VOL. 117, PARTS 1 & 2

4 JUNE, 1993

Transactions of the

Royal Society of South

Australia

Incorporated

Contents

Hutchings, P. A., Ward, T. J., Waterhouse, J. H. & Walker, L. Infauna of marine

sediments and seagrass beds of Cpr: Spencer Gulf near Port Pirie,

South Australia - - - - z : £ : 4

Barker, S. Seventeen new species of Australian Buprestidae (Insecta: Coleoptera) and a

host plant of Castiarina uptoni (Barker)- - - - - -

Beveridge, I. & Durette-Desset, M.-C., Adult and larval stages of Paraustrostrongylus

ratti (Nematoda: Trichostrongyloidea) from Rattus fuscipes

Campbell, R. A. & Beveridge, I. New species of Grillotia and Pseudogrillotia (Cestoda:

Trypanorhyncha) from Australian sharks, and definition of the ey

Grillotiidae Dollfus, 1969 - - - - E : 2

Baker, G. H., Barrett, V. J., Grey-Gardner, R. & Buckerfield, J. C. Abundance of

life history of native and introduced earthworms (Annelida:

Megascolecidae and Lumbricidae) in pasture soils in the Mount baie

Ranges, South Australia - - - - - - -

Gatehouse, C. G., Jago, J. B., Clough, B. J. & McCulloch, A. J. The Early Cambrian

volcanics from Red Creek, eastern Mount Lofty Ranges, South Australia

Austin, A. D., White, T, C. R., Maelzer, D. A. & Taylor, D. G. Biology of Etiella behrii

Zeller (Lepidoptera: Pyralidae): a pest of seed lucerne in South Australia

Lee, D. C. Incabates Hammer and Setincabates gen. nov. (Acarida: Cryptostigmata:

Haplozetidae) from South Australian soils - - = i :

Sprigg, M. & Bone, Y. Bryozoa in Coorong - type lagoons, Southern Australia

Wells, A. & Cartwright, D. Females and immatures of the Australian Caddisfly Hyalopsyche

disjuncta Neboiss (Trichoptera), and a new family placement- -

Brief Communications:

Johnston, G. R. & Richards, S. J. Observations on the breeding biology of a taierenyh

frog (Genus Oreophryne) from New Guinea- - -

Crossland, M. R. & Richards, S. J. The tadpole of the Australopapuan frog L Litoria

nigrofrenata (Gunther, 1867) (Anura: Hylidae) - -

PUBLISHED AND SOLD AT THE SOCIETY’S ROOMS

SOUTH AUSTRALIAN MUSEUM, NORTH TERRACE, ADELAIDE, S.A. 5000

27

37

47

57

97

105

109

TRANSACTIONS OF THE

ROYAL SOCIETY

OF SOUTH AUSTRALIA

INCORPORATED

VOL. 117, PART 1

TRANSACTIONS OF THE

ROYAL SOCIETY OF SOUTH AUSTRALIA INC.

CONTENTS, VOL. 117, 1993

PARTS 1 & 2, 4 JUNE, 1993

Hutchings, P. A., Ward, T. J., Waterhouse, J. H. & Walker, L. Infauna of marine sediments

and seagrass beds of Upper Spencer Gulf near Port Pirie, South Australia

Barker, S. Seventeen new species of Australian Buprestidae (Insecta: Coleoptera) and a host

plant of Castiarina uptoni (Barker) - - - - - - -

Beveridge, I. & Durette-Desset, M.-C. Adult and larval stages of Paraustrostrongylus ratti

(Nematoda: Trichostrongyloidea) from Rattus fuscipes - - -

Campbell, R. A. & Beveridge, I. New species of Grillotia and Pseudogrillotia (Cestoda:

Trypanorhyncha) from Australian sharks, and definition of the family

| Grillotiidae Dollfus, 1969 - - - - - - - -

Baker, G. H., Barrett, V. J., Grey-Gardner, R. & Buckerfield, J. C. Abundance and life

history of native and introduced earthworms (Annelida: Megascolecidae

and Lumbricidae) in pasture soils in the Mount Lofty Ranges, South

Australia - - - - - - - - - - -

Gatehouse, C. G., Jago, J. B., Clough, B. J. & McCulloch, A. J. The Early Cambrian

volcanics from Red Creek, eastern Mount Lofty Ranges, South Australia

Austin, A. D., White, T. C. R., Maelzer, D. A. & Taylor, D. G. Biology of Etiella behrii

