desiy- 
mated by Stover & Evans 1974) 


Trivalporites anguriin 
Partridge in Stover & Partridge [973 
IG, 172 


Comments, These specimens fit the diagnosis except thal 
the genenaly indistinct ora of the diagnosis wie haedly ves 
ble here. Size, 31-47 puns 22-29 yon, 


Strangraphie Runge, Southeust Austyilian carly toiu-late 
Eocene (Stover & Pareridge 1973, 1982). Lake Eyre Bursin. 
mid Bovene (Sluiter $981, this study. 


Tricalporiies lewros Partridge in Stover & Partridge }973 
FIG. L7WOX 


Comments, These specimens fit the diagnosis well. 
except thal on some specimens, the “indistiner or? eannar 
be seen at all. On other specimens, the exine i the interns 
dial areas may be 1-1-5 pm thick, compared with 2-4 jn in 
the diagnosis, and on these, the ora miy protrude so Mie 
they luck the polygonal autline of the thicker walled speci 
mens. Size, 20-25 pun. 


Straugmiphic Range, Southeast Australi. mid bocene to 
mid Miocene (Stover & Partridge [974, Maephail 1996), 
Lake Eyre Basin. late Paleocene (his study) nel reid 
Fovene (Sluiter F991, Alley ef al, 1996), 


Genus Pricelporpotienites 
Prluy in Thomson & Pug 1983 


Type species: Triealparepoltiadtes cefinn (Potonie) 
Plugs }953 


Tricalporopadledites etdobaliens Melotyre 196s 
FIG. LIRLS 


Stratigraphic Range, Southeast Austtalig, mid Hocene 
inta Plingene (Marti M87, Macphail 1996), Lake Eyre 
Basin, Early-mid Eocene (Sluiter 9b), oid bocene chs 
stacy ). 


Genus Trieciiey Cookson es Couper 1954 


Type speeles: Wiortios Hagnificuy Cookson LYS), desis 
nated by Couper 153 


Triavites minixentiy Melntyre L965 


FIG, iso 


Description, Grain iiporite, pores 23) unt duneter, 
axing bum thick, (wo layered. psiate except for fit pat- 
tern around pores. Size. 13 pm, 


Comments. The morphology of the specimen fits the 
deseription of Triprites miniseuluy given by Melntyre 
(1965). 


Stralivraphic Range, New Zealand, Pauleguene (McIntyre 
19651, Lake Eyre Basin. lute Paleocene-carly Eocene 
(Shutter 1991). late Puleocene (this study). 


PALY NOLOGY OF THE POONARUNNA NO. 1 WELLL. 135 


." 
. 
om, 
fe 
a 
— 
\ 
' 
™: 
onl 
a” 


Fig. 17. Tertiary species continued. A. B. Polyporina granilata. C, D. Simplicepolliy meridianus, E, Rhapites alveolatus. b. 

Santulumidites catnoceieus. G. Agleoridia quatamts. Us Lo Trieolpites sp. cl To asperumerginatus. J, K- 
Sapotuceoidacpollenites voiunduy. L. Quintiniapallis psilatispora. M,N. Tricolpites sp. cl. T. discus, O. Tricolpites. sp. 
eh T confessus. P,Q. Rhopites sp. ch R. alveolatus, RLS. Tricelporopallenizes endobatieus, VT. Tricalpites phillipsii. U. 
V. Tricalpites thamusit. WX, Tricolporites leuros, Y. Triperopollenites canbiguus. 7. Tricalporites angurium. AA, BB. 
Trivolparites sp. 2. CC, Tricalporites sp. 1, Seale bars = 10 ym. 


130 HA, MARTIN 


Triarites sp. 
FIG. LHI 


Description, Grain circular, triporate, pores 5 pm wide 
with ragged margins. Exine | pin thick, clearly two layered, 
with faint scabrate surface pattern. Size, 25-25 pm, 


Comments. The morphology of this species is smilar to 
that of some species of Ulmaceae, especially Apuntfes 
philipensiy. The former, however. his a ragged edge to the 
pore Whereas the litter has a well defined rim. Some species 
at Celtiv are summlar to the fossil also, e.g, C. glabra and C. 
eccidentaliy, but the latter show distinet columetlae and the 
pore has an annulus. 


Distribution. Late Paleowene, 


Genus Iriporopollenites Pflug & Thomson 1953 in 
Thomson & Pflug 1953 


lype species: Triporopollenites coryloides Pile in 
Nhomson & Pflug 1953 


Triparapotlenites anibiguas 
Stover in Stover & Partridge 1973 
PIG. 17¥ 


Strahyraphic Range, Southeast Australia. carly Locene 
ite kite Eocene (Stover & Partridge 1973), Murray Basin. 
mid Bocene inte early Miocene (Macphail 1996), mid 
Focene (his study), 


Genus Tuhiulifleridites Cookson ex Potonié 1900 


Type species: Tibuliflortdites antipadica Cookson, des 
ignated by Potomed [960 


Tubulifloridites sp. 
FIG. TKR 


Suahyraphic Range. Triporopollenizes bellus Zone. law 
Miocene (Stover & Partridge 1973) to the present as the 
daisy family, Asteraceae, Oligocene through Pleistocene 
(Macphail) 1996), Pound only in the Plio-Pleistocene of this 
sludly, 


Unidentified tiixa 
Dicolpopollis Pflangl emend, Potomie 1966 


‘Type species: Dicolpapaltis kockellt Plhinzl 1956. desig- 
mitted hy Potonié 1966 


Diculpopoilis sp. 
FIG, |&F 


Description. Grain has two, gaping colpi, Exine, 1 pm 
thick, has nexme, columellate layer and tectum, all of 
approximately equal thickness. Surface pattern line, uni- 
form puncto-reticulum. Size, 40 x 32 pm. 


Distribution. Middle Eocene, 


Tricalpites sp. 


FIG. 180, P 


Deseription, Grain prolate with broud poles, long colpi 
with ragged margins, Exine | pim thick. two layered with 
very fine, faint columeslae. Surface pattern finely puncto- 
reticulate. Size. 20 16 cin 


Distribution. Late Paleocene. 


Tricalporites sp, | 
FIG. 17CC 


Description, Grain oblate. tacolporate, distinct thicken- 
ings uround the pores. Exine | um thick, with two layers 
approximately equal thickness. no discernible siructure, 
Grain covered with minute voni spaved about | jun apart, 
Sive, 15-17 pm, equatorial diameter, 


Distribution. Lute Paleocene 


Tricalparites sp. 2 


FIG, I7AA. BB 


Description, Grains prolate. tricolporate with long colpi 
reaching ulmost to poles, pores indistinct. Exine 1-5-2 pin 
thick, with nexine, very finely columeltlate layer and tectum 
as thick as nexine, Surface extremely fine granulac/eeticu 
late pattern with larger (up to (5 um) fovilae through tee- 
tum. Size, 26-30) um x 22 pn, 


Distribution. Late Paleocene 


Tricalparites sp. 3 
PIG, Isl 


Description. Grains, protate, iivolporiute with mndistines 
pores und granules aligned alony borders al volpi, Exine. | 
2pm thick. is thicker over poles, vie 1 fim in equatorial 
region, 2 waver poles. Nexine, very fine columedlite layer 
und iwetunn of equal thickness, Surface seabrate. Size, 24 x 
{Om 


Distribution. Late Paleocene 


Tricelparites sp. 4 
FIG, [SEL 


Description, Grams oblate. trigblporute with weakly 
defined colpi, pores with ragged matuins, Exine 0 Se) um, 
ho discermble layering on thinner walled specimens, bul 
thicker ones show (wo layers. thinning towards colpl, 
Surface faintly scubrate, Sie, 12-15 pon. 


Distribution, Lite Paleocene, 


Triealparies ap. 5 
BIG. IRMLN 


Description, Cirain more or less spherical, triculporate 
with smooth colpal membranes, weakly delned pares, 
Exine | pun thick with two equal layers. Sexine reticulate. 
lumina aboub fo ym. muri (5 pi in intercelpal areas. 
becoming very finely punera-reticulate over poles and 
towards colp. Size, 26 pn. 


Distribution. Late Paleovene. 


Pancolpate sp. 
FIG. [SE 


Description, Grain presumed originally spherical, now 
llattened. There are about 16 colpi over surface, arranged 1 
form square or polygonal shapes. Exine | pm thick, with 
thin nexine, distinct columellale layer, thin tectum, Surface 
has scullered coni, O.5-1 um high, spaced about 1-2 pm 
apart, Size. 58 urn. 


PALYNOLOGY OF THE POONARUNNA NO. | WELL 137 


Pig. |S. ‘Tertiary species continued. A, Pediasirum sp. BLD. Barrvocoecus braunti. B. Small form. D. Large form, C, 
Morkaldovst pyrdniidaits, B. Pancolpare sp. F. Dicolpopoliis sp. G. Panporate sp. Ul. Trierites sp. L. Triealperttes sp. 3. 
JL. Trieolporues sp. t. MON. Pricelporites sp. 3. O, P. Tricalpites sp. Q. Trierires miniserilus, 8. Tubaliflavidites sp. Seale 
bars = [0 yn, 


138 H. A. MARTIN 


Comparisons, This type differs from Lyningronia which 
has a thicker sexine and a rugulate/verrucate pattern 
(Pocknall & Mildenhall 1984). Portulacaceae gen. et sp. 
indet. (Martin 1973b) lacks the coni of this pollen type, 


Distribution, Pliocene/Pleistocene. 


Panporate sp. 
FIG. 18G 


Description. Grain is broadly elliptical, panporate, with 
about 16 pores, 6-8 um diameter. Exine 3 um thick, with 
thin nexine, thick densely columellate layer and thin tectum 
which has small perforations, < 0.5 tim, spaced 1-3 jim 
apart. Surface pattern granular. Size, 78 kum. 


Distribution, Pliocene/Pleistocene, 
Microplankton 
Genus Borryococcus Kiitzig 
Type species: Borrvecoccus braunit Kiitzig 


Batrvoceceus braunti Kiitzig 
FIG. 18B, Db 


Comments. Botryecoccus braunii ix a cosmopolitan and 
extremely variable species with a number of races. Cookson 
(1953) remarks that only one race, the small form (Pig. 
ISB) has been found in Australia. In this form, the algal 
cells are 4-6 im in diameter and the colonies are tightly 
packed, There is also a much larger form (rig, 18D), where 
the algal cell is cup-shaped, 10-12 ym deep and 8-9 wn 
wide at the top and colonies are more or less fan-shaped. 


branching dichotomously, with the branches 6-9 pm wide. 
This larger form probably constitutes a separate race 
(Blackburn 1936). Botryococeus braunit usually inhabits 
freshwater ponds and lakes. Sometimes it may be found in 
brackish waters and coastal lagoons. It may be extremely 
prolific and was responsible for boghead coal (Cookson 
1953), In the Poonarunna well, B. brawuniii is extremely 
abundant from 67-91 m. 


Stratigraphic Range, Ordovician to the present, 
Genus Morkallacysia Harris 1973 
Type species: Morkallacysta pyramidelis Harris 1973 


Morkallacysta pyrainidalis Warris 1973 
FIG. L&C 


Comments, This species is rare and the specimens are 
usually crumpled, 


Stratigraphie Range. Paleocene (Harris 1973; this study). 
Genus Pediastrum Meyen 1829 
Type species: Pediastrim duplex Meyen 1829 


Pediasirum sp. 
PIG. TSA 


Comments, Pediastrum is usually found floating amongst 
aquatic plants, rarely in deep water. Tt may be found in lakes 
and small ponds where the water ts rich m= nulients 
(Pentecost 1984), 


Stratigraphic Range, Early Cretaceous to the present 
(Evil 1963). 


TRANSACTIONS OF THE 


ROYAL SOCIETY 
OF SOUTH AUSTRALIA 


INCORPORATED 


VOL. 122, PART 4 


RHOPALOMYIA LAWRENCIAE, A NEW GALL MIDGE SPECIES 
(DIPTERA: CECIDOMYITDAE) DEFORMING LEAVES OF 
LAWRENCIA SQUAMATA (MALVACEAE) 

IN SOUTH AUSTRALIA 


By PETER KOLESIK* 


Summary 


Kolesik, P. (1998) Rhopalomyia lawrenciae, a new gall midge species (Diptera: 
Cecidomyiidae) deforming leaves of Lawrencia squamata (Malvaceae) in South 
Australia. Trans, R. Soc. S. Aust. 122(4), 139-145, 30 November, 1998. 

A new gall midge, Rhopalomyia lawrenciae, is described from swollen leaves of 
Lawrencia squamata collected on Hindmarsh Island in the River Murray estuary, 
South Australia. Inside each of the infested leaves is a chamber occupied by one larva 
of the new species. Males, females, pupae and larvae of the gall midge are described. 
All specimens of the host plant lodged in the State Herbarium of South Australia were 
examined for galls and this revealed a wide geographic distribution throughout the 
state. A key to adults of the three known Rhopalomyia species occurring in Australia, 
R. lawrenciae, R. goodeniae, a native species damaging stems of Goodenia lunata and 
R. californica, an introduced American species damaging flower buds of Baccharis 
halimifolia, is provided. 

Key Words: Gall midge, Cecidomyiidae, Rhopalomyia lawrenciae, Lawrencia 
squamata, saltmarsh flat, River Murray, South Australia. 


Transaetioms of the Roved Sociery af S Ausr (1998), 12204), 139-145, 


RHOPALOMYIA LAWRENCIAE, A NE 
CECIDOMY UDAE) DEFORMING LE 


GALL MIDGE SPECIES (DIPTERA: 
AVES OF LAWRENCIA SOUAMATA 


(MALVACEAE) IN SOUTH AUSTRALIA 


by Peter Kolesik* 


Summary 


Robesik. POCT998) Rhopalonvid lawrencia’, & new wall midge species (Diptera! Ceeidamyiidac) deforming 


leaves of Lewrneia syne (Malvaceae) in South Australia 


November, 100% 


A new gall midge, Rhopalomvin lawrence, is desc 


Tray. R, Soe, S. Aust, 122(4), (39-145, 30 


ribed tram swollen leaves of Lenerevweia sauce 


collected of Hindmarsh Istana in the River Murray estuary, South Australia. Inside cach of the infested leaves 


isa chamber oceupiew! by one larvieol the new species. S 


Jales. females, pupae and larvae of the gall inidge are 


described. All specimens of the host plant lodged in the State Herbariurn of South Australia were examined for 


valls and this revealed a wide geoyraphie distribution (he 


Rhopuloniyie species occurring in Australia, R. fenwnenede 


oOughout (he state, A key to adults of the three known 
te, KR, goodeniae, a ative species damaging stems of 


CGoadenia hima and Ro californica, an introduced American species damaging Mower buds of Baveherts 


Julintifotiog is provided 


Kiy Worbs: Gall midge. Cecidonmytidae. Rhopulony 
River Murray, South Australia 


Introduction 


Lawrencie is an Australian plant genus comprising 
I2 species of perennial berbs and small shrubs 
Jessop LO86). Lawrence squamate Nees in Lehm. 
is ad rigid shrub up to | m high, eecurcing in all 
miuintand stiles Jessop 1986), In South Australia, it 
grows on saltmarsh flats, sand dunes and rocky cliffs 
along the coust and on sandy soils and marshes 
tikind. The plant forms part of the shore vegetation 
on the saltmarsh flats cn the estuary of the River 
Murray where in September, 1996, on the south- 
eastern coast of Hindinarsh Tstind. many leaves of 1. 
squamata were found to be swollen (Fig. 1). ‘The 
swellings were caused by kirvac of ab unknown gall 
nidee desenbed here. The new species is placed in 
Rhopalomyia, a large, worldwide genus. The new 
species becomes the second gall midge deseribed 
from South Australian saltmarsh Cats, the lirst. 
\splhomadvita inflate Kolesik (1997) having been 
described Just year. 


Materials and Methods 


Branches of Lewvenei squemare plants bearing 
leaf galls were collected on Hindmarsh island. South 
Australia on & September. 1996, The branches were 
brouehh tothe hiboratory and the galls processed. in 


‘Deparment ool berigutnore, Vircutrin aitl Qernatogy, Wate 
(Srpies Tye Tiniversity ot Adeliide PMB | Glen Osinond 
S Aust Wert 


fa fawreneiae. Lawrencia suena. saltrmarnsh Mat, 


Galls of Afiopaloniyia lawrenciae sp. nov. on 


Vie. | 
Fawreiicia squuieta, White arrows mark whole galls, 
black arrow marks a gall cut open, presumably by birds 


Seale bar = Winn. 


one of two ways. A small number was dissected and 
the larvae and pupae were preserved i) 70% ethanol, 
A larger number was lellon the branches and kept in 
plastic bags Lo develop to adults. Pupation toak plaice 
within the galls. Emerged adults were preserved i 
Wie ethanol after their colour had) been 
Canad balsam mounts of type specimens were 
prepared according tO the technique outlined by 
Kolesik (1995). The type series, and other (terial 
rehuned in 70% ethanol, are deposited in the South 
Australiana Museum. Adelaide (SAMA, and the 
Australian National Inseet Collection, Canberra 


noted, 


HG PO ROLESUs 


JANIOC). A dried sample of an aifested plant is 
deposited in the State Herbarium of South Australia, 
Adelaide [SUSA[. All measurements refer to the 
holotype and paralypes, Investigation of the 
veographie distribution of the new species was bused 
on examining the presence of galls on dried 
speelinelis of the host plant deposited in SHSA, ‘The 
galls were easily recognisable and some still 
contined pupal skins of the new gall midge, 


Cienus Khopufouyia Ribsaamen, [S92 


Rhopoulomvid Ritbsaumen, |h92: 370 

Woe species: Oligetrophis tanaceticola Karseh, 
1879: VIL Ther. west, Prov Ver, Wiss, Kunst) 27 (des, 
Kicftor, R96: 89) 


Rhapalomsia is a large. worklwide venus of the 
Wibe Oligotrophini with an undivided eibth feniale 
ibdominal tergite ald completely setu}ose 
vonostylus. Most ofthe known species hive a one- ls 
three-seymented pulpus. aim one speeres. (he 
Aastralian A. wandeniie Kolesik (1996), has a three- 
or lourseamentedl palpus, 


Rhopalomyla lawrenclae sp. Ww, 
(FIGS 1-19) 


Holoiypes oh. Utidmarst tstand, South Australia 
J35° 34S. 138° 53 BY, Yin, 1996. PL Kolesik, reared 
fromr a leah all on Lemvreneie sgaeneta Nees in 
Lehn. gall collected §.i8. 1996, [SAMA, 121394], 


AWS pupae (SAMA, [21395- 
40), 244, . 2 pupae [ANIC]. same data 
bul gurersed 8-1} 7a, 1996. 4° larvae, (SAMA, 
PAO PAIAOS|, 3 larvae [ANIC], collected wath 
holaiype, 


Pararyyrens 2" 


Other murvad> 279 Be ARR pupae, TD) bervie 
JSAM seine dit as paratypes: gall. coteeted with 
holaiype, ADIGA SLO? [STISA],. 


Mei (Pigs 27) 

Colowe Head and thomas brown, abdomen with 
splerotiseadl paris. browu anh fon-selerotised parts 
wry 

Head: Antennas seape broadest distally, as long as 
dicta! brewdth. [Sx length pedieel pecieed broader 
Hie lone Mopellomeres ba 14 re rumiben fest and 
sevehd fol Tased, neck about ey lenwih node: 
CHCUHHH COMprisMme (WO transverse and two 
Jonna! Bands. PaAdpus three-seaypented, Bye 
facets pounded, close together, spapser al verles, eye 
bridve 5-6 fitels lone. Labelle rowehly tere 
spherical. literally with 2-5 selae. Prony with [2-20 
selie per side, 

Thorns: Wing length 2.2 mim (hO-24 n= 50. 
width) LO im (O,8-1be Rs same thickness: cntine 


length. slightly curved posteriorly. joining C anterior 
to apex; Ry joining C near wing mid-length: Se cell 
pigmented and together with Ry and adjacent part of 
R; bearing scales. Claws toothed, empodinn as long 
as claws. pulvilli hall length empodiun. 

Abdomen: All tergites with pair of sensory sete in 
anterior corners and row of setae posteriorly. lergites 
Tand 8 additionally with few selue scattered meso- 
laterally; stermites 2-8 with pair of sensory setae 
anteriorly, a row of sctue posteriorly and a band of 
sctue mesally. Genitalia: gonocoxites cylindrical, 
ventral articulation with wonostylus longer than 
dorsal articulation, setose and setulose: gonostylus 
uboul same width entire length, sctose and setulose 
throughout, with strong tooth, comblike distally; 
cerer separate, selose and setulose: hypaproet 
bilobed. with one seta apically on cach lobe, 
setulose: purdametes setulose, with G-8 setose apical 
papillae: wedengus come, 


Fumule (Vis 813) 

Colours lead and thorax brown, abdomen with 
sclerotised parts brown and non-seleroused parts ced, 

Head: Flagellomeres 12-13 i mamber. terminal 
ones sometimes fused. neck about Yj length node. 