Zeller (Lepidoptera: Pyralidae): a pest of seed lucerne in South Australia

Lee, D. C. Incabates Hammer and Setincabates gen. nov. (Acarida: Cryptostigmata:

Haplozetidae) from South Australian soils - - - - -

Sprigg, M. & Bone, Y. Bryozoa in Coorong - type lagoons, Southern Australia - -

Wells, A. & Cartwright, D. Females and immatures of the Australian caddisfly Ayalopsyche

disjuncta Neboiss (Trichoptera), and a new family placement - -

Brief Communications:

Johnston, G. R. & Richards, S. J. Observations on the breeding biology of a a ti

frog (Genus Oreophryne) from New Guinea - -

Crossland, M. R. & Richards, S. J. The tadpole of the Australopapuan frog L Litoria

nigrofrenata (Gunther, 1867) (Anura: Hylidae) - - -

27

37

47

57

67

77

87

97

105

109

PARTS 3 & 4, 30 NOVEMBER. 1993

Shiel, R. J. & Koste, W. Rotifera from Australian inland waters. IX. Gastropodidae,

Synchaetidae, Asplanchnidae (Rotifera: Monogononta) - - -

Tian-Rui, Lin & Jago, J. B. Xystridura and other early Middle Cambrian trilobites from

Yaxian, Hainan Province, China — - - - - - - -

Pell, S. D., McKirdy, D. M., Jansyn, J. & Jenkins, R. J. F. Ediacaran carbon isotope

stratigraphy of South Australia — an initial study — - -

Nicholas, W. L. Two new species of nematode (Nematoda: Enoplida: Thoracostomopsidae)

from Lake Alexandrina, South Australia - - - - - -

Hoste, H. & Beveridge, I. Interspecific and intergeneric relations between nematodes parasitic

in the stomachs of kangaroos and wallabies —- - - - _

Ryder, M. H. & Bird, A. F. Effect of Acrobeloides nanus (Nematoda: Cephalobidae) upon

the survival of Pseudomonas corrugata (Rubattetia) in ey pasteuy ae soil

from Kapunda, South Australia < > : +

Zbik, M. The Big Rock Donga Metcorite: A new H5 Chondrite from South Australia

Tyler, M. J. & Godthelp, H. A new species of Lechriodus Boulenger (Anuras Leeptodicty lide)

from the Early Eocene of Queensland — - -

Barker, S. A new Australian species of Calodema (Coleoptera: Buprestidae) - - -

Brief Communication:

Wallman, J. F. First South Australian record of the carrion-breeding blowfly Calliphora

nigrithorax Malloch (Diptera: Calliphoridae) — - - - - -

Errata, Corrigenda, Addenda - - - - . ¢ + 2 3 * -

insert to Transuctions of the Royal Society of South Australia, Vol. H?, parts 3 & 4, 30 November, 1993

171

179

183

187

191

193

194

INFAUNA OF MARINE SEDIMENTS AND SEAGRASS BEDS OF UPPER

SPENCER GULF NEAR PORT PIRIE, SOUTH AUSTRALIA

BY P. A. HUTCHINGS*, T. J. WARD**, J. H. WATERHOUSE** & L. WALKER***

Summary

The infauna of six intertidal and shallow subtidal habitats were sampled in summer and winter near

Port Pirie, Upper Spencer Gulf. The habitats included seagrasses and non-vegetated sediments.

Over 12,000 individuals belonging to 372 taxa were collected in the survey. The species

composition of this fauna had very little overlap (only 24 species) with that of a more recent survey

of Northern Spencer Gulf. The density of the fauna in our study area was least (280 m‘~) in the

unvegetated intertidal habitat, and highest (2612 m’°) in the intertidal seagrass habitat (Zostera), and

was dominated by polychaetes, molluscs and crustaceans. The fauna of the Zostera and Posidonia

habitats is compared with other similar habitats sampled in southern Australia, in terms of the

number of species, density of individuals and feeding guilds of the fauna. Differences in species

diversity and densities between this study and other studies are likely to reflect the intensity of

sampling, as well as differences between locations in the density and species of seagrass and in the

prevailing hydrological regimes.

KEY WORDS: Posidonia, Zostera, seagrass beds, unvegetated sediments, infauna.

Transactions of the Royal Sociery of S Ausz., (1993), LIT(), 1-45

INFAUNA OF MARINE SEDIMENTS AND SEAGRASS BEDS OF UPPER SPENCER

GULF NEAR PORT PIRIE, SOUTH AUSTRALIA

by BR A. HUTCHINGS* T. J. WARD**, J. H. WATERHOUSE** & L. WALKER***

Sumumury

Hurcutnas, P_ A., WARD, T, J,, WATERHOUSE, J. H., & WALKER, L. 1993) Infauna of marine sediments and

seagrass beds of Upper Spencer Gulf neur Port Pirie, South Australin. Trans. R. Soc. $. Ausr. Wy, 1-15, 4

uns, 1993.

The infauna of six intertidal and shallow subtidal habitats were sampled in sammer and winter near Port Pirie,

Upper Spencer Gulf, The habitats included seagrasses and non-vegetated sediments, Over 12,000 individuals

belonging to 372 taxa were collected in the survey. The species composition of this fuuna had very litile overlap

(only 24 species) with that of w more recent survey of Northern Spencer Gulf. The density of the tauna in our

study arca was least (280 m) in the unvégetated intertidal habitat, and highest (2612 ny) in the intertidal seagrass

habitat (Zastera),.and was dominated by polychaetes, molluscs and crustaceans. The fauna of the Zostera and

Posidonia habitats is compared with other similar habitats sampled in southern Australia, in terms of the number

of species, density of individuals and feeding guilds of the fauna. Differences in species diversity and densities

between this stady and other studies are likely to reflect the incensity of sampling, as well as differences between

facunons in ihe density and specios of seagrass and in the prevailing hydrological regimes.

Key Worps: Fosidonia, Zostera, seagrass beds, unvegetated sediments, infauna.

Introduction

Upper Spencer Gulf near Port Pirie (33°19'S,

138°E), South Australia, is shallow (penenilly less than

20 m deep) and fringed with large expanses of intertidal

mudflats, intertidal seagrasses (Zostera spp.), and

subtidal beds of the scagrasses Pasudona australis and

P sinwosa, These shallow expanses of seabed are

fished commercially and recrestionally for a number

of species of fish, including garfish and whiting, and

for crustaceans including prawns and crabs (Jones

1979'; King 19777). Although these. and many other

species, use the seagrass beds and intertidal flares ar

various stages of their life cycle Jones 1979'), there

have been few ecological studies designed to document

other species of importance occurring in these habitats

of the Gulf.

Upper Spencer Gulf refers to the arca of the Gulf

between Port Davis Creek (just south of Port Pirie)

*« tnvertebrate Zoology, The Australian Museum, P.O, Bon

A285, Sydney South, NSW 2000

** CSIRO Division of Fisheries, P.O. Box 20,. North Beach,

WA 6020

‘e* jormerly of The Australian Museum now of 306,

Moleswurth St,, Lismore, NSW 2480

WONES, G. I. (1979) Biological Investigations of the Marine

Scale Fishery in Spencer Gulf. (Dept Agriculture and

Fisheries, South Australia). 72pp.

“KING, M. G. (1977) The biology of the Western King Prawit

Fengeus latisulcaius Kishinouye and aspects of the fishery

in South Australia. MSc. thesis, University of Adelaide,

unpubl

and Ward Spit, and Northern Spencer Gul! is defined

as the portion of Spencer Gulf north of Ward Spit, 1.c.

north of 33°S (Stefanson 1977). Spencer Gull, is

hypersaline, with salinities gradually increasing from

35 “/ nto more than 60 °/ oo near Port Augusta at the

head of the Gulf (Bullock 1975), Upper Spencet Gulf

has an intermediate salinity regime, varying annually

From about 41-47.2 °°... and water temperatures ringing

from 10-26.2°C (Dow 1980°). Studies of fish and

crustaceans in this area by Ward & Young (1982) have

shown that they appear to be unaffected by the constant

high salinines.