Thorax: Wing length 2.0 mm (ha, = 5), 
wiih OS min (0,6-0.9). Tergite § Wath single pare of 
scosory setae anteriorly, sclerotisation undivided, am 
shape of Jeter “sa. Ovipositor: cert fused into 
single, termingh spheroid lamelli, setose and 
setulose; Aypoproct rounded apically i dorsn 
ventral view, bearing (wo selue posteriorly, setulose. 
Other characters as in male. 


Pupa (Figs (4-16) 

Colours antennal and frontal hoths pigmented, 
brown, remaininie parts anapimented, Leagil) 26 
mm (2.5 2.8.0 = 5). Antennal hops strana, biti, 
JO pm 6172-206) long, Proms on eae sides are 
frontal ford, pare oot papillae on lower fice. one 
selose, one asetose: Fiplet ot lateral taeil papillae 
gue setose. wa uselose, Prothoricte spitiels with 
several irregular protuberanges apreally. (ches 
ending between hall and distal third of spracte. 
Intesurment of ubdeming segatents euvered with 
spice. very dense dorsally. ii daesul spities 
Present, 


Lest inyter teary Cees be, 08) 

Colours piokish ped. hength 2.5 iy t2a-2.6, 1 = 
0). Integuiment covered with spieuhie, ead with 
postero-lareral aporlemes shorter (tan heal length, 
No sparahe present, ATL papillae with shark setae, 
Thoricie and first ablomioil segments will pain al 
ventral pupillae. (wo pairs of pleneal papilhte. tree 
pairs of dorsal pupillie, Abdomind segment & with 
pain of yeotal papiliie. Ave pairs of pleural papillae, 


ANEW GALL MIDGE FROM LAWRENCIA SQUAMATA I41 


Figs 2-7. Male of Khopalomvia lawrenciae sp. nov. 2, Head in frontal view. 3. Last three flagellomeres. 4. Sixth 
Magellomere, 5. Last larsomere with claw, empodium, and pulvillus. 6. Genitalia in dorsal view. 7, Wing. Scale bars = 
100 tim 2, 3; 50 pm 4-6; 200 um 7. 


142 P. KOLESIK 


Rises fh. 


; / 


oe 


10 


‘o"o 


@ Wr ity te 
Hein rtater My 
Mee Pelee 


13 


Figs 8-13. Female af Rhopalomyia lawrenciae sp. noy, 8. Posterior end of abdoinen in dorsal view, 9, Posterior end of 
ES | ve I 
ovipositor in ventral view, 10, Last three flagellomeres (paratype 121397), 11. Sixth flagellomere. 12. Posterior end ot 
abdomen in ventral view, 12. Last three flagellomeres (paratype 121398). Scale bars = LOO pm &. 10, 12. 13) 50 pum 9: 25 
fim TL. 


fay > ae a ee, 


A NEW GALL MIDGE FROM LAWRENCIA SQUAMATA H4i 


Figs 14-18. Rhopalomyia lawrenciae sp, nov.. 14-16, pupa, 17,18, larva. 14, Anterior part in ventral view. 15, Anterior part 
in lateral view, 16, Prothoravie spiracle. 17. Two terminal segments in dorsal view. 18. Head capsule. Scale bars = 100 


pt 14, 15.17: SO um 16, 18. 


pair of dorsal papillae. Terminal segment with four 
pairs of terminal papillae. Anus ventral, 


Etvinalowy 
The specific name meuns “of Lawrencia’, the host 
plant, 


Gall and biology 

Leaves of Lawrencia squamata infested by this gall 
midge are several times larger than normal in volume, 
4-6 mm long and 3-4 mm wide (Fig. 1). Each gull 
contains a chamber occupied by one larva. The 
chamber wall is lined with a thin, hard, pale-brown 
layer of tissue at the time the larva is fully-grown, 

Pupation takes place inside the gall. The pupa raises 


two thirds of its body outside the gall before the adult 
breaks through the antertor end of the pupa. On 8 
Seplember, 1996. on the southeastern coast of 
Hindmarsh Island (Fig. 19), the galls contained larvae 
and pupae, with the first adults already emerging. On 
this occasion, the host plants were about 20 cm high 
and about 50 cm in diameter and approaching the end 
of flowering. Lawrencia squamata accounted tor 
some 10% of the ground covering of the dense, herbal, 
coastal vegetation at this locality. The population 
density of the new gall midge was high, comprising up 
to 10 galls per host plant. Many galls were found cut 
open, possibly by birds, a phenomenon described for 
other cecidomyiid galls previously (Struble & Osgood 
1976; Tscharntke 1990), 


Id P KOLESTE 


Pie 18. Tindtidest Iskind Soul Australia — aerih voew at (he pypie loealiny nf Réepeedanein Javed spe iia | 
(iimdinarsly feline 2, Sty Riehid Poriisuta. 3 Younstiistend Pernaali White aerawe tates (he type hieality 


Cre toile disnledion 

Calls or We pew wall unide Were teumel on 
huicreneta aynrneta phils collected from the 
tollawiay lnoiities on South Austratias SW ol Marte 
jr SYS lav SU BY wack to Fisher [abe 204 
Pa Sb 6S ke Wool’ Nullarbor [3b 27S. bane 
hb a0 hm Sor byre Highway [sl 47'S, tale 82 
Pit km Soot Voomibre [AVP Ses, Pate ES by, Veet 
Sinelaie [32 7S. ate ST ey. Thevenand (S27 YS, 
[AV WU del sineky Busy Jar? 1 So 14s" Pe], 
Vale Pies Dice |evEe bat 1av° 4 El, Nit luce 
Stalin (22° 2358, (35° Heh Camere [42° 2655, 
PAM? 32 Fay, Fue Psheret |92" 28S, 138 TY by]. Pee 
Istund AD IY’ SS. 1S" AE] bs kin No iliscn 
(vw Ww 19S ay be. St brivets stand ae 3148 
PAR TR EL). Mastin bela | $2" 40S. 144s 7 Eb] 
Teal Ian pat? sao Sas 17 RL Redein 
Suevey are pat 4S Lee al UY Strike Bay 
ja de Ss. (oe SE] 6 kin Et Cyneens [9° 43/ 
Hh. Ir UE  Vellistens [33 GMS, Pade SA) bf, 
Cywell (S5° 45S, 136° 55 be), Buea Creek Plain elas 
kit NS of Mt Mary bh] 34° S95 A. 18? 26], AW side eal 
Lake Lhamntin [93° 97S. 138° 8 leh. Port Hoel 
JS42 05S. 1578 38 De), Mit Mlory Pade i Sade on 
i), Acetate (liter Llarbour |34° 48 KR ) AR 20) FY] 
Weel Cupe (iynes National Burk) [38° 1S. 1 3e° 50 
1) Minkilon (rt! das, 147° Ae), Pom Linch 
jae 44S. LAS S211, Porrens Isham La4° 44° 5, 
ISK" 32 EY Part Adelaide (24° S05 8, Fee SE], 
Bort Noamlinga [Sa 098° S. Lage 28" A]. Pondaliwie 
Vay [AS ES. 146 SOUT. Pinditmeste Istana [3 
Th S, Lane S07 |) Malinois [3° 96S. 139° 40 by. 


IX kin N of Meningiv [35° 37° S. 1aHe 2h ty 
Vennavhar Porat (Rangsran isto |a5° 33S. ban 
4? E|, Coovong [40° Js S, LAV 43 TE] 

The highest qibuevlamee al calls wep Camigh ou 
plats wolleetwal Ta PY fy NooM Wace urn tbe 
exposed roeky Head hind of Dog blued whine plans 
Were subject to salt spray duping stariis Che phans 
From (his area whicll ve depasited if SHSA aye pall, 
dense shrubs. whoo! (OOo figh gid E50 nein 
diameres, eaelt bering seme LOU cally at tlie new 
sll inntee Oher loealions wilh fel uhundunee pl 
ahs were The Coorany ephints collected tos 1 8A! 
by ME. aR, Sti) yd Added iter Harb 
(plants collected Tah PT by A. Gh Spronenh 


Renrirks 


BNO TS WE GT Getty SSE SEE EES 
Species farming walls an plants af ihe Lanny 
Asteraceuc. J lene ane Pear alist ete Hypa icons ul 
sriips fin ie eens, one Conlinis species (list Maye 
hima with spatula tbseat ind pape will) alent 
hones presenp pie orher ennniins spee wes thar five 
eve WOT Spatula Brose Tih crid) puijoue: Watlh sate nital 
Horiis shscnt (Gane FOOD) Dre nw species belts 
te tips faeiier group The enly ather kien 
Australian atte Afapaferayig, gente rites 
Rolesik (1990), a species defurtinie stone at 
Cromena Tame J. Black (Quodenimeeney in the 
lathe Fayre region. beloms te tle latter sedup Boil 
spevies belongs 1 Syivews (1975) biulegical eroup V 
of promaey eall prdueers will lurvue eed 


A NEW GALL MIDGE FROM LAWRENCTA SOQUAMATA 145 


solitarily und pupation taking place in the gall) The 


gall of Ro goodenide comprises & conglomerate of 


individual chambers whereas the gall of R. 
luwrenciae sp. nov. consists of a single chamber. 
Because only these two species of Rhopalomyia are 
known to be native to Australia, it is loo early for a 
general characterisation of the genus on this 
continent. Below. a key is given to adults of the two 
nalive species and R. californica Felt, an American 
species introduced into Australia to control 
Bacchari. 8 helimifolia L, (Asteraceae) (McFadyen er 
al.': Gagné & Boldt 1995), 


Key to adults of Australian species 
of Rhopalomyta 


1. Tarsal claws toothed... R. lawrenciae 
Tarsal cliuws simple 
2. Palpus with 3 or 4 segments: length of papillae on 


male parameres !/s - '/> paramere Width... 
wR, goodeniae 
Palpus with | or 2 seements; length of papillae on 
male parameres about '/2 paramere width.......... 
-R californica 


Acknowledgments 


W. R. Barker and M. C. O'Leary. both of the State 
Herbarium of South Australia Adelaide, courteously 
identified the host plant species and assisted in 
examination of dried host plant specimens, 
respectively. D. Eastburn, Murray-Darling Basin 
Commission, kindly gave permission to print the 
photagraph in Figure 19 1 thank J. D. Gray, 
Department of Horticulture, Viticulture and 
Oenvlogy University of Adelaide. and R. J. Gagne, 
Systematic Entomology Laboratory USDA 
Washington DC, for commenting on an early draft of 
the manuscript. 


References 


CivGnr. Kd. (1904) * The Gall Midges of the Neotropical 

Region” (Cornell University Press. Ithaca, New York). 
— & Bonn, PE. (1995) The gall midges (Diptera: 

~ Cecidomyiidae) of Baccharis spp. (Asteraceae) in the 
United States, Pree, Aur, Sac. West, 97, 767-778. 

Jessor J.P. (1986) Family Malvaceae pp. 821-848 In 
Jessop. J Po & Toelken, Ho R. (Eds) “Flora of South 
Austratia,” Part 2 (South Australian Government 
Printing Diyision, Adelaide), 

KOBPAER IJ 1896) Newe Mitthedungen iiber Gallmiicken, 
Wiew. bat. Zi@ VS. 85-105, 

Komrsin. P(1995) A new species of Evctiectioarnia Pelt 

(Dipteras Cecidomyiidae) on Eyecafypiis faseiculasa in 

South Australia, /. Aus ent See. 34, 147-152, 

(1996) Rhopulomvie geudeniin, a new species of 
Ceeidomylidae (Diptera) damaging Ceonenin linata 
(Goodlentaeeae) in inland Australia. Trans. Ro Soe. 8. 
Vass. 120, (SS-160, 


PMePAb VES. Poh. Danses, Ch Bod “Tompnas AL J (P9RSy 
Bielujieal control ab grommet bush: pp. 2a 40 fn Harvey, GI 
(edo) “Australian Weeds KReseurch Newslenor' (The Ala Ploncher 
Research Sein. 


TSCHARNTEKE, TT. 


(1997) Two new species of Asphondylia (Diptera: 
¢ ‘ecidomyiidae) from Halosarcia spp. (Chenopodiaceae } 
in South Australia. (id. 121. 59-66, 

RUBSAAMEN, BEL 41892) Die Gallmiicken des Kanighchen 
Museums fiir Naturkunde zu. Berlin, Berl, Ent 237. 
319-411. pls VIEXVUEL, 

Stkiiser, BD. B. & Oscoon, F. A. (1976) Predation on 
larvae ot the balsam wall undee, Desinewra bulsanicala 
(Diptera: Cecidomyiidae), within galls in Maine. Ceneel, 
Ent. 18, (443-1444. 

SYEVEN, bk. (1975) Study on relabonships between hibits 
and external structures in Oligotrophidi larvae ( Dipterin: 
Cecidomyiidae). Zvel. Scripta 4, 55-92. 

(1990) Vogellruss heemtrachtigt dic 

Dichteregulanion einer Gallmiickenpopulation dureh 

Parasitoide: Weeliselwirkung ewischen vier trophischen 

hbenen. Min, Deutsch Ges. Alle. Anwew. Biri. 7 552- 

544. 


DASINEURA WAHLENBERGIAE, A NEW SPECIES OF GALL 
MIDGE (DIPTERA: CECIDOMYIIDAE) DAMAGING SHOOT 
TIPS OF WAHLENBERGIA STRICTA (CAMPANULACEAE) 

IN SOUTH AUSTRALIA 


By PETER KOLESIK* 


Summary 


Kolesik, P. (1998) Dasineura wahlenbergiae, a new species of gall midge (Diptera: 
Cecidomylidae) damaging shoot tips of Wahlenbergia stricta (Campanulaceae) in 
South Australia. Trans. R. Soc. S. Aust. 122(4), 147-151, 30 November, 1998. 

A new South Australian gall midge, Dasineura wahlenbergiae, that damages shoot 
tips of Wahlenbergia stricta (R.Br.) Sweet, a common plant of grassy habitats in 
Australian and New Zealand, is described. Two leaves of the shoot tip of the host 
plant are malformed into a globular, hollow, hairy, partially discoloured gall, 2-5 mm 
in diameter. The male, female, pupa and larva of the new species are described. The 
new gall midge is the fourth Dasineura species known from Australia. 

Key Words: Gall midge, Cecidomyiidae, Dasineura wahlenbergiae sp. nov., 
Wahlenbergia stricta, South Australia. 


Transactions of tre Reval Society of §. Aust. (1998), 122(4), 147-184. 


DASINEURA WAHLENBERGIAE, A NEW SPECIES OF GALL MIDGE 
(DIPTERA; CECIDOMYHDAE) DAMAGING SHOOT TIPS OF WAHLENBERGIA 
STRICTA (CAMPANULACEAE) IN SOUTH AUSTRALIA 


by PETER KOLESIK* 


Summary 


KOLUSIE, P1998) Masitenia willenbergiae. w new species of gall midge (Diptera: Cecidomyiidac) danagine 
shoot lips of Waitenbergia s(ricta (Campanulaceae) in South Australia. Trans, Ry See 8. Auyt, 12204), 47 151. 


30 November, 1998, 


Anew South Australian gall midge. Dayinenta wohlenbervice. that damages sboot fips ob Walilenbergia sirickt 
(R, Bry Sweet. common plant of grassy habitats in Australia und New Zealand, is described. Two leaves of the 
shoot tip of the host plant are malformed into a globular, hollow, hairy, partially discoloured gall, 2-S min in 
diameter The male, female. pupa and larva of the new species are desenbed. The new wall midge is the fourth 


Dasineura species known (rom Australia, 


Key Words: Gall midge, Ceewlamyiidue. Dasinenra wahlenbergiae sp. nov... Wehlenbergia siricni. South 


Australia, 


Introduction 


The new gall midge deseribed here was found it 
malformed shoot fips of the tall blue bell. 
Wahlenbereia stricta (Ro Br) Sweet (Campan- 
uluecae) at) Moriah Cooservation Park, fear 
Adelaide, Wahlenbergia stricta is a perennial herb, 
(00-9000 nim high with large, blhe flowers and is 
common al grassy sites in various yegetation types 
throughout Australia and New Zealand ¢Siith 
1986). The plants grow on slopes at the Morialta 
Conservation Park and in the spring the shoot buds 
of many of them are moditied ime globular, tary 
gills. Some plants have all their shoot tips galled and 
consequently do nor reproduee, 


Materials and Methods 


Shoot tip galls on Wallenberaia strict were 
collected at Moriali Conservation Pairk on 1S 
September, 1996 ind brought to the laboratory where 
a few of the galls were pecled open and the 
developmental stages Of the gall inducer examined. 
Same of the galls contained young larvae, some 
malure larvae, some cocoons and others were emply 
The cocoons contaned either larvae or pupae. A 
small number of the mature larvae wits preserved in 
70% ethanol A few eocoons were torn open and the 
larvae and pupae preserved as above. The majority of 
the galls was laid on wet sand within a pot to allow 


' Deprrtmeat of Hortealiove, Vitieultne ain Oenology. Waite 
Cuyats, The University of Adelaide PMB | Glen Osmond 
S, Aust. 5064 


them to develop into adults. Pupation took place 
Within the galls. Emerged adults were preserved in 
70% ethanol Canada balsam mounts of the type series 
were prepured for microscopic examination according 
lo the technique outlined by Kolesik (1995), 
Measurements refer to the holotype and paratypes. 
The type specimens. and other material retained in 
ethanol, are deposited in the South Australian 
Museurn, Adelaide [SAMA], the Australian National 
Insect Collection. Canberra JANIC| and the Swedish 
Museum of Natural History [SMNH]. Dry samples af 
the galls are deposited inthe State Herbarium ef South 
Australia, Adelaide |SHSA], 


Genus Dasineura Rondant, US40 


Dasineura Rondant, P40; 12 & 17 
Proposed type species Vipile aisverbrii: Schrank, 
1803) Gagné eral. (1997) 


Dasineura is a large, cosmopoliti Yenus UF some 
2) species containing Oligetraphini with four 
scemented pilpi, toothed tarsal claws, an Ry wing 
vein that meets C oanteriar to the wing apex, and the 
female ciehth tergite divided (hte two longitudinal 
sclerites, 


Dasinenra wahlenbergiae sp. nov. 
(FIGS )-15) 


Holawpe: 3, Morialta Conservation Park, South 
Australia [34° 54° 8. 138° 44° BJ, 20.14.1996. P. 
Kolesik, reared from a shoottip gall of Wallenbergia 
aricia (R, Br.) Sweet collected 15,i,1996, 121384 
|SAMA],. 


148 P, KOLESIK 


Figs 1-6. Male of Dasineura wahlenbergiae sp. nov. |. Head in frontal view. 2. Last three flagellomeres. 3. Wing. 4. Last 
tarsal segment with claw and empodium, 5, Sixth flagellomere, 6, Genitalia in dorsal view. Scale bars = 100 um 1, 6; 50 
hm 2.4.5: 500 um 3. 


ANEW GALL MIDGE PROM WAMLENRERGIA STRICTA du 


Parniypes: 39 2.3 pupae [SANA, 121385-12 1390), 
PP 2 7 2 pupae fall ANIC] same dina but 
emerved 17-25.i8, 19962 3 larvae |SAMAY, 3 larvae 
[ANIC], collected with holotype, 


Omer materials 34 5 [SMNUL, same dati as 
holotype bul enierged 20,-25ax,1996: 37 Taryae. 5 
pupae within cocaons [SMNI|. eollected with 
Holotype: gall [SUISA, AL99747 199). collected with 
holotype. 


Deseripron 


Male (Bigs 1-04 

Coluur eyes bhicks bead. thorax und) abdomen 
orlinesred: leas, auitennie, pilpi setae and settles 
ufey. Walleles oralige brown, 

Hew; Antero seape square in frontal view; 
pedicel sphernide 10 fiwetlomeres. Crstund second 
fused, nevks as long or slightly longer than podes: 
CHCHMTKG coliprisnig Uwo Uupsyerse yn bwe 
Jongitudioal bands, Palpus four-seamented, segrrents 
progressively longer, Bye fieers rounded. eluse 
together excep on veriex where small area ol ne 
Hicets separtites the eyes, Labelle tapered clistally, 
Kilenlly wall © setae, Proms wath) 23-26 serie per 
stile. 

Thorax: Wing length 2.) mim (2.022. n = 2), 
wWidih OO inn (8-91 Rs joining C anteriorly to 
upers Ri jouing C slightly anteriorly to mid-leneth; 
Ks not obvious. Claws (nothed, emporia as lone as 
chiws, 

Abdomen: Tergites |S with pair of sensary sete 
ih ablerlor copners, lereites 1-7 With stile setal poy 
posteriorly and seules scutlered evenly, fergile & ta 
form of mimrow. selenatised, anterior bund wathoud 
seme. Stermites 2-8 with pair of sensory sete 
anteriorly, setie i Wide band anterierly ated 
narrower band posteriorly, area between Lwo bands 


of setae mere weokly selerorised. Genitalia: 
gonueoxile evlindrival. setose and setulose; 


BOHOSEVIUS Lupered distally, sparsely setose, Setulose 
basally up te “/) oF ats Tengu ventrally and '/> 
dorsally, sparsely striate beyond, bearing distal 
comb, cere) large. cach with several selue uipieally, 
selulose: hypoproct deeply and widely divided, with 
one sel on each Jobe, selulose: parameres sheathing 
vcdeagus. with subylobulit disteusions dorso- 
basally. with --S selose papillae upiealhy; aedeagus 
long, stout, 


Female (vise 7-10) 

Colours ais in nile. 

Heid: 16 (lagellomeres, evlindrical, with necks /\- 
yy node's leowth, creumbilte comprising iwe 
transverse and lwo longitudinal bands. distal 
Irinsvere band with loop, cireumililin attachment 


points very dense. Labella with 7-10 setae laterally. 
troos with 22-28 sete hiterally. 

Thorax: Wing length 2.) mm (2.42.3, p = 5), 
Wilt O24 nin (O8-O.9), 

Abdomen; Tergiles 1-8 with pair of seasory setae 
inanterior corners, tergiles 1-7 with single setal row 
posteriorly and seales seatlered evenly, tergite 8 
ciyicled THto two donaitudinal selertes, Stermites 2-7 
wilh pair Of sensory selae anteriorly, setae im wide 
hand anteriorly and narrower band posteriorly, urea 
between Iwo bunds of selue more wenkly selerotisud. 
sternite Snot developed. Qvipasitas protractiles 
elongate, O.7 mm (0.0-0.7) long (anterior Linnil of 
genial chamber to werminal tip distance), 31% (29- 
45) of wing length: cerer fused medially into single. 
prolonged, terminal lamella, setose und setulose: 
AV poproet WHA Wo setae, setalose. 


Prpe (Pig, 1) 

Colour antennal horns brawn at upex. remaining 
puts yellow, Length 2.00 mit CLS-200 nm = 5), 
Antenml horns small, pointed, Frons on each side: 
three frontal papillae two of thent setose. asetose one 
someumes ducking) three usctose lateral facial 
papilla, Cephalic papilla with seta 94 pin (LS8Y 
201) Jong. Prothorucre spiracle 230) pint (220-244) 
ling, trachea ending at apex. Tnresument af 
abdomind segments covered with spiculac shelly 
lonver dorsally, second through seventh abdominal 
seenicutls with group of dorsal spines ad anterior 
hall) First through eighth abdominal segments with 
wwe pairs oF dorsal asetosxe papillae, one pair ol 
Setose pleural papillae, Gve pairs af asetose ventral 
papillae, 


Last destar larva (Figs 12-141 

Colour: red. Lenwth 24 gin (2028 w= 6) 
integument covered with rounded phites about LO pin 
ti dhameter, veninilly wth several transverse rows of 
spiculae on anterior fall af thorieie ane abdominal 
semments, Plead swith postero-hiterul apodemes as 
Jon as hed lenguh. Spatula bilobed, with long shat, 
longth $47 jn (177-169). Papitlae charuererisue of 
Dasineura laksa (syWwen L975), 


Livinology 
The name wellenbersiae is derived trom the 
genene mome vl the host plait. 


Gall and hiolaen 

The new gall Widwe modifies bwo deaves ob the 
shoot bp at Wehlenbergia stricto into a globular, 
hollow, hairy, partially discoloured gall, 2-5 main 
diameter (Fig. 15). On (5 September, 1997, at 
Morita Conservation Park most galls contained 
mature lurve. but some gulls contined young 
lurvue, Some Cocouns With larvae or pupae within or 


150 P. KOLESIK 


‘ ; 
egy 


~ a 


Figs 7-15. Dasineura wahlenbergiae sp. nov.: 7-10 female, | 1 pupa, 12-14 larva, 15 infestation symptoms. 7. Posterior end 
of abdomen in dorsal view. 8. Posterior end of ovipositor in ventral view. 9. Sixth flagellomere. 10. Last three 
flagellomeres. 11, Anterior part in ventral view. 12. Spatula with adjacent papillae. 13. Head. 14. Two terminal segments 
in dorsal view. 15. Gall on Wahlenbergia stricta (R. Br.) Sweet. Scale bars = 100 um 7, 11, 14; 50 um 8-10, 12, 13; 10 
mm 15. 


A NEW GALL MIDGE FROM WAARLENKERGIA STRICTA IS] 


empty cocoons, und others Contained no remnants of 
the gall inducer, Up to 20 larvae were found within a 
gall. The adults reared in this study originated from 
larvae pupated within the galls. 


Discussion 


Dasinetiva, the largest genus of Cecidomyiidae, 
comprises species oecurring in all zoogeographical 
regions of the world, Four species are known from 
Australias DL aedeiaelongifoliae (Skuse, 1890) 
(Gagné & Marohasy 1993) and 2D. diedyé Riibsaamen 
(1916) which damage Mowers of Acacia longifolia 
(Andr.) Wild. (Mimosaceae) and A. evelopy Cun, ex 
Dou tespeetively, D, divbaath? Kolesik & Skuhravit 
(1997) which ts an inquiline in Mower galls on 
Hybanrhus floribundus (Lindley) Muell, (Violaceae) 
Wduved by an unknown gall midge. and the new 
species described here. Dasineura wallenbergiae sp. 
nov. belongs to Sylyen’s (1975) biological group EH 
of gall midges whose larvae are primary yall 
inducers, feed gregariously abd pupate in both the 
soib and the plant. ‘The adults of the new species 
reured if) the present study originated from larvae 
that pupated within galls, bul the fact that some galls 
were found empty with neither cocoon remnants nor 
parasitoids within. suggests that part of the turval 
population pupates in the soil. This conforms with 
the behaviour of Sylven’s (1975) biological group LI, 
Dasiniera diybanthi. the only other Australian 
species of this genus described in detail. belones to 


gruup LL of gall midges Whose Jarvae are inquilines. 
feed gregariously and pupate in the soil. The new 
species differs ftom D. liyhanthi in several 
morphological characters. In D, wahlenbergiae, the 
wing vein Rs is not obvious, the tooth on the tarsal 
claw is much smaller than the claw, the [emale 
Hlagellomeres are much longer than wide, in the male 
genitalia the gonostylus is tapered distally, Ue mitle 
cerci and parameres are nearly as long us lhe 
aedeagus, and the larva has no medial papillae 
between the terminal papillae. In D. Ayhanrhi, the Rs 
is evident, the tooth on the tarsal claw is nearly as 
larve as the claw, the female flagellomeres are as 
long as Wide. inthe male genitalia the gonostylus is 
abou! the same width through its entire length, the 
Male cerei and parameres are much shorter than the 
wedeagus. and the larva has several medial selose 
papillae between the terminal papillie, 


Acknowledgments 


] am grateful to H.R. Toelken. South Australian 
Stite Herbarium for the  identbication of 
Wahlenhersia sricta A. Stark, Halle Germany 
courteously provided a copy of Ribsadme's paper, 
Special thanks go lo J. D, Gray, Department of 
Horticulture. Viticulture and Oenology University of 
Adelaide, R. J. Gagné, Systemuue Entomology 
Laboratory USDA Washington DC, and E, Sylveén, 
Swedish Museum of Natural History Stoekholin for 
commenting onan early draft of the manuscript 


References 


GAAGNE, 1, J, HARRIS, KL ML SKEET A, ML. Satins. M 
amt Svives, Be (L987) Daamenra Rondam, [840 
(lise. Dipreni: proposed designanon oat lipali 
yviibel Solmaink, 1802 as the ype species. (Case 2986, 
Ball. Zoal, Nein, 54, 92-44 

& MAROUASY 1. (1993) The gall Widges (Diptera: 
Cecidomyiidie) of Acccig spp. (Miimasacene) a Kenya. 
lnseota Mundt 7, 77-| 24, 

Ko esik, BCP9oS) A new species ol Locinetieorme Fell 
(Diptera: Cecidomydidue) on Lucedypray fusetenfesa in 
South Australian. A Aus. enn Soe. 34, 147-152. 

& Skuneava, M. (1997) Dasmenra lybanthi 
spec, nov. wo Hew inquiline species of Ceeidemytidac 
(Diptera) from galls on Hybanthas  flaribundas 
(Violieeae) in Australia Sradi Dept 4. 240-246, 


RISAAMEN, le Lh (M916) Bentrae zur Renniniis 
ausserecuropitiseher Gallmiicken, Sifsaiueberichte ir 
Gesellachaft Natieforsehender Ereninde oi Bertin W915. 
43)-48 1. 

SKUSE, BAL AL (TRY0) Diptera of Australia Neratacens. 
Supplement 1, Pree, Linn, See, NSW (2nd series) F. 97 A- 
4l3. 

Svrvi, Bot (M86) Pamily Campunukivaie pp. tA76-| 443 
Mt Jessop. d. Po & Toelken. H.R. (Rds) Plor of South 
Australia, Part 1 (Polemoniiceae - Compositie)” (South 
Australian Government Priiting Division, Adelaide), 

Syivin. E. (1975) Study on relationships berween habits 
und externyl straciires in Olivotrophidi luryae (Diptera, 
Cecidomyiidae). Zoel. Seripta 4, 55-02, 


DEVELOPMENTAL BIOLOGY OF UPEROLEIA TALPA TYLER, 
DAVIES & MARTIN, 1981 (ANURA: MYOBATRACHIDAE) 


By MARGARET DAVIES* & GRAEME F. WATSONT 


Summary 


Davies, M. & Watson, G. F. (1998) Developmental biology of Uperoleia talpa Tyler, 
Davies & Martin, 1981 (Anura: Myobatrachidae). Trans. R. Soc. S. Aust. 122(4), 
153-157, 30 November, 1998. 

Uperoleia talpa is a small fossorial frog restricted to the southwestern portion of the 
Kimberley Division of Western Australia. The frog breeds in the monsoonal wet 
season, and lays clumps of eggs in single capsules in ephemeral ponds. Larvae hatch 
at stage 19. Later-stage larvae have strongly arched tail fins, a sinistral spiracle, 
extremely large, cavernous external nares and a larval tooth row formula of two upper 
and three lower rows of labial teeth. Labial papillae are clearly interrupted both 
anteriorly and posteriorly. Larval life span is about 71 days. The large and 
conspicuous external nares have been found in a further five species of Uperoleia and 
are suggested as a possible diagnostic character for some larvae of the genus. 

Key Words: Uperoleia talpa, larvae, embryos, generic character, life history, tadpole, 
Myobatrachidae. 


Tramaetions a the Raval Secien of S Yast (1998), 122(4). 153-157. 


DEVELOPMENTAL BIOLOGY OF UPEROLEIA TALPA TYLER, 
DAVIES & MARTIN, 1981 (ANURA:MYOBATRACHIDAE) 


by MARGARET DAVIES & GRAEME F, WATSON’ 


Summary 


Davirs. M. & Warsus. GF (1998) Developmental thology of Uperuleia talna Tyler, Davies & Marrin, 1981 
CAnira:Myobatrachidie). Jeans A. See. S. Aust $2204). 153-157. 30 November. [908. 

Uperoleia talpa iss small fossarial frog restficted to the southwestern portion of the Kimberley Division of 
Western Australia. The [row breeds in the monsoundl wel season, and lays clinips ofeges Ti sige capsules in 
ephemeral ponds, Larvae hath arstige 19, Literstage larvae have strongly arched tail fins. a sinistral spirale, 
extremely large. cavernous external nares umd a larval loath row formule of two upper und three Inwer rows of 
lubial teeth, Labial papillie are clearly interrupted both anteriorly and posteriorly Larval life span is about 71 
days. The lacie and comspicious external nares have been found ina further five species of Uperalear sind are 
suggested asa possible diagnostic charter for some binvae of (he genus, 


Key Worns: Uperoleia telpa. larvae. embryos, generte chacaeter, life listory. tadpole, Myohatrachidae 


Introduction 


Uperalen Gray. L841 is a genus of small bur 
rowing, myobulrachine frows with a wide-ranging 
distribution deress Austealia in areas Of poor winter 
rainfall, Prior la the revision of Tyler efal (1st), 
the genus comprised three species. bul with the 
description ol Uperieia altivsine (Davies er ul. 
1993), now ineludes 24 taxa. However, very Tile is 
know of the tarvad biology at the genus 

Moore (1961) deseribedt the larva ol U) merrvrereter 
(how eonsidercd ta bet! deevigar Keterstein, 1467 
(Davies & Liulejohn 1986)) whilst Watson & Martin 
(1973) described the larva of what was thought to be 
UU. mermorala. bul whieh is now considered to be a 
representative Of LL tyler? (Davies & Lillleyohn 
}O86). Tyler er el. (1983) recorded the lHfe history of 
Hh jaundape Vyler, Davies & Martin, 1981) Davies e 
al, (1986) described the larva of 00 dittomoda Tyler, 
Davies & Martin, 198T and Richards & Alford 
(1993) provided a description of (he larva of U. 
mintila Davies, McDonald, Corben & Ingram, LO86, 
Pull life bistory data of Lhese species are scarce, 

Uperoleia tipa ‘Tyler. Davies & Martin, I98T is a 
large member of the venus (males 26-40 mim 5-V, 
females 35-38 qn) CPyler en al 1994), with a 
restricied distribution in the southwestern portion of 
ihe Kimberley Division of Western Australia. The 
species Was originally described from three [rogs 
colleeted ona very dry night south of Derby (Tyler ey 
dh 198)) and the description has been amplified by 


) Peptol Agology. Liliversity ob Adelaide Astle 30Gb). 
( Deplol Zamlony, University of Melbourie Parkville Vie. 052, 


Davies & Martin (L088), who provided addigonal 
morphological, ostevlowieal aid distributional in- 
formation and described the call, In early Pebrusey 
1994 we collected ampleetant pairs of Lh talpe the 
spawn of which Wwe reared to metumnorphosis. thus 
allowing the description af the Tle history of the 
species (hat we report here, We also discuss sone 
features that may aid in generic recognition of larvae, 


Materials and Methods 


The sevies of Uperoleta talpa was reared. trom 
spawn deposited in phistc bags by ampleetant pairs 
collected in the field, Larvae were initially reared tn 
dwrated water at ambient temperature in the field 
hefore being transported to Adelaide where they 
were mulled i) a COnstail temperature room at 
30 41° C in dechlorinuted tip water, Larvae were fed 
on boiled organie lettuce leaves supplemented with 
commercial goldlish Wakes (Bioser), Material was 
preserved in Tyler’s Hui (Tyler 1942) and 
Hlustrarions were mude with the aid of a Wild M8 
sterea dissecting microscope with attached camer 
lucida, Mewsurements were mie using ai eyepiece 
micrometer 

Developmental stages are those of Gosner (1960), 


Material eanmined: Davies collection: Uperoleia 
talpa series: U. lithomiuda serws, U, allixsime series; 
U. dnundata series (basis of data used hy Tyler er al. 
1983); larvae of Crinta (Rentdella) scanifera (Girard. 
1853), 6. (Ro) riparia (aitleyohn & Martin, 1965). 
Peendophiyie VPitvinger, 1843. Tadpoles of &. 
luevivala were provided by Harold Phmann and 


| 54 M. DAVIES & CF WATSON 


(poles of (/ russedie (Loyeridge. 1933) were cx- 
amined i the collection of the University ol 
Michigan, Museum of Zoology, Ann Arbor, 
Michigan 


Results 


Pwo amipleenint pairs of Uperaleia talpua were 
enilecied al 2345 hoon 3.11, 1994 at a site (2.2 km 
south af the Gibtr River Rd tumoll on the roud south 
of Derby in the Kimberley Division of Western 
Australia (Figo 1). The night wits extremely hol, 
humid and stormy althouzh na rai fell in the 
Inmmednie area. Two other species of Uperalera (Ui 
mberad (Andersson, 1913) and 4 aspera Tyler 
Duyies & Martin, ISS) were calling at the same 
pom Upereleta dap was calling trom the Ury 
yeuetation tanhest trom the water, Uo sijeberus was 
calling from the edee af the water and Oo aaperie was 
calling From the intermediate areas, The choruses ol 
Oo defpecanel Uh apera were substantial whilst thar 
OF LL nyiherey was less viworous. Loarta rubella 
(rry. TAZ) und c velarene cistraliy (Gray. 842) 
were also culling around the pond. When we visited 
the same site the following evening. there was much 
less aeHivity With a single) feldme and cery tew L 
aspera calling, Avila time, tewly mecinorphiised 


in 


C. arvptotiy Tyler & Martin, 1977, CL loauipes Tyler 
& Martin, L977 and Cy australis Were located, 

The captive pars were retained in pond water in 
inflated plastic bags, supparted by feeereum 
containers. The U. Kepe spawned vurly on 6.111994, 
AL O700 on 71,1994, the eges hid reached fate 
scustrulit slige b2. A single capsule surrounded the 
oyun Mean capsule diameter of six egies wits LS 
mm (range 1.78-2.04) and the ova had a mean 
diameter of 138 mim (ring 1.30-1.60 mim), At 1300 
on 71,1994 embryos were at stave 17 (tail bud) (Pig. 
2) with the tail being better developed thar the Head. 
Hatehing (stage 19) was completed by TITS an 
Ai 1994 (Mig. 3). The newly hatehed birwae fad vo 
eaterial gills: (he eyes were very dillicull to defect 
ind (he mouth had not perforated: adhesive glands 
Were HOF pigmented ul this stage. 

By HOS on 11.1, 1994, same preserved lirvac webe 
already at Stage 25, The spiracte hud formed and the 
adhesive whinds were pigmented, There was no 
keraunmsation of the beak ar teeth (Wis. 4). 

Mulerial preserved on [241 1994 inelided some 
Iirvie stil at Stage 22/23 (hig. 3) in which the 
nostrils hil not perforated althouph the adhesive 
ghinds were pigmented, The spiracle hud oor toriiea 
aU ulthouwh the mouth was perlorated there was no 
kervinisation on (he beak or teeth 


Pin. bo Site abo wlttct ampleetan pairs OF Moerutera tafpa were collected. Se kit S Guth Bier Relient on Piehiway t 


eH al Pder liv, WA 


DEVELOPMENTAL BIOLOGY OF UPEROLEIA TALPA 155 


_ 


Fig. 2. A. Lateral and B. Dorsal views of Stage 17 (tail bud) 
embryo of Uperoleia talpa at 2200, Scale bar = | mm. 


Fig. 4. A. Ventral and B. Lateral views of Stage 25 larva of 
Uperoleia talpa. Scale bar = | mm. 


Sal eal 


Fig. 5. Lateral view of Stage 25 larva of Uperoleia talpa. 
Scale bar = | mm. 


Fig, 3. A. Lateral view of newly hatched larva at Stage 19. 
B. Lateral view of larva at Stage 22 of Uperoleia talpa. 
Scale bar = 1 mm. 


Tarte l. Measurement (in min) ef body and total length of larvae of Uperoleia talpa as mean and range. 
N = number of individuals. 


Stage (Gosner 1960) Body length (mm) Total length (mm) N 
11,-20.11,1994 25 4.19 9.66 5 
(3.84-4.64) (7.84-8.16) 
14.11.1994 26 3 8.0 4 
(3.2 - 3.6) (7.84-8,16) 
26.11.1996 27 5.44 12.8 - 13.8 2 
2.-29. 11,1994 28 5.46 13.72 7 
(4.8-5.92) (12.96-14.58) 
6.-29.i11.1994 29 6.46 15.33 5 
(5.44-8.48) (14.24-17.44) 
29.i11,-13.iy, 1994 30 6,99 16,69 3 
(6.08-7.52) (15.36-17.6) 
[3.iv. 1904 3] 6.08 16.00 | 
I3.iv, 1994 32 7.84-8.64 18.56-21.8 2 
13.iv. 1994 33 9.28 21.92 i] 
13.iv. 1994 34 8.72 21.8 4 
(8,0-9,28) (20.32-22.72) 
[3.v.1994 35 9.28-10.4 23,2-26.4 1 
13.iv.1994 36 10.72 26,08 3 
(10,24-11.2 (24,8-27.2) 
I3.iv. L994 37 9.92-11,2 25.28-26.58 2 


U3.iv, 1994 38. 10.8 25.12 i 


156 M. DAVIES & G. EF WATSON 


Fig 6, A, Lateral and B. Dorsal views of Stage 36 Larva of diperetedt tulpu, Seale bar = 5 mm. 


Fig. 7, Oral disc of Stage 46 larva of Upeviteta talpe. Seale 
bar= | mn. 


Larvae al stave 25 preserved at 1115 an 14.11.1994 
(Fig. 5) had perforated nostrils, which were round. 
but these were not as conspicuous us is seen at later 


Stages (see below), The horny beak was keratinised 
aod kerauinisation of one upper and 2-3 lower rows 
of labial teeth was beginning. Yolk sull remained in 
the ut. although the cloaca was open dextrally, 
Larvae remained at stage 25 until after 20.0).1994, 
period of up to 9 days since the onset of this stave. 