In this study we examine the mainly invertebrate

fauna Lying in the sediments in shallow-water habitats

found in Upper Spencer Gull near Port Pirie, and its

species composition. These results are compared with

those obtained from sitmilur habitats elsewhere in

southem Australia

Similar hatitats which have been studied in detail

elsewhere in southern Australia are those of seagrass

beds and intertidal habitats. The infauna of seagrass

beds has been examined in several studies in eastern

Australia (Collett et af. 1984; Hutchings & Recher

1974), und in Western Australia (Hutchings et al. 1991,

Wells et al. 1985) and reviewed by Howard es al,

(1989), The infauna of intertidal habitats in Northern

Spencer Gulf has recently been studied by Ainslie ev

al, (1989),

DOW (1980) Redcliff Project — Deaf Environmental Effects

Statement. 178 pp. (Dow Chemical (Australia) Limited,

Adelaide SA,}.

te

Materials and Methods

Samples of sediment were collected at high tide from

18 sites representing six habitats, by divers using PVC

corers (220 mm diameter and 160 mm depth) in winter

(25Nii.1979 — 17.xiii.1979) and in summer (1.ii,1980

— 1.ii1.1980). Each corer had a sharpened leading edge

and was capped at the other end by a PVC plate welded

in place, A 50 mm hole in the cap covered with 1 mm

fibreglass. mesh allowed water to escape as the corer

was driven into the sediment, and, after coring, was

sealed with a rubber bung. No attempt was made to

avoid seagrass leaves when they occurred at a sampling

=——

¢

“=

—

P. A. HUTCHINGS, T, J. WARD, J. H, WATERHOUSE & L. WALKER

site, and the sampling technique resulted in substantial

amounts of live and dead seagrass leaf material,

rhizomes and roots in the samples. The habitats

sampled (Fig. 1) were from about mid-tidal level (on

mudflats adjacent to the mangroves) to sediments in

10 m water depth, and are the same habitats in which

detailed studies of sediments and epibenthic fauna have

been carried out previously (Ward & Young 1981,

1982). The habitats included bare intertidal muds (Sites

1A, 1B, 1C), intertidal seagrass beds of Zostera

mucronata (Sites 2A, 2B, 2C), subtidal beds of

Posidonia australis (Sites 3A, 3B, 3C) and. Posidonia

sinuosa (Sites. 4A, 4B, 4C) and bare unvegetated

Ward Spit _.-

SPENCER GULF

Fig. 1. The Port Pirie study area in Upper Spencer Gulf.

mangroves:

Port =

“Yy~pPirie

smelter af River a

PORT PIRIE

INFAUNA OF MARINE SEDIMENTS AND SEAGRASS BEDS OF UPPER SPENCER GULF 3

sediments (Sites 5A, 5B, 5C and Sites 6A, 6B, 6C) in

deeper water (Fig. 1). At each site three samples were

collected haphazardly from each of three plots of about

4 m? separated by at least 5 m. This resulted in nine

samples from each of the 18 sites, collected in winter

and summer, except for summer, when one sample

from the Posidonia australis habitat was lost.

The samples were sieved in the field through 2 mm

mesh and all residues were preserved in 10% formalin.

In the laboratory the samples were sorted under water

in large flat trays with the aid of low-power

magnification. Hermit crabs were discarded because

of the difficulty of extracting them from their shells,

but all other non-colonial fauna was collected. Colonial

organisms such as bryozoans and hydroids attached to

the blades of the seagrasses were not sampled in this

study, Because of taxonomic difficulties and a lack of

funds the collection was stored for six years until

resources permitted a detailed evaluation. The

collection has been deposited at the Australian

Museum, Sydney.