Measurements of larvae at stages 25-38 of Gosner 
are viven in Table 1 

The following description is of a larvaat Stage 36 
(Pig. 0), 

Body ovoid, widest at midpoint of body. Snout 
evenly rounded i dorsiul and lateral view. Nares 
dorsal and extremely large and cavernous. Eyes 
moderately conspicuous, Spirucle sinistral, short, 
opening dorsally and visible when viewed from 
above. Anal tube broad and dextral to ventral fn, 
Dorsal fin more strongly arched than ventral fin. Pins 
rounded terminally. Dorsal fin commencing on 
posterior part of body, deepest about half way along 
its length, Veotral fir commencing posteriorly to 
body. approximately same width along its length, 
Tail musculature moderately thick, tapering to fine 
point. Oral dise small and ventral. Labial papillae 
widely interrupted anteromedially. Also. interrupted 


DEVELOPMENTAL BIOLOGY OF LIPEROLEIA TALFA 1s7 


posteromedially. Two rows of upper leeth and three 
rows Of lower. the second of which is divided (Pig, 
7). Short P3 row supported on flexible flap, ‘Tail 
musculature and fins moderately heavily suffused 
with pigment. Small dark-brown islands of 
pigmentation un the body. 

Larvae reached metamorphosis at stage 46 on 
I5.iy, 1994, 71 days aller spawning, 


Discussion 


We have now examined tadpoles of six species of 
Upervleia, as wells several olber myobulrachine 
species, [eis clear that the external pares of muny 
Uperoleia larvac are unusuilly lire and cavernous 
(Vig. 6). OF the species examined, this feature was 
present mi all but (. imurdare, Richards & Alford 
(1993) provided measurements of €. minuda, but 
these do not allow ialiree: comparison of the data we 
provide here, since diuneter in relation to the width 
of the head could not be ascertained, These authors 
do not conmiment on the relative size of the nares. 

H, however, the nares ure nol particularly large, 
this feature would be useful in separating larvae of 
C) iimile (rom ff fithomoda - a species: pair in 
whieh the adults are difficult ta Sepirate beth mor- 
pholowieally, and, a high temperature. by call OK. R. 
McDonald pers. comm. 1986) 

There is no generic tooth row formula for 
Uperoleda, Uperaleta minuta, U. lithomoda and 
tnundata share a formula of 2¢2)/3 whilst U/ 


laevigata has a formula including no undivided rows 
of 3. Uperoleia talpa has 2(2)/3(2). whilst that of 
(!. altissima is 2(2)3(1.2) (Davies & McDonald 
1998). The dark tail tip recorded by Richards & 
Alford (1993) in early stage larvae, whilst shared by 
UL mimula, UL lithonioda UL laevigata and UU. 
ariysima (Davies & McDonald 1998), is not present 
in. talpa, The heavy pigmentation of O. talpa 
larvae is Shared by O/ tvleri (Watson & Martin 1972), 
The flexible flap supporting P3 labial teeth is 
recorded inal Uperafeia lo date, but is not unique Lo 
Uperoleia, being found in Criniat (Reanidelle) 
sienifera, C. (Ro) riparia and as a larger structure in 
Pseudophryne species (Watson & Martin 1973; 
Davies unpub.). Thus the possibility of using thes 
leature for generic recognition foreshadowed by 
Richards & Alford (1993), cannot be sustaited, 

The large nares may he uselul in some species 
assemblages, being absent in only OU. (auadera 
iimony the species wwe have examined to date, 

The unusual tail bud stage in whieh the tail is better 
developed than the head was noted alsa by Moore 
(M61) in U4. feevigata. This developmental condition 
merits further investigation 


Acknowledgments 


Fieldwork was supported by the Anstralian 
Research Committee and we thank Mo. Tyler foe 
companionship in the field and the referees tor 
helpful comments 


Reterences 


Davins. ME Ae Li tortoise, Med (86) Phe brag sents 
Uperalda CXnutihepadienytidie: in southeastern 
Australia, dics A Soe So Mash EM TED ps. 

_ — oN Maris, “AOA, (PORK) Redetiniian ot 
Uperoele tala Tyler, Davies & Martio, 188) CAnurie 
Leptodiery tiie: Myobutraelinae), fe 112, 47-89, 

& MeDosarb. BK. R. (1908) Develupmental 
hiology Of Upertein vitissime Davies, Walsor, 
MeDonild, Lrenerry A Werren. 18a CAN Ura 
Mycbatinelieliwe. Malek 22, 167 172, 

& CORBIN, ©. (IYsO) The rents 
Hiperafaia — Ciray CAnFE Leploduery Hetiey i 

Queensland, Australia, Hae, R, Soe, Wek 98. 147 E88, 

Wastin, Ch B MIC DOs anne Ke Ro TRPNs ay. M 
ode WiRRE SS, CE 1 1L9UR) A new apecies of Myrreralola 
(Anura:  Leptodjetylidae:  Myobatrachiniey tren 
fortheastern Australia, “tem, Old Atay, 33. 167 U4. 

Goss, KROL. C00) A simplified table for staging anaes 
embryos and Larvae Wil totes on identiteatiarn 
Herpotalowion Ube (R20) 


Mouikh. AS UIYOD The Togs OF eastern New South Wales 
Kull, Ain, Mus. Nat. Hive, (20, 140-386, 

Rieiiykos, Sb AR ALO, RAL (1995) The tadpoles ul 
twe Queenshind frees GN: Pylithie. Myabut 
rachidne), Mea Ghi Wis. 33. 337 340. 

Ty MA 1962) On the preservation atanurun Gidpales, 
Nest, J Set 25, 222, 

eCROOK. CAL ADAVIES. ML CIOS a) Reprodietiye 
bidlvey of the frags of the Maeele Creek Systeni 
Northern Territory, Mec S. Ata Was B.S hi 

Davis, Mak MARTIN AL ACTOS TE Adstentiar 
frogs of the leptodaerylid venus Upenvena nay, Vive ot 
fool Stippl Ser 7 1 O4, 
Satie bo AL Ae Teminstosr, RO (POU bis 
ot Western Australia’ Revised edicin (Western 
Anatraliin Museum. Berth). 

Warsom, Gob & MARIN A, AV T973) Lite histary, leval 
morphology and rehitionships of Australian lopradaery ia 
Hows, Lraas A Saw, S. Aur. OT ON dA, 


A NEW SPECIES OF FROG (ANURA: MICROHYLIDAE) 
FROM CAPE MELVILLE, QUEENSLAND 


By MARGARET DAVIES* & KEITH R. MCDONALDT 


Summary 


Davies, M. & McDonald, K. R. (1998) A new species of frog (Anura: Microhylidae) 
from Cape Melville, Queensland. Trans. R. Soc. S. Aust. 122(4), 159-165, 30 
November, 1998. 

Cophixalus zweifeli sp. nov. is a relatively large member of a genus of microhylid 
frogs restricted to New Guinea and the Cape York Peninsula of Queensland. The new 
species is found in boulder fields in the Cape Melville National Park. Females are 
characterised by having flame-scarlet axillae, groins and hidden parts of the hind 
limbs. Males have not been observed. The finger discs are expanded. Morphologically 
the species is allied to C. saxatilis, but unpublished mitochondrial DNA sequences 
link it with C. infacetus. The description of this taxon brings the number of species of 
the genus in Australia to 13. 

Key Words: Cophixalus zweifeli, new species, osteology, Microhylidae, morphology. 


Transactions of the Royal Sactery of S. Aust. (1998), 122(4), 159-165 


A NEW SPECIES OF FROG (ANURA: MICROHYLIDAE) 
FROM CAPE MELVILLE, QUEENSLAND 


by MARGARET Davies’ & Kerrd R. MCDONALD! 


Summary 


Davies. Mad MeDonNanh, KR t}998) A new apecies of frog (Andra: Microhylidae) from Cupe Melville, 
Queensland. Tas, A. Soe S. Aut (2204). 159-165, 30 November 1998 

Caphivalus caveifedi sp. toy. is wrelatively barge member of a genus of microlylid Trays restricted to New 
Guinean and the Cape York Peninsule of Queensland The new species is found. in boulder fields in the Cape 
Melville National Park. Females are characterised by having Clame-scarletaxillie. geoios un hidden parts of the 
hind fimbs, Males have not heen observed, The finger dises are expanded, Morphologically the species ts allied 
oC) sexes, bul unpublished mitochondrial DNA sequences lok it with Co dafdeenis. The description of this 
Jason brings the nuniber of species of Ihe genus in Australia to 13, 


Kiy Wonos: Cophiveles sveffedi few speaes, ostealogy. Microhytidae, morphology, 


Introduction 


Faun surveys have been conducted in Cape York 
Peninsula by the Queenskind 
Bavironiment (ind its predecessors) sinee 1975, 
Information on the vertebrate fauna of the area bas 
been reviewed by Winter & Lethbridge 1995! us part 
of Stave }ool the Cape York Peninsula Land Use 
Stody, Subsequently fauna and fora surveys in Cape 
Melville National Park have lovaled significant pew 
records for mammals, reptiles, frogs, earthworms anil 
verelition types (Stanton 1994+: Little & Hall 1996; 
Stanton & Fell 1996": Jamieson 197, McDonald 
1997, 1998, unpub.) The area of Cape Melville 
National Park was increased front 36 000 ha lo 137 
O00 hit in L995, thus incorporating @ greater diversity 
of habitats and an increase in the runge uF Hora and 
fauna in the park, The new area includes assemblages 
ol topography, geology and vexctation types unique to 
Cape York (Stanton 1944, so the Cape Melville 
National Park is can aued of proven and potential 
codenisnd (Covaeeviel & figean L978: Stanton & 
Pell }990% danieson 197: Me Donald 1997), 


© Dephel Zoology. Darversity ab Adetitley Ansisiiia S002 

) OM preecvalion Stiteey BAO. Depa hice ob Terrien iO 
Bos kad Alherin Olt ass 3, 
Wires J OW. Tragic POD TUS) Lerres tei) Gertolinte 
fot Cape York Penosebe Cape York Peatusuli tan tae 
Some Satgngh Resources Amatyae Prem, Bribie 
Cush, OF we at (ie Cee ordiitern Cigna 
SHhvstion. dh PECPOM @uypy Mebwilie Sutra) (rks Rasetirie 
Thar Cheer repiet Hor Hie Oticenshanel Departenn ot 
LWIPO nen ie Peri ite) 
Siwy 1 PMT DL EYSGO Kaniitiida) al Capa York 
(hikari rept faite Quive osha Deporte i) Deieaments 


Department of 


A large hylid tro (Liforia andiiremalin MeDonald, 
1007) was discovered in boulder fields of the Melville 
Ringe. In addition, a second few frog species was 
Tocated amongst boulders, This species wats 
recognised as a mentber of Microhylidae. a family 
well represented in New Guinea bur with Aastraliin 
representatives restneted to the — subtamily 
Genyophryinae in Wwe genera Cophivulus and 
Spheneplirvne. Aastralian microhylils are contined 
to hortheast Queensland, with the exception of 
Sphenophryne adelphe Zwerlel, TUS. a species 
found in the north of the Northern Territory (Tyler & 
Davies 1986), Australian microhylids were reviewed 
by Zweilel (1985) who recognised 16 species. sever 
of whieh he deseribed at that time. Richards ef a, 
(1994) described Cophivaluy monticola (rom the 
Carbine Tablelands, northeast Queensland and here 
we deseribe a further Cophivalus from Cape Melville. 


Viaterials and Methods 


The omatertal stodticd is deposited) in the 
Queenstind Museum. Brisbune (QM) and the South 
Australian Museum. Adelaide (SAMA). Measure- 
ments were made with dial calipers reading wo 0.01 
mm, Measurements tiken (ii mind) were > tympanum 
diameter (T), eye lo naris distinee (hb), eye 
diameter (TL). foot (F). hand (Hi), head with (EW), 
fiewd lengih CHL), internarial span (IN), snout ty vent 
length (SV). tibia length (PL). width oF third finger 
dise and of penultinite phahinx. width ot fourth toe 
disc qind of penuldimate phaliis. length ob hand and 
lenwth of foor and follow Zweifel (1985) tind Tyler 
(1068), Material was cleared apd stained using at 
modification of he method of Dingerkus & Uhler 


lal) M DAVIES A KR MCDONALD 


(O77), Description aid discussion ol Ostvolory 
Lollow ZAwenlel (1985), 


Results 


The new speeies ts assigned to Copliveluy on the 
basis of the following features: denuiries mot in 
conlict; vertebral Coluinm prococlous: longue !/y [ree 
hehind with ne furrow or pouched pocket: maxillie 
nat ome comtiet (relationship) with  premuasilite 
indaiernmnable). ‘This combination af feattires assigns 
the species to the Genyophryninae (Zweitel 1971), 

In addition, the species jacks procoracoids and 
vlavicles. has snout that is not narrow er clongate 
ind lueks at hypertrophied serous sland an the snout, 
The other defining feature oF Caphivalis, the aliry 
Process being typically slender and not merging 
insensibly into the body of the bone, could not he 
determiiued, 


Cophixalus zweifeli sp. nov, 
(FIGS 14) 


Holoiye: \' QM Jodess (formerly QNPWS 
N29789) Cape Melville National Park, 14° 15! 3S, 
eh? 27 40 BE, altitide 60-80 mi. 17,1, 1995. Coll, Ke. 
R, McDonald and L. A, lackson, 


Paruiypess & SAMA RS1L080 16.41.1995, Same 


location and collectors as holotype: 2 QM J64889 


y+ 


Hi LP yatidiee fet sp. aM VTE ES Woah Sain 


(formerly (QNPWS N73038) Cape Melville 
National Park, Permwinent Camp Qld (rear type 
locality), allitude 40 m, T4.411,1995, Coll J, O'Shea 
(cleared and stained) 


Definiiion 

A large species (YY 40,1-45.4 mm SV) wilh long 
legs, large finger dises with third finger dise larger 
than fourth toe die. ai elonyate snout: dorsal 
eolounmtion brown with Mamescarlet axilla, thigh 
Nushes tind ventral lon markiigs. 


Deserimion of Holonype 

Head slightly narrower than body) legs moderately 
long CPL/SV OST), snout truneate from pbove, 
straieht and slightly projecting in profile (rigs 1,2) 
canthus rostralis straight, loreal region steeply 
sloping: nares anterolateral on Up al snout eye to 
hares stance ereaer Chon iitornianial span CliEN/TN = 
1.125), eyes moderately urge, commend! outline cleautly 
visible from beneath, interorbilal width greater than 
widlh of upper eyelid. Tympanim linge. obseure 
dorsally, diameter greater than hall eve diameter. 

Relative lenuths ab fingers 3=d=2=1, the first 
slender and approximately hall the length of the 
second (Fig, 3). Dises of lingers 2-4 greally enlarges 
and trumeate, That of first barely extending, beyond 
Width OF penultimate phalans (Fig. 3): sabarticuler 
tubercles founded. moderuely prominent. bow 


a 


4 


NEW SPECIES OF COPHIXALUS If] 


Fig. 2. Cophixalus oweijeli sp, nov, A. Lateral and B. Dorsal 
views of head of holotype (QM J64888), Scale 
bar = 5 mm. 


ovoid inner and outer palmar tubercles. Relative 
lengths of toes 4>3>5>2>1 (right foot). toe three 
abnormally short on left foot; length of first toe 
approximately half that of the second. All toes with 
enlarged truncate dises with terminal grooves. Discs 
on first and fifth toes smallest and approximately 
same size. Toe dises smaller than those of fingers 
2-4 (Big, 3). Subarticular tubercles rounded, 
moderately prominent. Low elongate ner meta- 
tarsal tubercle. no Outer metatarsal tubercle. 

Dorsal and ventral surfaces smooth. 

Colour and pattern: dorsum tan with darker brown 
pigment spots above insertion of arm, alony flanks 
and superior to inguinal region and along 
midvertebral region; large faint mark between and 
posterior to eyes: dark canthal stripe [rom tp of 
snout, through nostril and eye and above and slightly 
posterior to tympanum. Pale crescent along anterior 


R) 


Fig. 3. Cophixaluy cweifeli sp. nov. A, Palmar view of left banc. B. Plantar view of right foot of holotype (QM 64888). 
Scale bar = 5 mim. 


162 M, DAVIES & K. KR, MCDONALD 


rim of tympani and paler stripe along lower rinr al 
eye, Dark brown pigment patehes on dorsal surfices 
of hand and forelimb, Lesser pigrient spots on dorsal 
surkive of Took dark patches along anterior edee of 
tibia, Throat very lightly dusted with pigment, more 
concentrated around margin of jaw and speckled 
with white, 


Measurementy 

SV 40,L0TL 20.6; IW 13. 9 TLL 13.4: B45; 73.1; 
RN 3,6; IN 3.2; third finger dise 2.2 (penultimate 
phalanx 1.0); fourth toe dise 1.9 (penultimate 
phalanx 0.71); hand 11.7: foot 7.3: ThISV O31; 
HW/SV 0.35; HL/SV 0.33: TIN/IN 1.125; HIL/AW 
0.96, BESY Ob IN/SY 0.08. third tager dise/SV 
O.054; fourth toe dise/SV¥ O,047; hand/sV 0.20: 
foot V O45: ENS V (0902 7/E ULAe 


Colour ia life 

Dorsal surhice beige when lirst observed at night 
darkening to lan during Ihe day with widely seatleredt, 
irrenulie brown speckles. Brown moti on aim wd 
thigh dorsal surface, Black canthal streak trom snout 
(rough eye and above tyimpianin, Lateral dark 
brown mottled marking between axilla aml groim-. 
Axithi gromn. hidden parts of thigh. ventral dbia aod 
miner halt of Foot flume searlet (Smithe 1975). Ventral 
surtice densely mottled Fight purple on throat and 
ches! becomme more ditfuse posteriorly. Ventral 
sutfiaee of femuciund arm mottled with brown, Dull 
yellowish waste on lower third oF abdomen and ander 
the femur, Brown ventral surface to hand and foot. 


Varnintan 

The iwe paritypes have the following meisure 
Mens: 

SAMA RSIO80: SY 41.5; TL 22.2: IW 128: TT 
Ide BOS05 LE 43s RN 40k IN 4.2: third Hinwer dise 
25 (penultimate phalang 12)e fourth foe aise 17 
(penullimare phakins O.6% hand lds fet TT, 
PLS 0.54: DIW/SV 0,40; HEYSV 0,45; AYES O95: 
TNS TW 113, BYSV 0.12) TIN/SY O18 (hint finger 
ise/SV 0.06; Fourth toe dise/SV O.04; laml/S'¥ (29: 
thovSV 0.46: EN/SV (1.006, T/E 0.66. OM [6g885. 
SV sh Th 20.3 IW 14 2: HE IAL be oP 34: 
PN SAIN Ay did finer dive 245 (penultimate 
phaking Lay: forth te clise 19 (penualtiniite plains 
Uoer hand bOo8: foat 1S: TES Gate: HAW SV O32: 
HLiSV 0.475 BN/IN 108; TILZTIW 0.92; BySV 0),U8: 
TN/SV O77. hid finger tite’ ¥ INS 3; fourth toe 
HiseYS¥ 0.0425 hand/SV 0.242 thovS¥ 143, PNVS\! 
WNT be ORS 

Dori) caleut os more hrewe than tin i SAMA 
ROVORD ciel He nurkitges are more clistiet The 
SCenN a! SUF. In puticnloralthe thine and pheno 
WHOM er ds flore eavaly ails irrenuda hl) prerie ted 
walla Laral witike stripe amehialhy. The undersurtice 
ul the (highs ts rere heavily spechted, 


Comparison with ather species 

Cuophixalus cweifeli sp, nov. is uw very large species 
of Australian mierohylid comparable only with ©. 
sevadiliy Zweifel & Parker, 1977. In addition, the 
(hird finger dise of the Hew species is hunger than (hat 
ofthe fourth tue, a feature shared by Cy sewerifiy aml 
©. vrnares (Pry, 1912). This latter species is smatler 
than eiher CL aweifeli ar C. saxarilis, The eanthus 
rostralis is straight in C, Sweijfels compared with a 
pounded canithus in Cy seediliy, The snout of ¢, 
cwelfele is longer than thal of ©. seveailis. The 
distinctive Thame-searler colouration on the hidden 
surfaces af legs is not found in any other Caphivalies 
it Australia. Peniies of ©. yeaediliy ure canary 
yellow at night, darkening tow light tan during the 
day, Unpublished cata of ©. Hoskin from 
mitochondrinl DNA sequences show C. pieifels lr be 
i sister taxon to CL dafacenis Zweitel, 1985 and ia 
separate clade from ©. savediliv, Cophivatuan 
infeicetis is asmall species (females te 17-6 mm SV) 
with a rounded eanthus rostralis. features not shared 
by C\ cweifeli. In life, CL infaeetis is dark grey on (he 
uiderside, compured with [he purplish colour of the 
(hroat and chest ole. cweiletr. 


Osreology 

One paritype wis cleared und stained, but 
untortinitely becuse of poor preservation. the 
material did not remain iitwet throughout the 
indeeration process. However, charicteristie and 
diagnostic features were obtainable, 

Skully The skull is lootiless with well-deyelaped 
and well-ossified nasals and frontoparielals. The 
ybanrtojigal arrieuhwtes with the minithi ‘Che 
eleutheroenathine condition af the premaxillae, 
typical OF the Genyophryiie, could nou be 
confirmed, The oloceipilal region (prootic and 
exoceipial) is ossified and the bones are vlosely 


aansociited with each other The vemers have aw well 


developed transverse arin (probable fised vomer yc 
pulling) arising froth an expanded arew in the 
inidline of the palate. and an anterior ani that passes 
mesial to and then anterion to the jiternal mares. he 
lraunisverse arm teaches The manithiry shell althoueh 
remaining lied Uy the maxilla by cartilaee (Pig. 44, 
The pleryuoid is extremely robust 

Mhete ts u thickened median portion al the lye 
plate and. (he posterior cornu huve well-developed 
Nanges it ig 4). 

The pewtoral girdle lacks chreictes wid a Sery saul 
medial projection niw represen| a yestioal 
ormostembin (Pig. 4). Cale icanen is whsenr in the 
mesosterimal region 

Presnerml vertebrae ure nate-inbricnte, Relative 
Willlhs af Lrninsverse pricisses andy ISS DelVotl 
SVSVISVIISVITL Vestiziol taunsverse priesses lin 
APPUreAL on the Urosty le 


NEW SPECIES OF COMIINALUS lo 


Fie 4. Caphiivelis sweifeli sp. nev. AJ Right yvomerine bone in ventral view. B. Dorsal view of hyoid plate. C. Ventral 
elements af pectoral girdle, D. ‘Terminal phalanx of finger (Paratype QM Jo4889). Scale bars = 1 mm. 


‘The tps of the terminal phalanges of the hands and 
feet are ‘T-shaped (Fig. 4), 


Comparison with other species 

Zweitel (1985) examined the osteology of 1) 
species of Australian Cophixvalus and the current 
comparison is with these data, 

The otoceipital region of C. zweifeli is similar to 
that of CL savatilis, C. concinnus Tyler. 1979, and C. 
exiguus Zweifel & Parker, 1969. Cophixelus 
infacetus and C. hasmeri Zweitel, 1985 have the 
ossification of the prootics restricted to buried 
nubbins, as seen from above. The other species 
examined by Zweitel have an intermediate condition 
between these two extremes, 

The fused vomers and palatines of C. cweifeli 
approach those of C. coneinuus in their relationship 
with the maxillary shelf. whilst the mesial extension 
upproaches that of C. orntatus, The anterior portion 
of the complex approaches that of C. saxurilix 
although itis more robust in C. zweifeli. 

Zweitel did not recognise characters in the hyoid 
us being useful in interspecitic comparisons. 

Some Cophixelus (including C. sexarilis) have a 
small cartilaginous protrusion on the anterior ventral 


midline of the pectoral girdle (7 vestigial omost- 
ernum). A smaller process is apparent in C. cweifeli. 

The terminal phalanges lack a median notch found 
in C. infacetus, C, saxatilis and C. ornatus. 


Distribution 
The species is known only from the type locality in 
Cupe Melville National Park. 


Hahirat 

The habitat of C. zwe/feli sp. nov. is restricted to 
boulder fields of Altanmoui granites (Fig. 5). The 
holotype and paratype (SAMA 51080) were located 
at the base of rocks at night near a ereek flowing 
through the rock formation, No calling was heard. 
Paratype (QM J64889) was found on a rock in a 
stream flowing out of the boulders. 


Kivmology 

This species is named for Richard G. Zweitel, 
former Curator of Herpetology at the American 
Museum of Natural History, New York, whose 
revision of the Australian microhylids is a standard 
reference. We honour his contribution to herpetology 
and his friendship. 


lod MM. DAVIES & Ik. 


Discussivn 


Morpholiwically Cupliccdiay swerfell appears lo he 
a SISTET Spec wetoc \waeilis Bathare Titer Vr iN; ihe 
lest of any oVisteahian microhylil and semnilar m 
only jUOpartions Che fame-searler colgutuen m the 
AN are amt on the legs is uniqoe WO) cwetfele 

bh auldition to morphologiont appemrenee. 
ciel und C. wawveeliy uclive Similar habitats al 
eranitic boulder fhelds with patches at 
veeehilion iemost pockets (Pig. 5: 
Parker 1977, Pics 7 for the habitat of ©. 
This Taem oF taba is restrivted to the Melwatle 
Range ind Black Mountam im Cape York Peninsula, 
Sith sonal aveas of just a few hectives are found 
in nimenws locelons it eastern Queenstind 
(Sianton $994) The direer distanee Tront Black 
Mountain ty the Melville Ringe is (75 ki, Ramfall 
araynd the Cape Melville Rane is estimated 1 be as 
Hiwh) as 2000 mim (seme 700 iim bisher than the 
surrounding country) (Stanton booed’). 

Nouvithsiinding the merphologienl taku abe 
owedeli with CL savartlis, Tloskin™s dane trom 
mitochondriil DNA sequences fink the specurs wilh 
C oinfveems in separate clade from ©, wavarslis 
Zweitel (LOSS) atempled ta denve a tree ot 


closed 
see Fawejlel & 


WE WALES). 


melitionships iimonest Cuphivalis usm extertol 


{Me DONALD 


Ms 


momphohwicul characters bul fem thas foo be 
“ursaislying (Aweilel PRS p. 470) 


net belidve that uny ane af his miest plrsinenions 


Zwerlel did 


lites Woe detemsible, Given We Porn-congiueroe 
mourphilesicnl anu biechermtenl lata 


here. ito ots vleue tr a robust 


beiw cen 
micheuted more 
marpholowical study ust other morpholoeres (ie 
exrenvial featares aS Needed eas a lostat the roObustess 
of the mitochondrial DNA diliie Th the allie ane 
copious, Indepemdem and evenly cisiributee aevess 
the branches of the tree, phylogenies Cream any dary 
(494) gine 
coneruenee between trees trom diferent cutie sets 


set fend to converge (Mishler stich 
provides strong ewilente far any hypothesis ol 
phylugenerie® bistary 

Hos clear bowevor thar whitewer the data set ised 
i derive felanonships, monophy fy iy 
Coplivdas Hest most be deniwnstrited 


4iistralun 


Acknowledgments 


We thank L. Jickson and J. O'Shea for feld 
assistamwe and Co Lloskin for permission ti quete 
unpublished dita from his BSe tHoms) Chess 
and further Studies, M. J, Tyler critically read the 
manuseript We oalo thank the refenses fot can- 
SPH Wet SUSOUSTIOIA 


Mark where Caliitilies Wedfel sy HOY IN hold 


NEW SPECIES OF COPHIXALUS 165 


References 


CovaAcevicH, J. & INGRAM, G. J. (1978) An undescribed 
species of rock dwelling Cryptoblepharus (Lacertilia : 
Scincidae). Mem. Qld Mus. 18, 151-154. 


Dincerkus, G. & UHLER, L. D. (1977) Enzyme clearing of 


Alcian Blue stained whole small vertebrates for 
demonstration of cartilage. Stain Technol. 52, 229-231. 

JAMIESON, B. G. M. (1997) Some new and_ previously 
known earthworm species from Cape York Peninsula 
(Annelida: Oligochaeta; Megascolecidae) Mem. Qld 
Mus. 42, 233-270. 

Litre, A. & HALL, L. S. (1996) Preliminary observations 
on the bats of Cape Melville National Park. N. Qld Nat. 
34, 53-57. 

McDonatp, K. R. (1997) A new stream-dwelling Litoria 
from the Melville Range, Queensland, Australia. Mem. 
Qld Mus. 42, 307-309. 

(1998) First Queensland record of the burrowing 
frog Cyclorana cryptotis Tyler & Martin, 1977 
(Anura:Hylidae). Trans. R. Soc. S. Aust. 122, 85-86. 

MISHLER, B. D. (1994) Cladistic analysis of molecular and 

morphological data. Am. J. Phys. Anthropol. 94, 143-156. 


RICHARDS, S. J., DENNIS, A. J.,. TRENERRY, M. P. & WERREN, 
G. L. (1994) A new. species of Cophixalus 
(Anura:Microhylidae) from northern Queensland. Mem. 
Qld Mus. 37, 307-310. 

SMITHE, F. B. (1975) “Naturalist’s Color Guide” (American 
Museum of Natural History, New York). 

TyLer, M. J. (1968) Papuan hylid frogs of the genus Hyla. 
Zool. Verhandl. (Leiden) 96, 1-203. 

& Davies, M. (1986) ‘Frogs of the Northern 
Territory’ (Conservation Commission of the Northern 
Territory, Alice Springs). 

ZWEIFEL, R. G. (1971) Results of the Archbold Expeditions 
No. 96. Relationships and distribution of Genyophryne 
thompsoni, a microhylid frog of New Guinea. Amer. 
Mus. Novit. 2469, 1-13 

(1985) Australian frogs of the family 
Microhylidae. Bull. Am. Mus. Nat. Hist. 182, 205-388. 

& PARKER, F. (1977) A new species of frog from 


Australia (Microhylidae:Cophixalus). Am. Mus. Novit. 


2614, 1-10. 


DEVELOPMENTAL BIOLOGY OF UPEROLEIA ALTISSIMA 
DAVIES, WATSON, McDONALD, TRENERRY & WERREN, 1993 
(ANURA: MYOBATRACHIDAE) 


By MARGARET DAVIES* & KEITH R. MCDONALDF 


Summary 


Davies, M. & McDonald, K. R. (1998) Developmental biology of Uperoleia altissima 
Davies, Watson, McDonald, Trenerry & Werren, 1993 (Anura: Myobatrachidae). 
Trans. R. Soc. S. Aust. 122(4), 167-172, 30 November, 1998. 

Uperoleia altissima is a small fossorial frog restricted to upland areas in northeast 
Queensland. The frog breeds in the monsoonal wet season, and lays clumps of eggs 
that fall to the floor of ephemeral ponds. Larvae hatch at stage 19. Later-stage larvae 
have moderately strongly arched tail fins, a sinistral spiracle, large, narrow, 
cavernous, external nares, and a larval tooth-row formula of two upper (second 
divided) and three lower rows (first and second divided). Labial papillae are strongly 
interrupted both anteriorly and posteriorly. Later-stage larvae are strongly pigmented 
although the strongly-pigmented tail tip of earlier larvae is less so. Larval life span is 
about 39 days in captivity. 

Key Words: Uperoleia altissima, larvae, embryos, life history, tadpole, 
Myobatrachidae. 


Fransaettons of the Royal aectery ofS. Aust (1998), E224), 16772 


DEVELOPMENTAL BIOLOGY OF UPEROLEIA ALTTSSIMA DAVIES, WATSON, 
McDONALD, TRENERRY & WERREN, 1993 (ANURA:MYOBATRACHIDAE) 


by Marcarer Davies’ & Kern R. McDoNaALb 


Summary 


Davies, M, & McDonanion KR. (1998) Developmental bialagy of Uperoferds aid Gin Davies. Watson. 
MeDonald, ‘Prenerry & Werren. 1993 (Anurw Myobatrachidie). Pras. Ae Soe. S. Ause 122(4). 167 172. 30 
November, (998, 

Unperaleia alixsines is asmall fossorial frog resinictes 10 upkind areas in northeast Queenshtud, The frog breeds 
ithe monsoonal wer season. and keys elueips of eggs that fall te the Toor of ephemeral ponds. Larvae Taich al 
stage 19. Lalter-stage larvae ave moderately strongly arched tail Tins, a sinistral spiracle, large. narrow, 
cavernous. extemal nares and a larval tooth-row formula of lwo apper (second divided band three Tower rows 
(lirst and second divided), Labial papillae are strongly interrupted both anteriorly and posteriorly. Taiterstage 
larvae are strongly plemented although the svongly-piemented tail lip oF earlier Lirvae is Tess so, Larval tle span 


ts about 39 days in caprivity 


KY Wokds) Uperledd altivstine, hurvaed. crbryas, lle listory. tadpole, Myabatraehidie 


Introduction 


Uperolerd altissiita Davies, Watson, McDonald, 
Trenerry & Werren, 1993 1s a Woothed member of a 
speciose venus of small fossorjal frogs distributed 
across mainiind Australia except for the southwest 
of the continent. Uperaleia altissimea is confined to 
elevated sites on Ue western Wet Tropics 
Bingeoyraphic Reyion (Stanton & Morgan 1977) 
from Princess Hills, Lumbolz National Park. north bo 
the Windsor ‘Tablelind of northeastern Queensland, 
The species is found tn moist cucalypt forests and 
woodlinds above 600 metres. Although deseribed i 
1995 from freshly collected material, the species had 
been collected but vot identified previously. Little 
was known of its breeding biology. 

In carly February 1997, We cneountered a breeding 
chorus af the froe following heavy miinkdl at a site 
onthe Atherton Tablelands. Amplectant pairs which 
luier Spawned were collected, and the resultant 
larvae were reared to metumorphasis. 

The deseriptian of this life history adds to the 
scarce dita available on dite histories of the 24 
species of Uperolere (Moore 1961; Watson & Martin 
1973; Tyler e/ al. 1983: Davies ef al, 1986; Richards 
& Alford 1993: Davies & Warsan 1998), 


Materials and Methods 


The series of Uperalera altissmia was obtained 
from spawn deposited by amplectui adults collected 


Depl al Aoolwy. Criiveriiy of Adelie Anata S005, 
7 Conservalidn Strateey Branch. Peparnnientol Lnvironmedt, Po 
Box S47 Wherton Ole 4883_ 


in the field. Larvae were initially reared in aerted 
water at ambient temperature (water temperature 
approximately 24° Cy) in the field before being 
iransported (a Adelaide Where they were maintained 
in wv constint temperture room at 3h + 1 Can 
dechlorinated tap water. Larvae were led an boiled 
organic lettuce eaves supplemented with come 
mereial goldfish flakes (Biosera). Material wis 
preserved mi ‘Tyler's fluid (Tyler 1962) and illus 
trations were made with the wid ola Wik M& sterco 
dissecting microscope with attached camera lucida, 
Measurements were taken using an eyepiece miero- 
ineter, Developmental stages are those of Gosner 
(1Y60), 


Results 


Four amplectant pairs of Uperdlita alussiie were 
collected on 1.4.1997 at a quitery near Carriigton 

“alls (17° 19" SI" S, 145° 26! 42" &). The site tsa 
quarry with gravel pits some of which have regrowth 
vewetation, We had visited the site on the previotis 
night utter min, but although O. adtissena was 
calling, no breeding was observed, However, Litaria 
rubella (Gray, (842) was calling and breeding took 
place later (hat night. Other species calling when Li. 
alussima was breeding included Crinfe remota 
(Tyler & Parker, 1974). 2. rasure (Gray, 1842), 1 
rothii (DeVis, 1884). 0. fallax (Peters. S81) and 
Limnodvaastes: terracresinae Very, VOUS. The night 
was humid following torrential rain in the nearby 
Herberton Range, although rain did not appear to 
have fallen at the site. 


Males were culling from the gravel areas 
surrounding the temporary pools (Pig. 1). often 


1M M DAVIES All. RL MeEDONALID 


Pig, 1. Callin tale Uperaleke aliisinie ab Careington Balls Quuirey site (SY approximately 24 nim}. 


ae 
SS". - 


hee 2 Atnpleenint (aierdeda adnan a Coecipton Pah quarry st (SY oat nile approwonately 24 mn 


UPEROLEIA ALTISSIMA DEVELOPMENT loo 


facing away trom the water toward the surrounding 
vegetation. Pairs in inguinal amplexus were found 
moving toward the shallow water (Fig. 2) 

The U. altissimea spawned early on 2.11,1997. At 
1310.0n 3.41,.1997, the eags had reached stage 12. late 
vastrula. A Single capsule surrounded the oyun. 
Mean capsule diameter of four eggs was 2.27 mm 


(range 2,22-2,32) and the ova had a meun diameter of 


1.36 mm (range 1.32-1.40 mm). At 1300 on 4.41.1997 
embryos were at stage 17 (tail bud) (Fig. 3). with the 
tail being better developed than the head. Adhesive 
elands and the stomodaeal pit were prominent, They 


Fie. 4. A. Dorsal. B. Lateral views of Stage 17 (tail bud) 
embryo of Uperoleia altisstna, Seale bar= | mm. 


had reached stage 18 by 5.11,1997 (Fig. 4). Hatching 
(stage 19) was completed on 6.111997 (Pig, 4). The 
newly hatched larvae had no external gills: the eyes 
were very difficult to detect and the mouth had not 
perforated: adhesive glands were pigmented at this 
stage. 

By 0900 on 7.i,1997, some larvae were at stage 
20. Adhesive glands were well developed on stalks 
and both the mouth and the external nares were 
perforated. Larvae were still at stage 20 at 1140 on 
8.111997, the cornea was not transparent. and heavy 
pigmentation was apparent on the tail fin. 

Larvae had reached stage 26 by L000) on 
14.11,1997. The horny beak was keratinised as were 
upper und lower labial tooth rows, The adhesive 
glands were reduced to patches of pigmentation. The 
nares were large and cavernous and the tip of the tail 
was particularly heavily pigmented (Fig. 5). Stage 2% 
was reached by 1100 on 19.i7.1997_ 

Later-stage larvae lacked the heavily pigmented 
tail tip. 


Pig. 4. A. Ventral. B. Lateral views of Stage 18 (muscular 
response) embryo. C, Lateral view of newly hatched larva 
of Upervleta altissima at Stage 19, Seale bars = | mm, 


170 M. DAVIES & K.R. MCDONALD 


hiv. 6 A. Lateral. BL Dorsal views of Stage 34 larva of Uperaler alrissinia, Seale bar= 5 mon. 


UPEROLEIA ALTISSIMA DEVELOPMENT 171 


Tarn tl. Measurements (in mm) of body and total lengih ef larvae of Uperoleia altissima as mean and range. 


N = number of individuals, 


Stage (Gosner 1960) Body length (mm) 


28 6.16 
(6.08-6.24) 
29 6.83 
(0.36-7.36) 
30 6.72 
(6.72) 
32 7.84 
35 8.42 
(8.00-8.64 
36 9.36 
(928-944) 
37 936 
(9.12-9.60) 
38 9.09 
(8.64-9.6) 
39 10.08 
(10.08) 
AQ 9.60 
4] 974 
(9,6-10.56) 
42 10.00 
(9.99-10.08) 
43 10.27 
(9.6-10.94) 
44 10.24 
(10,08-10.40) 
4s 10.44 
(10.4-10.56) 
4h 10.24 


(10.08- 10.400) 


Measurements of larvae at stages 28-36 are given 
in Table |. 

The following description is of a larva at Stage 39 
(Pig. 6). 

Body ovoid. widest at midpoint. Snout evenly 
rounded in dorsal and lateral view, Nares dorsal, 
Jarge. narrow and cavernous. Eyes conspicuous, 
Spiracle sinistral, moderately long, opening 
postertorly and searcely visible when viewed trom 
above. Anal tube broad opening dextral to veniral 
fin. Dorsal fin more strongly arched than ventral fin, 
Fins rounded terminally. Dorsal tin commences on 
posterior part of body and is deepest about halfway 
along 1s Jength. Ventral fin commences posteriorly 
io body and is deepest about halfway along its 
length. Tail musculature moderately thick, tapers to 
point. Oral dise small and ventral. Labial papillae 
widely interrupted aateromedially; less widely 
interrupted posteromedially, Two rows of upper 
teeth, secand divided: three Jower rows, first anel 
second divided (Fig. 7). Short P3 row supported on 
Mexible flap. Tail musculature and fins heavily 


Total length (mm) N 
13.68 2 
(13.12-14.24) 
16.53 3 
(15,68-17.12) 
16.56 2 
(16.32-16.8) 
19,20 I 
19.31 3 
(18.24-20.48) 
23.68 7 
(23.52-23.84) 
23.68 4 
(21.92-24.96) 
24.08 6 
(22.24-25,92) 
26.08 2 
(25.92-26.24) 
25,28 | 
26.26 y 
(25,76-27,36) 
2 
7 
qd 
4 


i 


suffused with pigment. Dark-brown islands of 
pigmentation on body. 

Larvae reached metamorphosis at stage 46 on 
13.11.1997, 39 days afler spawning. 


Discussion 


The complete larval biologies of Uperoleie 
altissima, UL inihdarea Tyler, Davies & Martin, L9st 
(Tyler ef af 1983) and Lo talpa Tyler, Davies & 
Martin, L98t (Davies & Watson 1998) are noy 
known as are tadpole morphologies of U, ryleri 
Davies & Littlejohn, 1986 (as UL maciearcuti. Watson 
& Martin 1973). Of) lithamoda Tyler, Davies & 
Martin, D98t (Davies ef al. 1986) and LL iminitla 
Davies, McDonald & Corben, L986 (Richards & 
Alford 1993). ‘These latter two species occur in or 
near geographic locations of OC. affissiima, hence a 
comparison of their salient features is of value for 
identification of tadpole assemblages. 

Uperaleia mime and U. lithamoda share a looth 
row formula of 2(2)/3 whilst that of LO eltissfra is 


172 M. DAVIES & K. R, MCDONALD 


f OUTTIMM NT 
“ é Wie 


Fiz. 7, Oral disc of Stage 37 larva of Uperofetu elfivsinu, 
Seale bar = ] mm. 


2231.2). The flexible Map supporting P3 labial 
teeth is recorded in all Upereleia to date, but is not 
unique to Uperolera (Davies & Watson 1998), The 


dark tail lip in early stages of U. mimulea recorded by 
Richards & Alford (1993), is shared by U. lithamoda 
and U. altisyima. The heavy pigmentation of later L’, 
altissima larvae may be greater than the diffuse 
pigment of U. lithomoda and U. minnula. and is a 
feature of other Uperoleia (Davies & Watson 1998). 
There are considerable differences in the length of 
the spiracle, that of U. alrissima being intermediate 
between those of U2 niimul/a and U. lithomodea. 