Results and Discussion

Species Composition of the Fauna

The samples contained 12,396 individuals

representing 372 taxa. These taxa consisted of 39%

polychaetes, 26% molluscs, 22% crustaceans, and 6%

echinoderms, with the remaining 7% being distributed

among the minor phyla (Table 1). The distribution and

abundance. of all taxa within the six habitats sampled

is shown in Appendix 1. Of the 26 dominant taxa.

defined as those represented by 100 or more individuals

in the collection, 18 were polychaetes, four were

molluscs, three were crustaceans and one was a

holothurian (Table 2). The taxa identified as

nemerteans and sipunculans may not be monospecific

because of difficulties in identifying individuals to the

species level. The most abundant species was Tanais

dulongi, a tanaid crustacean.

The six habitats sampled varied considerably in the

number of species present. The four subtidal habitats

all had a high numbers of species (173 — 193)

compared to the intertidal Zostera habitat (116), and

TaBie 1. Infauna of Upper Spencer Gulf! Numbers of species (5), density (d, number of individuals m°*), abundance (n,

% of toral individuals in habitat) and total abundance (N) of the major taxa in each of the six habitat types sampled. All

habitats were sampled with 54 cores (total surface area of 2,05 m*), except Posidonia australis which was sampled with

53 (total surface area of 2.01 nx’).

taxa

intertidal Zostera P. australis

polychaetes

Rn 57 94

d 243 1262 884

n(%) 87 48 76

crustaceans

Ss 5 20 34

a 16 907 120

t1(%) 6 35 10

molluscs

s 6 23 34

d- 13 330 §4

n(%) 5 B 7

ascidians

S 0 4 3

d - ul 6

n(%) = 05 0.5

echinoderms

8 1 3 »

d 0.5 39 37

n( 9%) 0.2 2 3

minor taxa*

3 6

d 8 62 27

ni %) 3 2 2

all taxa

26 15 Lil

N S74 5362 2329

d 280 2612 1157

habitat

P. sinuosa 5m bare Win bare total

9o7 1 88 148

630 501 242 626

73 76 49 62

44 23 18 BI

76 28 19 nH

9 4 4 i]

32 38 50 95

95 92 183 133

iT] 14 37 b

5 7 2 10

i 16 it 9

L 2 2 1

12 Q 10 20

38 14 32 za

4 2 7 3

4 3 4 18

ul 3 0 2

1 1 2 2

194 171 172 372

1764 Bs) 1017 {2396

860 660 497 1010

* Minor taxa includes fish, insects, sipunculans, poriferans, nemerteans, pycnuyonids, enteropneusts and hirudineans,

4 P A. HUTCHINGS, T. J. WARD, J. H. WATERHOUSE & L. WALKER

TABLE 2. Abundances (%) of the dominant species — total abundances of 100 or greater — in each of the the six habitat types.

ee ee a ee SE

intertidal Zostera P. australis

Tanais 26 (3) 819 (97) -

dulongii

Capitella 51 (8) 6l1 (92) -

‘‘capitata”’

Nereis - 31 (5) 577 (89)

bifida

Apseudes - 513 (87) 63 (H)

dustralis

Leitoscoloplos 31 (5) 414 (71) 121 (2))

Sp.

Nephtys 109 (20) 99 (18) 13 (2)

gravieri

Ceratonereis 248 (65) 8 (2) 34 (9)

sp. |

Pista - 340 (91) 6 (2)

violacea

Platynympha 2 (0.6) 291 (90) 22. (7)

longicaudaia

Barunwlla sp, = 88 (31) 57 (20)

Austromytilus - 45 (2) 2 (0.8)

penectus

Nematonereis - - 103 (41)

unicornis

Augeneria - 131 (55) 18 (8)

verdis

Tellina - 226 (95) -

deltoidalis

Zeacumantis 1 (0.5) 206 (99) -

diemenensis

Eunice sp. 3 - - Al (21)

Notomastus 2 (lh 96 (52)

sp. J

Tellina 6 (3) TL (41) 20 (12)

mariae

Leptosynapta = 74 (49) 42 (28)

dolabrifera

Agyychis sp. - - 5 (4)

Cirriformia - 43 (34) 43 (34)

sp. 1

Caulleriella - 20 (18) 31 (27)

sp. 4

Eunice sp. 1 2 - 5 (5)

Diplocirrus sp. - 92 (90) 3 (3)

Scoluplos 1 (i) 74 (73) 17 (17)

simplex

cas Pere ~ 7 (7) 35 (35)