The unusual tail bud stage in which the tail is better 
developed than the head was noted also by Moore 
(1961) in LL feevieata Kelerstein. 1867 and Davies 
& Watson (1998) in UO) talpe. 


Acknowledgments 


This research was supported by the Department of 
Zoology, University of Adelaide and the Queensland 
Department of Environment and Heritage, We thank 
M. Tyler for critically reading the manuscript and the 
referees for their helpful comments. 


References 


Davies. M., McDonacp. K. R, & CorRen, C, (1986) The 
genus Uperoleia Gray (Anura: Leptodactyhdae) in 
Queensland, Australia. Proc, R. Soe, Vict 98, 147-188, 

& Watson. G. FE. (1998) Developmental biology 
of Uperoleia talpu Tyler, Davies & Martin. 1981 
(Antira:Myobatrachidae), Trans. Ro Soe, Ausi. 122, 153- 
157. 
. MCDONALD. K. R.. TRENERRY. M. P. & 
WERREN, G, (1993) A new species of Upernieia (Anura: 
Le ptodaetylidae: Myobatrachinae) from northetstern 
Austraha. Ment, Qld Mus. 33, 1607-174. 

Gosnek, KL. (1960) A simplified table for staging anuran 
embryos and Jarvae with nates on identification 
Herperalugien Ub, 182-190, 

Moori. J, A. (1961) The frogs of eastern New Seuth Wales, 
Bull. Am. Mus. Nat, Hist. 121, 149-386. 


Riewaros, S.J. & ALFORD, R.A. (1993) The tadpoles of 
iwo Queensland frogs (Anura: Hylidae. Myobat- 
rachidae). Mem. Qld Mus, 33, 337-340. 

SrANTUN, J.P. & Morcan, M. G. (1977) "Project RAKES 
- a rapid appraisal of key and endangered sites, Report 
No. 1: the rapid selection and appraisal of key and 


endangered sites: the Queensland case study” 
(University of New England School of Natural 


Resources, Armidale}. 

Tyrer. M, J.(1962) On the preservation of anuran tadpoles. 
Atist. J. Set, 25,222. 

Crook, G. AL & DaAvins. M- (1983) Reproductive 
~ biplogy of the frogs of the Magela Creek Systeim, 
Northern Territory. Rec. S. Amst. Mus. 18, 415440, 

Warsos, G. F. & Martin A. A. (1973) Lile history, larval 
morphology and relationships ot Australiana leprocdacty lie 
frogs, Trans. Ro Sac. 8. Att, 97. 33-45, 


CHANGES IN A MANGROVE/SAMPHIRE COMMUNITY, 
NORTH ARM CREEK, SOUTH AUSTRALIA 


By PERI S. J. COLEMAN* 


Summary 


Coleman, P. S. J. (1998) Changes in a Mangrove/Samphire Community, North Arm 
Creek, South Australia, Trans. R. Soc. S. Aust. 122(4), 173-178, 30 November, 1998. 
Use of a computer GIS package to study aerial photographs of North Arm Creek 
(1979-1993) confirmed previous studies suggesting a landward migration of the grey 
mangrove, Avicennia marina, but seaward progradation was also apparent. Samphire 
communities were reduced in area by nearly two-thirds, with the majority of the lost 
area overgrown by mangroves. At the same time samphires colonised unvegetated 
areas and some areas previously occupied by mangroves. From 1979-85 the area 
colonised by samphire was similar to the area lost, but was less from 1985-93. It is 
suggested that several factors are responsible for the changes in distribution. 

Key Words: Avicennia marina, Halosarcia, Sarcocornia, mangrove, samphire, 
saltmarsh, temporo-spatial change, progradation, colonisation. 


Pranic tions af the Rove sacieny af S. Atist (1998), 122) 


(4), 173-078. 


CHANGES IN A MANGROVE/SAMPHIRE COMMUNITY, NORTH ARM CREEK, 
SOUTH AUSTRALIA 


by Peri S. J. COLEMAN’ 


Summary 


CoLeMAm®. PS. J. (1998) Changes in a Mangrove/Samphire Community, North Arm Creek. South Australia, 
Trans. KR. Soc, 8S. Aus, 122(4), 173-178. 30 November, 1998. 

Use of a computer GIS package to study gerial photographs of North Arm Creek (1979-1993) confirmed 
previous studies suggesting a landward migration of the grey mangrove. Ayieennia marina, bu seaward 


progradation wat 


also apparent. Samphire communities were reduced tn area by nearly two-thirds. with the 


majority of the lost area overgrown hy mangroves. At the same time samphires colonised unvegetated areas aud 


sume areas previously occupicd by mangroves. From 1479-85 the 


area colonised by samphire was similir to the 


area lost, but was Jess from }985-93, It is suggested that several factors are responsible for the changes in 


distribuion, 


Key Worps: Aticcnnia marina, Halosarcia. Sarcocurnid, mangrove. samphire, saltmarsh, temporo-spatial 


change, progradation, colonisation, 


Introduction 


North Arm Creek drains from the Wingfield/Dry 
Creck area of Adelaide northwards into the 
mangrove zone of Barker Inlet (Fig.1). The zone 
comprises a seaward tringe of the grey mangrove 
Avicennia marina (Porst) Vierh. var resinifera 
(Forst) Bakh., backed by a salt marsh comprising 
nixed samphires of the genera Hulosarcia P. G. 
Wilson and Sercocentia A, J, Scott. The mangroves 
und sumphires form bands of variable width on both 
banks of the creek. 


The ereek has been used for the reception of 


stormawaters, sewage effluent and trade wastes, The 
wel coustal ecosystem edging the creck has been 
considerably modified since European settlement, In 
the fite 1800s seawall embankments were builtalonge 
the mangrove/samphire interface, and the samphire 
Aone Way used as papturage, Salt production on the 
eastern side of the creek began in 1934 and 
progressively much of the low lying area inland ot 
the seawall embankment has been ponded, On the 
western side of the creek the low lying lam behind 
the seawall became a muanicipal refuse tip, In the 
JO70S 8 series OF groynes supporting power pylons 
Was butt thioueh te taingrove/samphire zone 
ubulling the creek, 

The more recent changes have resulted in changes 
to the water flows and tidal dynamics inthe area. tn 


ella Bavironimentil Cotati, b2 Beach Reak St kil 
Aus ST) 

Hinvoriey, POL CEM ROy Haseltine Stmdy and rely: bvaloetion 
OF TCT Solt bvapergden Ponds” Bitterns Discharge on the 
Mangrove Continuity. sith Supplement Repunt provided ta (C] 
Ncatoilias Ackelatde, CU api. ), 


Sttily 
Aten 


oe Nak 
Puinp 


Torrens 
Island 
Swat 
Ale 
Lefevre Af Mier: 
Peninsula € 
( Norlt ray 
ry 


jf 
Port 


Adelaide 


North Ne wich 


Fig 1. Map ot the region. 


J986 Bradlev! recorded large seale dieback of bath 
mangroves und samphires im the yiemity ol the 
power pylon grovnes and recent field inspections 
have revealed that the area is only slowly berry 
recoloaised, 

Aerial photographs of North Arm Creck, taken 
between 1949 and 1993. show changes in the 
mangrove and samphire communities. Some changes 
are marked. such as areas of dieback. or the inland 
wdyance of mangroves. The use of Geographic 
Information Systems (GLS) technology has allowed a 
closer look af the changes in ane small region of the 


174 PS.) COLEMAN 


North Arm Creek coustal wetlind, the drainage area 
uf the Dry Creek Salttiells' No. & Pump. 

The area is hounded On the east hy wt seawall and 
om the vest by North Arin- Creek, Running ecntally 
veross the area is a small greek (hat has been formed 
by the discharge of bitterns (brine that remains after 
salt erystalligation is complete) from the saltiields 
‘The No. & Puinp and its supply drains are clearly 
visible on aerial photographs, 


Previous studies 


Burlon (1982) studied mangrove development 
north of Adelaide to the River Light using aerial 
photogriphs eovering the period from 1935-1982. 
Ve noted that the manyroave stinds showed dilfereat 
direvtions of growth at the (wo extremities af the 
study zone. Crenerally the northerly inungroves were 
prograding (extending seawards) while the southerly 
Inangeraves near Swan Alley were reureutine tlie 
aerass The samphire Mats. Burton's paper discusses 
Ihe possible causes of tis difference, i parlieukar 
discussing ferrigenbus supply and relative scu-level 
tise (eustatie rise and land subsidence). 

Durie 1985-6 Bradley! examined the mingroves 
in the vieinity of the Noo S Pump on North Arn 
Creek, He tsed Visual comparisons of aerial 
photographs of the area laken jn 1879 ard 1085, He 
peeved two trainscer fies weross the uresaued ntupped 


a Faget 
bi anus = Advpttine 


* SAUNA 


orth 143 
vn 
free 


Pinan 


f 0 all uD Lin 


Tie 2 Tie neue ey ae PP ad Od 


the distribution, heallh and age oF the veweralion 
along these, Plastie 30 cm rulers were attached to the 
transect pees to deterniine passible sedimentation 
patterns. The alignment of the substrate on the rulers 
wis recorded, 

A further study of the North Aro Creek te Swan 
Alley area was undertaken by Blackbuen® in 109d, 
He used GIS (eebniques to asvertain distribution 
changes within the mangrove and ssimphire 
communities. Blackburn? did nol physically visit the 
North Arn Creek. but the photographs he examined 
indieated both landward and seaward progradation Gl 
mangroves, 

The present study re-cxamines the area reported by 
Bradley! (Pig. 2). The review of the area combines a 
GIS analysis of vertil photographs (1949-1993) with 
4 ground survey using Bradley's! existing: transects. 
The study was constramed by growth ob Maneroves 
making access dilficull, lass of some sediment rulers 
and loss of transect pegs nearer the seaward fringe: 


Materials vod Methods 


Department of Environment and Natural Re- 
sources | 1480 seale cokugements of four aerial 
survey photographs dated 1 January 1949. 19 
March 1979, 18 February LO8S and & December 
1993 were manually dribsed tate a orn saitable fay 
use in the GIS mapping package TNTonips Lile. The 
three more recent photowraphs. duting, from 1Y79 cw 
1993, were veorcetified Using man-made structures 
onthe neihbouring sallfielvl as control ports, alone 
with Woluwted nmingrove GAL meri) lees Liat were 
ideniltiuble through the series of photographs. ‘Lhe 
1949 photogriph shows a lindscape so different 
from the present thal geurceuilication cold only be 
waccomplished by matching Ue angles of narrow 
‘barrow-pits” along the sea-wall. so vate from ir 
were Guy used ina general manner im the present 
Study, 

The prmeipal components af the mapped ares were 
dete is mangroves, samphires.ar jeither (water or 
bane mud An amilysis of the Timuts of vegetavon 
hver each OF twee periods (U7 )O85 und 185- 
YOU wos underiaken to boy blo vdewromine whit the 
dyvnuimics af the verehiuve vhange wore, 

Iieldwirk wh 1296 ineluded Winding: the Leaner 
pegs plived by Bradley’ in 1985. Vhe vexetution 
alone he Twe trynsects Was recorded and its heigl 
Weaswred Wy meres MSIE Lape 
Keudines were alse rewonled where sediment mifeds 
were still in puree 


Het oa 


Brarnues, Bo ciel) Mopiige Mayepase yin Salrenarss) 
Commmoties Sear ee Bleeets Digelwwe Area kintall 
Poti Atldale Report te Peirce Seen Mroelitere CE a piber 


CHANGES IN A MANGROVE / SAMPHIRE COMMUNITY 7 


Typur |. General cheanves in the vegetation. 1979-1093, 


Mar 79 
Mangroves (17) 73902 
Saniphire (ns 14173 
Vevehtlive cover (m2) RSOTS 


a 


beb—-$S Dee-"93 Change, *79-"93 
TROOA N8459 + 14557 
12769 5340 ~ $833 
91377 93799 +5774 


Tash 2. Change rates ef the prineipal campenents af ihe vegetation. 


Area of etch chine ehiss 
of the yezetalion (a) 


Annnal rate of change 


1979-1985 


Mangrove to niangrove (6 change) 69175 
Saimphive te iminennve 202 
Mamnurove to samphire 2913 
Neither (o iuingrove 35) 
Mingreve to neither” 124) 
Neither’ (9 sumphire ARTO 
Siimphire to neither! 1924 
Saniphive jn samplire (ao change) S004 
Veretative loss 3163 
Vegetative dhercase 7396 


Rale of change in loti vegetative cover 


1945-1993 


TO7OTU8S IUSS. 1993 


THY74 - - 
GOUS a4 762 
20 46 7s 
3758 587 47th 
ahh 207 74 
W163 646 145 
AOR 32| 400 

2051) - 
4576 527 a72 
442] i254 613 


706 43 


* “neither” indicines areas of bare mud or water 


Because the A, mwrine trees had grown 
considerably during the TL years since the transect 
pegs were pliced, locating the pegs past the 80 mH 
mark on Transeet A and the 110m mark on Transeet 
Bomight possibly fave resulled in damiage to the 
mangal and so no dala were collected beyond these 
ports, 


Results 


Vevetarton imeppinge 

A comparison for the period 1979-1093) shows 
extension Of the mangrove canopy, and a reduction in 
{he aren covered by samphires. with an overall 
inerease in vegetated area (Table 1). As the study 
zone is delimited on the landward side by the seawall 
embankment, the gain must either be the 
overzrowing of previously bare mud patches, or 
some seaward uccession, 


The results of the analysis of the limits of 


vegelation Tram 7979-1985 and 1985-1993) are 
summarised in Table 2. 

The latwest change over the period was a jnerease 
of mangroves at the expense of the samphire 
community. tlowever, the extension of both 
mangroves and samphires over bare mud and into 
water areas ts alse oecurring, along With samphire 
colonisation ol arcus previously supporting 


mangroves. Lirosion is oecurring in some wreas of 


samphire, 
Samphire has given svay lo mangrove at their 
interface as the mangrove has. extended inland. 


Almost the entire central samphire zone has been 
suceeeded by mangroves and the trees have also 
oecupied many of the creck lines as well as 
colonising the bare mud dreas along the seawall 
embankment. Site visits ia duly and September |996 
revealed juvenile mangroves growing along the No. 
8 Pump discharge channel and specimens more that 
4in high growing along the seawall within 40m of 
the discharge pout 

Some mangrove areas have been replaced by 
sumphire or by bare mud. This has mainly occurred 
in the southern part of the study area hut alse along 
the bitterns discharge creck 

Along the seaward edge, progmiadation ol 
mangroves is apparent along the entire length of the 
study zone. The extension is most marked i the 
southern areas, with a maximum advance of 
approxinnitely 25-30 min the 14 years from 1979- 
1993, 

In the northern purl of the study zone the seaward 
progression consisted mainly of infilling the many 
invaginations and embayments around isolated trees 
and the advance was between 10 and 15 m. The 
cause of the slower progradation ol mangroves in the 
northern urea is uncertain but the milling of semi- 
enclosed areas suggests that low water flow rates in 
the sheltered areas were conducive to sediment 
wecretion, whereas the actual seaward fringe may 
have been exposed lo stronger wave action, 

The samphire community has also been extending, 
and has beeome established on previously bere mud: 
there ure now samphires along the seawall within [Sm 


176 


Taunt. 3. Sedimenrarion recdings along the trdasects. 


P.S.d, COLEMAN 


Distance A Transect GB Transeet 
along 
transect 
1985 1986 reading 1996 reading, 1985 1986 reading 1996 readin 
initial (change) (chanpe) initial (change) (chanpe) 
redding readin 
in Ts 63 nit 5.2 63 nit 
( tdbem) (-1) em) 
Hin bf K 3 45 46 7 
(-lL4em) (r5.0 em) (-U,1 cnt) (2.4 om) 
Aan 43 52 4 4 34 ww 
(-0.9 em) (44.2 em) (+0.2 em) 
50 m 8 7S ta 7 62 ie 
(40.5 cm) (+0.8 em) 
oom 6.1 6.1 Nib 63 63 3.5 
(no change) (no chanve) (+408 om) 


Note 1: 1985 & 1986 readings from Bradley (|986)!, 


Note 2: Readings are the alignment of the substrate agaist 30 ch plastic rulers attached to the transect pegs. Zero is to the 


lop af the ruler, 


of the discharge point As the creeks are being 
iitilicated by miatueroves, new areas for samphire 
colonisation have appeared. Much of the new growth 
is along the seawall and to the north of the study 
urea, Where the ETSA eroyne his altered the tidal 
circulation. 

The bitterns discharge does not appear to have 
affected growth Of samphires negatively. possibly 
because (he species are adapted to surviving in high 
salinity regimes, but samphire has been eroded away 
in some areas along the bitterns discharge creck. 


Treisects 

Figure 3 presents the 1985 ind 1996 heights of the 
vegetation along Bradley's! existing transects and 
shows the muturation of young stands of mangroves 
and the new colonisation (by mangroves and 
sumphire) of areas closer to the discharge point. The 
1996 date were collected along the transects in 
September, The forests are now so dense that 
accessing the pegs is difficull and so the iranseets do 
pat continue fo the original (50m point, 


Sedimentiaion 

The bitterns discharge creek has formed since the 
1949 geri! photograph was taken and Bradley's! 
report capressed same coneerh that erosion might be 
oecurring in this creek near the discharge point, He 
examined sedimentation patlerns away lrom the 
iminediate disebarge point by attaching plastic rulers 
to the transect pons and recording the celative hehe 
Of (he substrate at each location, 

In response to Bradley's! finding during the initial 
Observation period (1985-1986) that some erosion 
wis oveuriie lea’ the discharge point, saltfield 


personnel deposited concrete blocks in the drain to 
break up the flow. To determine the types of change 
that might have been occurring since 1985/86, the 
rulers were examined ip September |996 where they 
sull existed. The few remiining sediment rulers 
indicated that the hydrology of the area may have 
changed. These 1996 readings are presented (Table 
3) together with Bradley's! L985 and 1986 readings. 

The southern transect (Transect A) shows 
deposition to have occurred within tO m of and 
possibly closer to, the discharge point. The 
lopography of the transect is smooth, with no creeks 
crossing It, so sedimentation may be relatively 
uniform across the urea, 

The northern transect has several small evecks 
crossing il, and the Crosion/sedimentition pattern is 
more complex. The lack of rulers makes iCdifficult to 
Interpret, However, the urea closest to the discharge 
point has eroded somewhat over the list LO) yours, 
forming a creek tine, Al low tide any clischarse 
follows the existing creeks (slightly to. the north 
before turning westerly), which have become more 
detined as mangroves have colonised the Mats around 
them. The creek at 60 Mm is not recorded as having it 
sediment ruler on Bradley's! original sediment lable, 
but a reading Of the topographic plan of the transect 
done ih 1985 shows the creek to be about 15 em 
deep; the current reading is 14 em, The ruler an the 
JOO m Wansect peg in the main forested area along 
the northern transect shows a small sediment sain. 


Discussion 


The detailed GIS study was possible beeause 
sufficient Markers were visible iN veri photog raplis 


CHANGES LN A MANGROVE / SAMPHIRE COMMUNITY 


Distribution along Transect A 


1985 


Height of vegetation (mm) 


177 


60 70) 80 90 100 110 


0) 100 Ho 


10 20 30 40 Ss) 
A Distance along transect (m) 
Distribution along Transect B 
19&S 
4 1996 0 --2ee->- 

u 

x 2 

= | 

B Distance long tumseet (my 


Fig 3A. Heteht distribution of vegetation along Transect A” B. Height distribution of vegetation along Transect Bs, 


to allow precise yeorectification. The 1949 
photograph lacked some markers, reducing 


confidence in the precision of its georectification, 
However, this earlier photograph provides some 
insight into changes in vegetation patterns. The main 
differences include: 

1. a larger area of vegetation in 1949 benween 
the seawall and North Arm Creek: mangroyes 
extended further out into the creek, 
no creek in the location of the current bitterns 
discharge creek and the land inside the 
seawall was grazing land. 

a wide expanse of samphires, with 

mangroves penetrating trom North Arm 

Creek in) towards the seawall along 

depressions, and 

4. areas of mangrove dicback just behind the 
seuward [ringe of mangroves. 


i] 


to 


The 1949 photograph showed that the mangroves 
in North Arm Creek were already retreating inland, 
so the seaward expansion visible in the post-1979 
photographs must have started before 1979 but atter 
L949, 

It is postulated that there has been an advance and 
regression of the mangroves with relatively small 
changes in water flow patterns. According to 
Hodgson et al. (1966) North Arm Creek received 
the flow of effluent trom the Islington Sewage Farm 
from 1881 through to the opening of the Bolivar 
Sewage Treatment Works in the 1960s. 

During the operation of the sewage farm, nutrient 
rich water would have been released into North Arm 
Creek, The effluent may have supported algal 
blooms that could have caused the sporadic oxygen 
depletion in the waters of the creek recorded by 
Hodgson (1959)* . Induced anaerobic conditions are 


178 P. S.J. COLEMAN 


reported to cause the asphyxiation death of areas of 
mangroves (Diop et al. 1997) and this may explain 
the areas of di¢back visible in the 1949 photograph, 

The changes in the mangrove/samphire com- 
munities vistble in the 1979-1993 photographs 
confirm previous studies that have sugvested that a 
landward migration of A. marina is occurring in the 
southern reaches of Barker Inlet resulting in a 
reduction of the area of samphires, However 
mangroves are also prograding seawards and 
covering a larger area, suggesting that the growth and 
distribution pattern is not a response Lo a single factor. 

While land subsidence/sea-level rise (Burton 1982) 
may be responsible for the landward progradation, it 
cannot account for simultaneous seaward 
progradation. Sedimentation readmgs from. the 
transect rulers indicate that sediment is accumulating 
over much of the area and that any lowering of the 
Jand surface is likely to be a widespread Jandform 
settlement (PPK 19925) of the sedimentary coastal 


deposits rather than a lack of sediment supply or 
erosion per se, except in specific areas such as creek 
lines and patches of mangrove dicback. 

Samphire communities over the period 1979-1993 
were reduced in area by nearly */; despite the overall 
gain in vegetated area. Most of the lost area was 
overgrown by mangroves, However, the direction of 
change was not entirely one way, as samphires 
colonised areas previously occupied by mangroves 
plus areas of mud/water. During the carlicr period, 
between 1979 and 1985, the urea of new samphire 
growth cach year nearly matched the area lost. so 
that there was an apparent loss of only 200 m? of 
samphire annually. The later period (1985-1993) 
showed a slowing im newly colontsed areas of 
samphire. Although the area overgrown by 
mangrove or eroded each year remained about the 
same asin the earlier period, the rate of loss appeared 
higher (LO0O nv annually) because there was Itttle 
colonisation of new areas by samphire 


References 


Burpon. T. E. (1982) Mangrove Development North of 
Adelaide. 1935-1982. Trans. Ro Soo. S. Aust. 6, 183- 
TRY, 


Hopason, HL J. N., Lewis, RK. Wo. MILes, K. Ro, Jupp, Po& 
GILCHRIST, J, W. (1966) Report of the Committee of Hnguiry ite 
the Udlisation of Bffluent from Bolivar Sewage Treatment 
Works. Government Printer, Adelaide. (Unpub. ). 

' Honcson, H, JN. (1959) ‘Treatment and Disposal of the Sewage 
of the Adelitde and Salisbury-Klizabeth-Gawler Drainage Areas. 
Engineering and Water Supply Department, Adelaide. (Unpub.), 
PPR Consultants (1992) MFP Australia Gillman/Dry Creek 
Urban Development Proposal-Drafi Environmental tmpaet 
Statement prepared for the Premier of South Australia, Adelaide. 
(Unpub.} 


Diop, E. S.. SoumARE, A., DIALLO, N. & Guise, A. (1997) 
Recent Changes of the Mangroves of the Saloum River 
Estuary. Senegal. Mangroves and Salt Marshes 1, 163- 
172, 


NEW SPECIES OF SEURECHINA (NEMATODA: SEURATIDAE) 
PARASITIC IN DASYURID MARSUPIALS FROM AUSTRALIA 


By L. R. SMALES* 


Summary 


Smales, L. R. (1998) New species of Seurechina (Nematoda: Seuratidae) parasitic in 
dasyurid marsupials from Australia. Trans. R. Soc. S. Aust. 122(4), 179-184, 30 
November, 1998. 

Seurechina hobbsi sp. nov. is described from the stomach of Phascogale tapoatafa 
from Western Australia. It differs from S. chaneeti, the type and only described 
species, in being a larger worm (6-8 mm compared with 3.1-3.8 mm) with longer 
spicules (500-630 wm) for S. hobbsi compared with 185 ym for S. chaneeti. 
Seurechina spratti sp. noy. 1s described from the stomach and small intestine of 
Sminthopsis leucopus and Antechinus agilis and is most closely related to S. hobbsi 
from which it differs in having three lateral papillae extending into the caudal alae 
rather than two, oval rather than spherical eggs and the absence of a large projecting 
lip anterior to the vulva. 

Key Words: Seurechina, nematodes, Seuratidae, Echinonematinae, Australia, 
Dasyuridae, marsupials. 


Tronsaetions afte Ravel Societhy af S Aust (1998), T2204). 179-184. 


NEW SPECIES OF SEURECIIINA (NEMATODA: SEURATIDAF) 
PARASITIC IN DASYURID MARSUPIALS FROM AUSTRALIA 


by L. RL SMALES 


Sunumary 


Sviapes. LR. 1998) New species of Searveting (Nemiuoda 2 Scuratidie) parasitic ieckisyuridl iarsupiils lean 
Australia, Teaiis Be Soc Ss Alist E2204), 179-18, 30 November, 1998, 

Semrechine hebbsi op. WoW is described from the stomach ot Phiscogale tapoanife tron Western Australi. I 
differs Trou. ehaneer. (he (pe wid oAly deserbed species, i berg a larger worm (6-8 in compared: with 
3.1 3.8m) with longer sprees (500-630 pind for §. frefbat compared with TSS pm lor S cfemeed, Searectiia 
yyrrotté sp. nay, is described [rom the stomach and small intestine OF Saintiopaty dedeopis and Antechinesy agilis 
and is mast closely related (oS. frabAsd from whiek it differs in having three fateral papillae extending inte the 
caudal alae rather thon two, oval father (han spherival eves and the ubsence of a hinge projecting Hip anterion to 


the valyva 


Kry Wokbs: Seweniiaa. nematodes, Seuratidac. Eehinonematinge, Australi, Dasyuridae, marsupials, 


Introduction 


Nemmlodes of the furnily Seuratidie are parasites 
oF reptiles, birds, bats. redents and Australian 
marsuphils (Chabaud }978) The fumily inelides 
venerd in Which the mouth is dorsu-ventrally 
elongated and fimked by paired lips and genera in 
which (he mouth opening ts triingulir or hexaeonal 
(Inglis 1967) All three genera oecurring in 
Australian marsupiils. Sewreehina. Pradechine and 
Linstewitwned spp. are contained in the subfamily 
hehinonematinue Inelis, 1967, characterised by a 
lurve mouth opening with no lip lobes, the anterior 
end of the body being swollen as a cephalic bulb 
hearing hooks, no pre-cloucal sucker on the mile and 
cloacal region covered by cuticular granulitions 
Although originally placed in the Sehnerder- 
nematidae by lntlis (1967) the affinities of 
Linviowinema Smiues, 1997 (lormerly Behinonema 
Linstow. 1898 preaecupied) with the larvae ol a 
species of Senratiu Hall, (976 resulted in Quentin 
(1971) placing the Eehinonermiatinue in the 
Seuratidie, 

The genera Linstvwdnemea and lngleettineg 
Chabaud, Seupeau. Beveridge. Bain & Durette- 
Desset. 1980, vont species with a triangular 
mouth Opening on a swollen cephalic bulb beariny 
hooks. The monatypie genus Sevreching Chabuud, 
Seureal. Beveridge, Bain & Durette-Desset, [980 
however, has neither a tritngelar mouth opening nor 
a swollen cephalic bulb bearing hooks, although il 
does have other characteristics of the subfamily. 

In this paper, Iwo new species of Sevreciine are 


' School of Biolowieal ane havevnmenkd Sefenees, Ceara 
Quesmsbindl Criversily Rocs himpton Ohl 4702. 


described, The delinition of the sublinmily Hehi- 
Honematinae js re-evaluated and the yelationships 
belween the venera discussed, 


Materials and Methods 


Nematodes collected from Sriiitliapsin dencapuy 
ad Anlechinus auilis were fixed ta bot LO%e formalin 
and then stored in 70. ethanol The preservation 
history of the specimens from Phetsireete wepoctefit 
is unknown although they were stared i 7% 
ethanol. All hematades were examined after clearing 
in lactophenol. Measurements for more than tour 
specimens are given in micrometres, as the range 
followed by the mean in parentheses, and were made 
with the aid of an ocular micrometer or drawing tube 
ahd bap Measorer, Drawings were made with the aid 
Ob ad drawing tube. 

Type muterial has been deposited in the South 
Australian Museum. Adekude (SAMA) wid voucher 
specimens are held in the collection of CSIRO 
Wildlife und Ecology (CSIRO), 


Seurechina habbsi sp. nos. 
(FIGS 1-10) 
Types? Hulotype cd. allolype 2. paratypes 4 dd. 17 
oS) from stomach of Plawecavale tapoutafa (Meyer 
1793), Manjimup (34° 15'S. 16) 09" LE) WA, June 
1992, coll, S, Rhind, SAMA AHC 31262, ATIC 
31263 und ATIC 31264, respectively, 


Minerial examined: Prom Phiscogale tapectafea 
fypes. 


180 


L. Ro. SMALES 


Figs 1-10. Seureehine hobbsr sp. noy. 1, Anterior end optical section. arrow indicating laminae Uateral view). 2, Cephalic 
end, optical section (lateral view), 3. Cephalic end, optical section (dorsal view). 4. Cephalic ene, arrows indicating 
laminae (ateral view). 5. Cephalic end (en fuce view) 6. Male posterior end (ventral view). 7. Body spines. a. From 
oesophageal region. b. From mid body region. c¢. From posterior body region, 8 Male posterior end (lateral view), 9, 


Female posterior end (lateral view). LO. Vagiia (lateral view). Seale bars = 200 um ts 10d Ss 100 pov 6.8.9 FO: 50 ui 
7:25 pum 2. 3, 4 


NEW NEMATODES FROM DASYURID MARSUPIALS 181 


Deseripnon 

Small worms, body with tine transverse cutie kur 
unnulations, Cephalic extremity without spines, 
remainder of body with up to 50 rows of spines (nid 
body of female) at cach annulation: extending over 
Ye ohody dorsally to caudal ake ventrally of male. 
over entire body of female: spines beeommnig 
progressively smaller towards posterior end, 
Anterior extremity wilh mouth opening and oral 
cumily, elonsted dorse-veoually. bearine 2) pairs 
double cephahe papillae, pair aniphids: without lips 
or lip-like structures, |Sntertar end af vesophagus 
cupped by 2 oval, dorso-ventrally aligned sclerotised 
tings, enlamwed dorsally and ventrally. Qesophaztis 
surrounded at untertar end by 4 pairs hunminue &O 
lon, Oesophigus simple chivilorm oh. body 
lenwth, Nerve fing aid seeretory-exeretory pore net 
sce, deirids linge. eonival. al level-al Sth row oF 
spines. 


Mite (i=5 unless olheryine shite) (bigs 1-8) 

Length 0-70 (0.5 mim), width 300-370 (440), 
Ocsuphugus 502-569 (536) Jong, Deirtidls 77-94 (85) 
from anterior end. Spieules equal sine. without 
whe, 5006630 (590) lone, about i) body Jength. 
Gubermaculum 50 (n=1) long. Tyo pies lateral pre 
wloavcul papillag. | pair extending inte lateral alae: | 
pay lateral ud-cloicul papillie extending inte lateral 
alice, 3 pais postelotcul papillae, | pair phasnuds 
well posterior to Glaaua near tail tip, Pettelouwal 
pupillae not seerr “Tail | 30-170 (150) long. 


Fenle (v=35 unless otherwise stated) (Pigs 9, 10) 

Length ZUR U8 vin), width SIO-3580 (550). 
Oesophiwus 536-670 (910) long, Deiids nov seen, 
Vaying 550 (n=l) long: vulva opening behind a large 
projecting Lip, 2600-3450 (4000) from anterior end. 
Monoudelplig, (gil 215-280 (255) long. Fags spher- 
ieal 1) 54 (47) diameter. 


Livniales! 
The species is named aller Mr R. Hobbs who his 
been Helpful iy providing material for.this work 


Kemiarks 


The method ot fixation used Tor this materith wats 
not ideal most specimens being contracted and 
distorted, [Was impossible to determine the number 
and arrangement of the peri-cloueal papillie on mule 
speciinets but lateral pre- ad- and) post-cloucal 
papillae could be seen, Their number and artange- 
ment are similar to those of the bype and only other 
species, SL avaneen. Semvehind Hobbst. 6-8 nan 
Jone. is at kirwer worm thin dS. efazeen. 3.13.8 pin. 
With longer Spicules (500.630 in 5. febhyt conipured 
with 185 in & eheneety. The posterior ventral body: 


spines cover the entire ventral body surliee of mate 
§. hobby whereas those of & chuneetd terminate in 
two dateral bands (Chubuud ef af 1980, Fig. Lp p 
430), In S Nehbse the lemale tail (215-280) is longer 
than that ofS. efaneeté (120), the spherical cues are 
larger (47 diameter compared with 40x35), the vulva 
iS pre-equaturiitl compared with a post-equatorial 
vulva in So hancert. Sencechina habhsi 1% 
monodelphic, whereas 4. chareet/ is didelphic, 


Seurechina sprattt sp. voy 
(FIGS 11-19) 


Types: Voloype 2. atlolype So trom: stomach) of 
Sminthopsis leucopus (Gray. (842), Sidlings Swamp 
South, Vimbillica State Purest (47° TBS, 149° 45' Ty, 
NSW, 25.483, coll. P Hiyeork, SAMA AHC 31265 
und AFIC 31266, respectively. 


Meterial evanied: Prom Snitnihepsis lencapuis: 
NSW, tpes: From stomach Aavtechinus cagilis 
Dickman, Parraby. Crowrher & Killa. (998: 20 4 
7 Y 2. Sidhinus Swamp North, Timbillrea State 
Forest. NSW) Law 87. S188, coll PO Haycock 
and EB. L. Walter, CSIRO N2841, N2977, 


Dese riplion 

Simall worms, body with fine transverse cuticular 
annulitions. Cephahe extremity without spines. 
reminder ot body with up to 46 rows spines (mid 
body of fenale) an eaeh aipntilation, extending Over 
YY hody dorsally ta caudal akre ventnlly, of mle, 
over enti body of female: spines becoming 
progressively Soiuller towards posterior end, 
Ajiterioy extvemity with mouth opening and oral 
cavity cloned dorsa-ventrally, hearin 2) pains 
double sub-median cephalic papillae. pair lateral 
aniphids: without tips or lip-like structures. Anperiot 
cud Of oesophagus capped hy 2 oval derso-ventrally 
aligned selerotised rings. enlarsed dorsally vind 
ventrally, Ousophaus surrounded al anlerior end by 
4 pairs laminae 110-165 long, Ovsophagus simple: 
chayv-ifort, Yee '/je body length. nerve rin aileron 
lo. detrids. deirids Linge, Comma aitlevel of about ath- 
7th row of spines: Seerefory-exerelary pore nol seen, 


Mole Oneasiwenients of falotype falliwed by 
measurements of 2 cde fron A. wu7fis) (Plas Fh 12, 
14.17. 18) 

Length 44. 45-6 pimsy width 220. 270-340, 
Oesuphagus 470. 355-470 long. Nerve ving #3: 
derrids VEO, RO-T10) fromt antetior end. Spicules 
equal, sitar without ale, SOQ. 350-000) long, 
about My body length. Gubernaculiunt fot seen i 
holotype, SO-38 in specimens trom A. again. Two 
pairs of hiteral pre-cloacal papillae: | pili extending 
mtg fateral alae, 2 paws lateral ac-cloacal papillae 


182 LR, SMALES 


AS 
a) 


han 
he 
Me. 


Figs }1-19. Searechina sprati sp. noy. 11. Antertor ead, optical section (lateral view), 12. Cephalic end. optical section 
(lateral view). 13. Anterior end (en face view). 14. Body spines, a. From oesophageal region. b. From mid body region. 
15, Cephalic end, optical section (dorsal view). 16, Cephalic end (dorsal view). 17, Male posterior end (lateral view). 18. 
Male posterior end (ventral view), 19, Vagina (lateral view). Seale bars = 100 pm Th. 19: 50 pm 12, 17, 18; 25 pin 14, 
15.16: 10 pm 13. 


NEW NEMATODES FROM DASYURID MARSUMALS 183 


extending lhto Literal ahie, 3) pairs peri-cloucal 
pupiliie; 3 pairs post-cloweal papillae, 1 pair 
phasmids well posterior to cloaca near tail tip. 
Anterior lip oF eloaea with swallem taterul edyes 
Simulating pair of supplementary papillae. Tail | Ab), 
{O5-200 long, 


hemale (neastiienients of allitype tollawed 
measurements Ob 7 Ly from A, audi) ies ba. 1 
16, (9) 

Lenath 6.6, 6.07.0 nit (6.9), with 470, 300-630 
00). Oesophugus G00. 380-570 (485 lomo. Nerve 
runt, Secretory-excretoey pore, demds not seen, 
Varia 340, 450 (n=1) long, Vulva 2950, 2975-3485 
(3150) trom anterior ead, Monydelphie, Tail 240, 
1-240 (2501) hone. Pugs oval 34-53 147) lone by 
23-33-(27) wide. 


ny 
3, 


Eivinelosy 

The species is named after Dr BoM, Spratt in 
revo Tion af ls Conte bathed Lo Gur understinding 
of the hehotinths of dasyurids. 


Renirks 


The scurclory-cxereiory pore. oay un coneedled 
between body spmws close fa the anterior end. is 
vilen dieult lo defeet in echionernatines. Ta lis 
species, The anterioremls of all worms from A, aerliy 
were Contucred, tod ereater or lesser extent during 
Futon, obscuring the seeretory-exerelory pore. 
This may have occurred beeduse the heads. of the 
worms were embedded ii ihe mucosa al post- 
mortem cxamimnaion (DO Mo Spratt pers. comm. 
}O98) Meustinanents of oesuphagesl length were 
Als uiffeeted by the state OF Fixation, those ol 
lermiles from A. aailis bere apparently slterter dati 
that of the female Prom S$. feneepius, Other 
measurements oF specimens Trontthe two hosts were 
consistent Wilh their belonging tow single species, 

Sevechind sprit most elosely resembles 


hohbsi a size, length oF spieates, distribution of 


body spines, posifion of yvulyi. bene monodelphie 
and the length of the tail in both the mate and 
female. AU of these characters distinguish both 8. 
habbsr and So osprane foin S. cheaneen. Searee hina 
spradé can be veadily distinguished from both 
hobhsi ind So chaiect® in having three rither than 
two fare dateral papillie extending (ito the caudal 
lace, Searechina sprend lias oval egas whereas those 
oF S. tabby: ate spherical. Senreefine spear lacks 
Hie lunge projecting lip anterior to the vulva Pound in 
&. hobbse, 

Semechinag chinteecn wis described front eaves 
allneatis. Gould, {842 from Koolan [shind, off the 
vost of fort Westerns Australia, 8. bobs? fron P 
fapodtate (Meyer 1793) front thre southert 


miintand of Western Anstralia and §. spre tram S- 
leucopus (Gray, L842) ahd AL eediy Dickman, 
Parnaby, Crowther & Ring. (998 fron southeastern 
New South Wales fear the Victorian border ‘The 
differences between S$. Chance and the other two 
species may be explained, at lease partly. by 
acngraphic separalion. The similarities between 4, 
hobhsi and So spreui would be the resale ot a 
common ancestor in coastal Vieloria and/or Nesw 
South Wales. where the ranges of the three host 


species overlap (Sumner & Dictkmun T4998; 
Soderquist 1993), 

Discussion 
Inglis (1967) created) the subfamily leh 


Hanematinae to wccommodate the genus Aeliauend 
(se). and pliced it within the Sehnercernenttiduc. 
niher than the Seurattdite because OF its long sinple 
spicules, short gubernaculuiri und a Lriradiate mouth 
opening without fips. The aftiity of the Beli 
nonemiulinae with the Seuratidie was diseavered by 
Quentin (1971) and confirmed by Chabaud. cn al 
(1980), They linked the presence of a simple, lip 
less. triraciate mouth opening. two pairs of doubled, 
subiediah eeplalic papillae. a very shallow 
vheilostome. and chanacteristie spines an the body 
culicle oF adults will the Seuralidae, sie ith) harwal 
development in Seapatua sp. 

When the genus Senpechiia wis erected by 
Chabaud ef a/. (1980), these authors diseussed tts 
lack of Gephalie hooks but placed it in the Eehi 
nonematinie wlones with the genera, Lonalondrena 
(formerly Achinonene prenee.) and dieleehine. 
Which also ocour ih dasyurid hosts. hey did not 
comment upon the fuct that Senreching has an oval 
mouth openiny wor upon the signitieatwe of the 
dorso-ventrally elongated culeahir structure 
behveen the oesophagus and the mouth opening AL 
present this chitinaus cup. the walls of which are 
made up of bwo Superimposed rites (Chabaud ef ef. 
JOSO). ds NOL Tterpreted as part ala cheiostonre ws 
defined by Inglis (1967) bul rather as assoeialed with 
un pesophastome, The four pairs of sublate rid 
laminae found in the cephalic hypodernis wene 
howe by Chabaud er af (1880) us appearing To be 
dilations of the hte! Telds. possibly playing a 
roke tk mechanism ty hold the eervien! Spies 
steady when they are embedded jn the gastric 
mucosa, These structures hive not been found in 
other echinonemiuatines (Chabaud er uf TY80; States 
LOOT), 

For the time being it is eanveniont To renin 
Seureehine within the Eehinoaematinae until the 
developmental relationships of the mouth, oese- 
phygus and assoviated structures have been 
detenmined, The senus could ether be moved ta 


184 L. R. SMALES 


the Seuratinae on the basis of a bilaterally 
symmetrical mouth opening, necessitating 
emendation of the diagnosis of the Echino- 
nematinae to accommodate adult worms with an 
oval mouth opening, or, alternatively new 
groupings could be established. 