Sp.

totals 475 (6) 4165 (52) 1354 (17)

total abundance (all species)

574 5362 2329

% abundance dominant/abundance total ‘

5

P sinuosa 3m bare 10m bare total

- - 1 01) 846

- - = 662

29 (5) 70) 1 (0,2) 645

2 (0,3) 7) 3 (0.5) 588

13 (2) 1 (0.2) - 580

U5 (21) 149 (27) 67 (12) 552

14 (4) 69 (18) 10 (3) 383

12 (3) 3 (0.8) 12 (3) 373

8 (2) - - 323

88 (31) 46 (16) 6 (2) 285

2 (0.8) 57 (21) 211 (76) 277

18 (47) 25 (10) 7 (3) 253

49 (20) 26 (11) 16 (7) 240

8 (3) 5 (2) - 239

= 7 - 207

an 2548) B®)

45 (26) [2 (7) 18 (10) 172

33 (22) - I (0.7) 150

wey 4

38 (34) 21 (19) 3 (3) 13

81 (74) 19 (17) 4 (4) 109

4

43 (43) 14 (14) 1 (1) 100

859 (H) 661 (8) 439 (6) 7953

1764 1350 1017 12396

49 49 43 64

aE ee eee ee

the intertidal mud, which was by far the most species-

poor habitat with only 27 species recorded (Table 1).

The pattern of densities amongst the six habitats was

generally similar to that of species richness except for

the Zostera habitat. This habitat had the highest density

of fauna (2612 individuals ny?) with the second lowest

species richness (115 spp.).

Overall, polychaetes and molluscs were the two

dominant groups in terms of both numbers of species

and individuals, The crustaceans were generally similar

to the molluscs in species richness and abundance in

ithe two Posidonia habitats and the intertidal mud, but

were considerably reduced in the two subtidal

unvegetated habitats. Echinoderms, ascidians and the

INFAUNA OF MARINE SEDIMENTS AND SEAGRASS BEDS OF LIPPER SPENCER GULF 4

minor taxa were represented by only Jew species and

typically few individuals in all habitats.

Of the 32,396 individuals collected 7,953 (64%)

belonged to only 26 (7%) of the 375 species. The

remaining 346 species were represented in most cases

by only a few individuals (Appendix 1). Ninety-four

species Were Tepresented by only a single individual

in the collection and 155 species were present in only

One babital. Similar patterns have been found in other

studies of the infauna of Austratian seagrass beds

{Collett er al, 1984; Hutchings et al. 1991).

The 26 dominant species were distributed unevenly

between the six habitats (Table 2). The two most

dontinant species (Timwis dulongi and Capitella

\‘capitata”’) were almost exclusively restricted to the

two ineertidal habits, and predominantly in the

Zastera. Several other species also occurred

predominantly in the Zostera habitat — Apseudes

gustralis, Leitoscoloplos wormalis, Pista violacea,

Playympka fongicwidata, Tellina delroidatis,

Zeacumantus diemdénensis, Diplocirrus sp. and

Scolaplos siniplex. Nereis bifida and Eunice sp. |

octurred oaainly in the PB australis and P sinuese

habitats respectively, while Asychis sp. was mainly

found in the Sm subtidal bare habitat. Only four of

the deminan! species (Nepthys graveri, Ceratonerets

sp, J, Tellina mariag, and. Scotoplos simplex) were

present in fll six habitats. Tweaty of the 26 dominant

were foutd in all the subtidal habitats. Thus the

dominant species could be separated into two groups:

those dominant in the intertidal and those dominant

in the sabtidal, with few species common to both

groups, A detailed analysis af the variability of species

composition amongst individual habitats, sites, seasons

and sediment characteristics will be presented

elsewhere (Ward & Hutchings in prep.)