Acknowledgments 


] am grateful to R. Hobbs and D. M. Spratt who 
made available the specimens for this study, and I. 
Beveridge who criticised an early draft of the 
manuscript. 


References 


CHABAUD, A. G. (1978) Keys to genera of the 
Superfamilies Cosmocercoidea, Seuratoidea, 
Heterakoidea and Subuluroidea No. 6 pp. 1-71 Jn 
Anderson, R. C., Chabaud, A. G. & Willmott, 5. (Eds) 
“Keys to the nematode parasites of vertebrates” (CAB 
International, Farnham Royal), 

. SEUREAU, C., BEVERIDGE, L., BAIN, O. & DURETTE- 
Desset, M.-C. (1980) Sur les Nematodes Echino- 
nematinae. Ann. Parasitol. hum. comp. 55, 427-443. 

INGLIS. W. G, (1967) The relationships of the nematode 
superfamily Seuratoidea. J. Helminthol. 41, 115-136. 

Quentin, J.C. (1971) Sur le cycle évolutif de Seuratian 
cadarachense Desportes, 1947 et ses affinités avec ceux 


des Nematodes Subulures (Ascaridia) et Rictulaires 
(Spirurida), Ann. Parasitol. hum. comp, 45, 605-628. 

SMALLS, L. R. (1997) A revision of the Echinonematinae 
(Nematoda : Seuratidae) from bandicoots (Marsupialia : 
Peramelidae). Trans. R. Soc, S. Aust. 121, 1-27. 

Soperouist, T. (1995) Brush-tailed Phascogale: Pliuss- 
cogale tapoatafa (Meyer, 1799) pp. 104-106 Jn Strahan, 
R. (Ed.) “The Mammals of Australia” (Reed Books. 
Chatswood). 

SUMNER, J. & DICKMAN, C. R. (1998) Distribution and 
identity of species in the Antechinus stuartii ~ A, flavipes 
group (Marsupialia: Dasyuridae) in southeastern 
Australia. Aust. J, Zool. 46, 27-41. 


SPIROXYS CHELODINAE BERRY, 1985 (NEMATODA: 
SPIRUROIDEA) AND CAMALLANUS CHELONIUS BAKER, 
1983 (NEMATODA: CAMALLANOIDEA) FROM FRESHWATER 
TURTLES (PLEURODIRA: CHELIDAE) IN 
QUEENSLAND, AUSTRALIA 


By MERYL A. FERGUSON*® & LESLEY R. SMALES* 


Summary 


Ferguson, M. A. & Smales, L. R. (1998) Spiroxys chelodinae Berry, 1985 (Nematoda: 
Spiruroidea) and Camallanus chelonius Baker, 1983 (Nematoda: Camallanoidea) 
from freshwater turtles (Pleurodira: Chelidae) in Queensland, Australia. Trans. R. 
Soc. S. Aust. 122(4), 185-189, 30 November, 1998. 

Spiroxys chelodinae was found in 22 of 77 Emydura krefftii, in three of six areas 
sampled. This is a new host record. Camallanus chelonius, previously reported only as 
occurring in the pleurodiran turtle Pelusios sinuatus from South Africa, was found in 
seven of eight Elseya latisternum, 19 of 77 Emydura kreffti1 and one of one Chelodina 
expansa from five of six areas sampled, new host and locality records. This finding 
provides a link between pleurodiran turtles on three continents. 

Key Words. Spiroxys, Camallanus, nematode, freshwater turtles, Pleurodira, 
Australia. 


Treecions of te Royal Seciely af S Navi (1998). T2204), PKS E84, 


SPIROXYS CHELODINAE BERRY, 1985 (NEMATODA;: SPIRUROIDEA) AND 
CAMALLANUS CHELONIUS BAKER, 1983 (NEMATODA; CAMALLANOIDEA) 
FROM FRESHWATER TURTLES (PLEURODIRA: CHELIDAE) IN 
QUEENSLAND, AUSTRALIA, 


by Meryl A. PERGUSON' & Lestey R. SMALES' 


Summary. 


Percuson, MA & Smares. (Re (1998) Spica ehelodinve Berry. }Y8S (Nematoda: Spiruroideay and 
Camallanis chelonins Baker, (983 (Nematoda: Camullanoided) fron freshwater turtles (Pleurodira Chelidae) 
it Queensland, Australia Tas. A bee Ss. Ati. 122(4), 185-189. 30 November. 1998. 

Spiraxyy cheloedinne was found inn 22 of 77 Enveira Reefs, in three OF Sis areas sampled. This is a new host 
record, Camadlanus chelonius, previously reported only us oceurring mi the pleurodiran torte Pedisies sinus 
from South Africa, wits found in seven of ent Elsevier dadiscernmmn Lol 77 Leeyedire hrefffit and one of one 
Chelodina expanse trom five af six areas sampled. new hoseand tneality records. This finding provides a link 


benween pleuradinan firtles on three continents, 


Kie Wonis: Spires, Canrrlfanes, nematode, [reshiwater tirtles. Plearodico, Auatiaha 


Introduction 


Two miuyor routes af orgins for the nemtode 
parasites of reptiles aod amphibians have been 
suuuested (Baker lO84), Groups with sporadic 
representatives in ainiphibians and reptiles are 
considered to tave beer captured from invertebrates 
or other vertebrates. The majority of nematodes, 
however, Fave evolved wilh their amphibian and 
repuilian hosts, 

The spirurid family Camatlinidae ts an example of 
the Hest mode of origi This family is suggested to 
have evolved in fish in tropical Asia. the region with 
the wreatest diversity and riehest eamallanid: hiune 
(Strombery & Crites 1974), Boecal morphology 
suigwests Hal He subdamily Camallaninae, including 
he venus Cimallanuy Railliel & Henry, 1915S is the 
tnost reeently evalved ane this is supported by the 
fier that 43% of Camella species oceur in turtles, 
frogs ane smukes (Strombery & Crites 1974). In 
Australis there are flo Species known trom 
freshwater fish ar turtles, although Serpinenn 
aclorugatas (Baylis, 1933) Petter, 1979 has been 
reportest front a eryptodiran turtle. Heavens 
uranix, in Malaysia (Baylis 1933). 

The-spirurid family Gnathostomidae, including the 
monovenerie subfamily Spiros ine. is an example ar 
the second mode of origin, Le, evelving primarily in 
imphibians and reptiles (Baker 1984), The genus 
Spiros Schneider, 1866 probably originated in the 


' Sehool ooh Bivlogieut vind Choveeenenil Senices, Cernig 
Oueenslind Ciniyersity Rock lmplon Olt 4702, 


holarctic or oriental regions us perssites of non 
Inaring cheloniins of the Suborder Cry ptodira (Berry 
LOSS), Spirovys chelodinde Berry, 1985 was first 
recovered from Cheledina sp. Prony Australia (South 
Australia. New South Wales. Queensland and 
Western Australin) and New Guinea | Berry 1985). 

Estat turtles are vrouped inta Lwe suborders on 
the basis of differences in ncek vertebrae Mexion 
during head retraction, AIP Australian turtles are 
members of the Suborder Pleurodira, a group 
characterised by sideways Texion al the neck 
vertebrae, whieh had a cosmopolitan distribution 
before the separation of Gondwanakine and Lanriesi 
120-100 million years ago (mya) (Paugh eral 1990). 
Theit modern distribution is restricted to the 
Pelomedusidie of Soath Africa, the Podocnemidie 
of South America und the Chelidae of Sout 
America. New Guinea and Australia (Cogper 1996), 
All remaining freshwater and marine titles are 
members of the Suborder Cryptodira, a group 
characterised by vertical Texton ofthe neck, 

Th this study. §. ehedadinae wis found in Lanvedeere 
kroffii Gray in Eastern Australia, Camella 
chelenius Baker, 1983 Is reported for the first time 
from the pleurociriun turtles Bvivdure krefitit, Blveva 
latixiernum Gray, Chelodina expansa Gray and ©. 
loneicallis Shaw from Australia 


Muterials and Methods 
A total of 86 turtles, | Chelading expansa & Elseya 


lufisiermun and 77 Endura Krefiri, Was taken fron 
6 Queensland catchments using wt batted crah pot or 


86 M.A, FERGUSON & 1. R. SMALES 


COOKTOWN 15°5 
fag 
CAIRNS @ 
. 
TOWNSVILLE @, 
20°S 
ROCKHAMPTON 
. 
BUNDABERG #; OS'S 
Hig. 1 Distiburaan ol the Nemimdes Craellanis 


Chitty and spires ehelvdtiae (ran Baevdure 
Arelini Plsewa hvisternin and Claladime expanse tall 
locales sumpled HW Queenstind from May [9% to Dee 
WOT B= Camaflinay elretunins, dy = Spiracys 
echelon 


fiand line. The lollowing localities were sampled: 
Cooktown, 1S 28° 8, 145° (So RE (2 EF, letivtermunys 
Couns 16° 45°58, 145° 47° E C1 A. leatisternuin | Ben 
Aeeditiy: Townsville 19 16" S. 146° 49° RIS Ene 
Kveffit, Proserpine 20" 24'S. 148" 35' (Sb. 
lativieranin, WO Bin, krefftiiy;, Rockhampton 23° 22'S, 
(50° 32° BCL ©) expanse, 500 Eat Arefftity and 
Bunduberg 24° 52'S, 152" 2 ECL Ean hrefftii) (Fig, 
}). "The turtles were cothanased by cervical injection 
of at least 2.5 ml oof Pentobarbitone sodium 
(Nembund o) per ke of turtle, Che turtles were 
dissected then ulboruns were examined with the acl 
Ol a dissecting microscope, All nematodes found 
Were washed in saline, fixed in near boiling 10% 
formalin then stored jn 70% ethanol, Speeiiens 
from the Queensland Museuin. Brisbane (OM) fron 
©. lungicallis, localily unknown (G207571). Tran &. 


favisternin, Mulgrave River, Glo Atish and 
Edinontou. Queeushiwad (G2132398, G213239, 


G213241). and Troi Min, Aresfii, Mulgrave River, 
Queensland (G213240), were alse examined for 
comparative pHeposes, Specimens for detailed 
Microscopic eauminulion were cleared in 
Juctophenol, All measuremenis are in im unless 
olherwise indicated vind were made using an 
eyepiece micrometer. Voueher species, dos 


Photomicrograph af antecior at Camathanrnes 
tetas Fron Eaadure krefiiiin Arrow 
chitnous bueeul nidees, Seale bur = 35 yun, 


b 
Nip 2 
Tichicales 


G213990-G 14004. bave heen deposited i the 
Queenshind Museum, Brishane. Ecological 
terminology conforms to the definitions of Marsalis 
ebal, (WY82), 


Results 


Adult Apacs efeladie were tecovered Lom 
granulomas in the stomiuch of 22 of 77 hin, kreffiii 
examined (28.6% prevalence), but not from other 
species examined. Examination shawed that the 
granuiomas originated oi the mucosal layer, but in 
some cuses had extended into the submucosa une 
muscularis, and could be seen in the serosa on the 
extermil surface of the stomach. Eneysted hirvae 
were also found in the intestinal walls. mesenteries 
and liver, but numbers were not recorded. Adult 
Comins chelonius (Pig. 2) were recovered (hom 
the small intestine of 19 Of 77 Bin. Areffiti (24,7 
prevalence), 7 OFS FE, lafisternon (87.3% prevalence) 
and the single ©. expease examined. OF 77 
specimens examined. 4 bi Areffii were tnfectcul 


NEMATODES FROM AUSTRALIAN TURTLES 1K7 


TABLE L. Comparative body measmements, ov um unless otherwise stated, of male and female Spiroxys chelodinae froma 
Emydura kreffit fro Rockhampron, Queensland und the paratypes of S. chelodinae, (paratype measurements from Berry 


POSS |. 
Means in parenthesis. 
Specmnens from Enryeura kref{tit Paralypes 
Male Female Male Female 
n=5 iL=4 n=10 nw=10 
Total length Gm) [8-25 (21) [4-27 (22) 73-273 TY-28.6 
Masini width 429-516 (457) 374-563 (491) 179-545 171-860 
Lenuth pseudolabiam 36-495 (42) 40 (n= 1) 31-65 23-645 
Width pseudolabium 83-112 (101) 76 (n= 1) 53-114 hd-136 
Length muscular pharyax 435-415 (A80) 408 (n= 1) - - 
Leneth glandular pharynx 2500-3100 (2883) 2400 tn = 1) - - 
Length pharynx (mm) 3.26 (n= 1) 261 (n= 1) 1.534-3.61 1,72-3.59 
Nerve ring from anterior 536-71 (O09) 4&9 (n= 1) 332-21 435-643 
Secretory-excretory pore 
from anterior YOL-1173 (L037) 424-K 11) AIO-89 | 
Deirids from anterior (02-1224 (1099) 624-| 334 O44-[312 
Leneth gubernaculum 115-168 (142) 108-295 - 
Length spicule 1230/1630 (1455) 770-2410 - 
Length tail (74-201 (189) 245-268 (254) 96-281 121-459 


TABLE 2, Comparative bouv measurements, tr unt iwiless othenvise stated, of male cud fenuile Camatlanus cheloaius fron 
Elseya fatistermum fre Queensland ane C. chelonius fron South Africa (measurements from Baker PYs3). 


Means in parenthesis. 


Specimens from 


Specimens from 


Queensland South Africa 

Male Female Male Femiale 

n=5 n=5 n=4 n-4 
Total length (mm) 9-14 (12) 13-24 (18) 10.6-16 17.2-23 
Maximum width 181-409 (261) 340-516 (388) - - 
Length buccal valves 101-127 (119) 134-151 (140) - - 
Width buccal valves 107-147 (121) 134-168 (151) - - 
Length muscular pharynx 429-502 (474) 489-594 (550) 506.595 518575 
Length glandular pharynx 608-1020 ($22) M84- 1054 (949) S3K-YS0 O31 1094 
Nerve ring from anterior 231-241 (237) 235-288 (270) 214-251 275-281 
Secretory - excretory pore fromanterior = 523 (n= 1) - 
Deirids from anterior 663-705 (714) 
Vulva from anterior (mim) < TUS) - h4-12.4 
Length right spicule 450-460) (455) - 522-572 - 
Length left spicule 450-460 (455) - 325-384 - 
Length tail 100-175 (132) {81-261 (214) 135-175 222-428 


with both C. chelonins and 8. chelodinae, 1S were 
infected with ©. eheloniuy only (19.5% prevalence) 
and 18 were infected with 8. ehelodindae only (23.4% 
prevalence), Splrayys chelodinae was tound at 
Bundaberg, Rockhampton and Townsville only. 
Camallanus chelonius was found at all localities 
except Bundaberg, but this is probably because only 
one host specimen was examined at this locality, 
Measurements of $8. chelodinae trom this study are 
in the range reported by Berry (1985) for 8. 


chelndinae occurring in Chelodina sp. from Western 
Australia, South Australia, New South Wales. 
Queenslind and Papua New Guinea (Table 1). The 
quality of the female specimens was such that few 
measurements could be taken, but no measurements 
were outside the range reported by Berry (1985). 
Specimens of C. cheloninus from this study 
contorm to the description by Baker (1983) 
especially regarding buccal yalye morphology and 
the number and arrangement of caudal papillae. The 


18S M.A, FERGUSON & LR. SMALES 


| 


Fig, 3, Drawing of tip of right spicule OF ¢eonallares 
Chelonins trom Emyedtire kref/iii showing differenee trom 
South African speeunens of Baker (1983), Scule bar = 25 
ju 


measurements OFC, chelonius (Table 2) contorm for 
the tost parr to these given by Baker (1983) 
although there are differences in’ spicule length 
(shorter in Queensland specimens) aod female tail 
length (shorter in Queensland specimens), Also the 
slender, sharply pointed clongate process on the tip 
of the right spieule as figured by Baker (1983, Fig, 
1p. 163) appears to be shorter in the South Atrican 
specimens than the Queensland specimens (Fig 3). 
However, these differences do not appear to be 
significant enough to propose a mew species. 


Discussion 


Although specimens of Llseya and Laver were 
exuumined for nematodes by Berry (1985), the sourees 
ob these hosts Were not given. Spiros cleloddinae 
wis not recovered by Berry in either host genus: but 
has now been found in ban Areffir fron Bundabers, 
Rockhampton and Townsville, The finding ofS. 
thelodinae in kay kreffii in Central and Northern 
Queensland is a new host and docality record, 


This is the first record of ©) chelates from 
Australiin turtles. Camallanus chelonius has now 
known been reported from Australian, (this study) 
and South African (Baker 1984) pleurodiran turtles, 
The only other report of a camallanid from a 
pleurodiran turtle ts Serpinema amaconicus (Riberio, 
1941) Petter, 1979 from South America (Riberio 
1941). All other Serpinemai spp. are found in 
cryptodiran turtles, with each geographical region 
haying its own species (Petter 1979). 

Camallanus spp. have large numbers of 
unseparated buccal ridges (Fig. 2). while Serpine nie 
spp. have smaller numbers of distinctly separated 
buccal ridges. The similarity of spicule morphology 
and distribution of caudal papillae between C. 
chelonins and Serporenia spp.. however, indicates i 
close relitionship (Baker 1983). Also, the buccal 
morphology of S. aaacenicus und C. chelonius has 
been suggested to be intermediate between Case- 
Hanus and Serpinema. This suggests that Sespinentia 
may have evolved first in pleurodirans (Baker 1983), 
A camallanid ancestor of S$. amezenicus may have 
been captured by early eryptodiran turtles and 
radiated with the major cryptodiran radiation around 
120-90 mya (Shafter et af. 1997). The geographical 
distribution of Serpineme in cryptodiran turtles 
shows that the greatest diversity of fauna ts in South 
and Central America and tropical Asia (Stromberg & 
Crites 1974). The finding of §. ammezenieus in South 
America. an intermediary form between Camallanus 
and Serpineme. suggests that South America may 
haye been a point of origin for this genus. which then 
moved into North America and Asia with its hosts. 

The oecurrence of C. chelonifay in both Australian 
and South African pleurodires suggests that this 
species originated before the separation of Australi 
and South Africa, approximately 120-lOUmya. The 
close link between South Africa and South Amerion 
us recently as 90 mya und the similaritics between 
chelonius, S amezenicous and other Serpinemea spp 
SuBBeSsL i COMMON Origin for these groups, 


Acknowledgments 


Thanks go to C.J. Parmenter for assistance in 
turtle collection and jdentifieation, and to L. Cannon 
and K. Sewell of the Queensland Museum for 
providing access ta Museum Specimens. 


NEMATODES FROM AUSTRALIAN TURTLES 189 


References 


Baker, M. R. (1983) Nematode parasites of the turtle 
Pelusios. sinuatus (Pelomedusidae: Pleurodira) from 
southern Africa. Syst. Parasitol, 5, 161-167. 

(1984) Nematode parasitism in amphibians and 
reptiles. Can. J. Zool. 62, 747-757. 

Berry, G. N, (1985) A new species of the genus Spiroxys 
(Nematoda; Spiruroidea) from Australian chelonians of 
the genus Chelodina (Chelidae). Syst. Parasitol. 7, 59-68. 

Bays, H. A. (1933) On a collection of nematodes from 
Malayan reptiles. Ann. Mag. Nat. Hist., Ser. 10 11, 615- 
633. 

CoaGcer, H. G. (1996) “Reptiles and amphibians of 
Australia” (Reed Books, Melbourne). 

Maraous, L. G., Escu, G. W., HOLMES, J. C., Kurs, A.M. 
& ScnaAb, G, A. (1982) The use of ecological terms in 
parasitology. J, Parasitol, 68, 131-133. 


Perrer, A. J. (1979) Essai de classification de la sous-famille 
des Camallaninae (Nematoda, Camallanidae). Bull. Mus. 
natn. Hist. nat., Paris, 4 sér, Sect. A 1, 991-1008. 

PouGH, F. H.. HkISER, J. B. & MCFARLAND, W. N. (1990) 
“Vertebrate Life’ (Macmillan Publishing Co.. New 
York). 

Riperto, D. J. (1941) Pesquisas helmintologicas realisadas 
no Estado do Para. VIL - Camallanus amazonicus n. sp. 
parasito de Podocnemis expansa (Schw.). Mem. Inst. 
Oswaldo Cruz 35, 723-727. 

SHAPFER, H. B., MEYLAN, P. & McKnicut, M. L. (1997) 
Tests of turtle phylogeny: molecular, morphological and 
paleontological approaches. Syst. Biol. 46, 235-268. 

STROMBERG, P. C. & Crites. J. L. (1974) Specialisation, 
body volume and geographical distribution of 
Camallanidae (Nematoda). Syst. Zool, 23, 189-201. 


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