Comparisons with other areas

Intertidal unvegeiated sediments —

A recent study by Ainslie et al, (1989) of the mfauna

of intertidal unvegetated inudflats of Norther Spencer

Gulf (about 60 km north of our suidy area) recorded

a total af 99 species, These 99 species. included 31

species of polychaetes, 31 molluscs, and 22

crustaceans. This is considerably greater than the total

of 27 species which were recorded from intertidal

unvegetated habitats in this study. The densities of

individuals recorded in the Northern Spencer Gulf

intertidal habitats (about 550 in*} was also

considerably greater than those recorded in this study

(280 m*). These differences may result from the

different scales of sampling in the two studies or to

real differences between the. locations related to

differences in the hydrological conditions between the

two Spencer Gulf jocations; Similar sampling

techniques were used in both studies although Ainslie

etal, used a} sms sized mesh, and 20 replicates were

collected af each site in summer and winter from Jane

1982 to June (986 inclusive, which is a far greater

intensity of sampling than carried out in this study. The

Ainslie et al. (1989) study saanpled a total area of

33 m? in Northern Spencer Gulf whereas we sampled

about 2 m2. This difference in total atea sampled and

the extended temporal scale of the Northern Spencer

Gulf study may have contributed to the grealér number

of species of infauna recorded. Alternatively, the Gulf

near Port Pirie may have a depauperate incertidal

infauna because of different salinity and temperature

regimes,

The species composition of the infauna found by

Ainslie et al. (1989) in Northern Spencer Gulf ts not

similar to that fund in this study, with very few species

being common to both areas. Ainslie et al, identified

a total of 100 taxa to species level, and in this study

we identified 178 taxa to species. Of these only 24

species are common to both Upper and Northern

Spencer Gulf. Obviously, some of the taxa ientified

only to genus may, after further study, be found to

occur in both regions. We suggest that this apparent

difference in species composition may be related to

the different hydrological regimes in these two regions

of Spencer Gulf. Northern Spencer Gulf ts

characterised by high summer salinities, up to 48 Soo

(Nunes 4 Lennon 1986), and high summer

teniperatures with wide seasonal Nuctuatlons LL —

25°C Yohnsen 1981*; unpublished data quoted by

Ainslie et al. 1989). Northern Spencer Gulf is

seperated from the Upper Gulf by a narrow channel

just porth of our study area, and this could be

responsible for maintaining a different set of

hydrological conditions between these two study areas,

ever though they are close to cach other

Posidovia australis seagrass beds

The low-intertidal seagrass habitat of Posidonia

australis and P. sinuosa, in Northern Spencer Gulf,

contained 147 species of infauma with a mean density

af bout 170 individuals mt* (Ainslie er al. 1989), The

fost similar habitat in our study (P aussreis)

contained! 18) species with an average density of 57

individuals m*. Ainslie ef afl. do not give any

indication of the density of seagrasses ii this habitat,

but, because Aozidania is basically a suftidal seagrass

species, we presume that thelt sample location

comamed only scattered seagrass on the trrargins of

more subsiantial subtidal beds of P australis,

P. australis seagrass beds elsewhere In southern

Australia have also been studied, In Table 3 the

distritutions of the major phyla present in P. australis

TJOHNSON, LE, (981) Hydrological dats from U

Spencer Gulf 1975-1978 Fisheries Res. Paper No, 3; 0

(Dept Fistieries, Adelaide, SA, unpubl.)

6 P. A. HUTCHINGS, 'T, J. WARD, J. H. WATERHOUSE & L. WALKER

TABLE 3. Posidonia australis: distribution of the major phyla (§ = number of species, d = overall mean density).

site Polychaetes Molluscs Crustaceans Echinoderms Ascidians Minor Total

Albany (Hutchings ef al. 1991)

s 47 26 bt a iL 4 113

a** 73) 691 656 B32 124 ~ 2327

Spencer Gulf (this study)

8 94 34 a4 10 3 6 J81

u** 883 84 9 37 6 27 56

NSW (Collen et al, 1984)

$ 134 1 106 # ? 32 323

a‘ 1656 285 lf) 13 2763

* amphipods not identified to species

# echinoderms and ascidiany included in minor

™ densities (number of individuals

Gulf = 2.014 m*, NSW = 5.74 m°*)

in Albany, SW West Australia (35°03‘27"S 117°03°

27"W) (Hutchings er al. 1991) and at nine sites along

the NSW coast from 32°13'S to 36°54'S (Collett et

al. 1984) are compared with those occurring in Upper

Spencer Gulf. Polychaetes dominated the infauna in

these three geographical localities, both in terms of

number of individuals and species, The average densily

of infauna in NSW Posidonia beds was twice that of

Upper Spencer Gulf, although the NSW data were

based on samples totalling 5:74 m? in area, collected

at nine locations along the NSW coast. Similarly, the

P. australis sampled in Albany (total area sampled

1.2 m*) had a mean density of infauna about twice

that found in Upper Spencer Gulf in this study.

The species composition of the infauina of P. australis

in (hese three geographical areas is different, indicating

that although P australis has a wide distribution,

occurring from Shark Bay in Western Australia around

southern Australia to Wallis Lake on the central const

Taste 4, Posidonia australis: distribution

of species (%)).

location herbivore suspension

Albany (from Hutchings ef al., 1991)

polychaetes 4 (9) 3.(7)

molluscs 7 (33) 7 (33)

cmistaceans - -

uscidians . ut (100)

echinoderms - 2 (iS)

totals fi (hy 23 (22)

Spencer Gulf (this study)

polychaetes u (12) 12 (13)

molluscs 15 (44) 13 (38)

crustaceans I (3) 1 (3)

as¢idians - 3 (100)

echinoderms 1 (10) 1 (0)

totals 27 (16) HW) (17)

NSW (from Collett ef al., 1984)

polychaetes - 28 (21)

molluscs 10. (19) 24 (47)

crustaceans 3@) (4)

tonal B (5) 56 (19)

phy it

mm) calculated from total abundances and total area sampled (Albany = 1.2 m2, Spencer

of NSW (Larkum & den Hantog 1989), the composition

and density of its infauna vary according to location,

The infauna may also be related to the density of the

seagrasses themselves, but at the Upper Spencer Gulf

and Albany sites the density and productivity of the

P australis beds are similar (Watd 1987; Walker et al.

1991). Collett er al. (1984) report only a qualitative

assessment of the density of seagrass blades which

cannot be related to the numeric values determined for

the Upper Spencer Gulf and Albany sites.. Therefore,

hecause of both different sampling. intensities and

seagrass densities, it is not possible to directly compare

the density or diversity of the infauna found by each

of these studies.

In this study we also examined whether the feeding

stmitegies employed by the infauna were similar

regardless of geographical location. The fauna was

assigned to one of five feeding categories (Table 4) for

Upper Spencer Gulf, Albany and the nine locations

of the fauna amongst feeding vategories at Albany, Spencer Gulf and NSW (number

deposit carnivore omnivore

20 (44) 18 (40) -

1 (5) 6 (29) -

5 (36) 9 (64) -

4 G1) 5 (38) 2 (15)

30 (29) 40 (38) 2 (2)

40 (43) 30 (32) 1 (1)

2 (6) 4 (12) -

19 (56) 13. (38)

3 30) 5 (50) :

64 (37) 52 (30) 1 (1)

53 (39) 37 (28) ¥ (12)

7 (Id) 4 (8) 1 (2)

28 (26) 20 (19) 44 (41)

‘88 (30) 61 (21) 61 (21)

INFAUNA OF MARINE SEDIMENTS AND SEAGRASS BEDS OF UPPER SPENCER GULF 7

along the NSW coast, The designation of feeding

categories was determined from the literature (mostly

for related species) and from personal observations,

Camivores, including scavengers, comprised 21 —

38% of the total infauna in a geographical region. The

proportion of deposit feeders tanged between 29%

(Albany) to 37% (Upper Spencer Gulf). Suspension

feeders, which may feed on suspended particulate

™matier as well as plankton, were between 17 and 22%

of the fauna (Table 4). There were major differences

in the herbivore and omnivore fauna between locations,

with ranges of 5% (NSW) to 16% (Upper Spencer

Gulf), and 1% (Upper Spencer’ Gulf) to 21% (NSW)

respectively. These data suggest that the distribution

of feeding guilds varies according to geographical

location, Perhaps future studies should consider the

feeding category of fauna as well as their abundance,

productivity and biomass in order to estimate the

ecological importance of a species in shallow marine

systems, Edgar (1990) used size structure of benthic

macrofaunal communities to estimate faunal biomass

and secondary productivity but he did not assign the

fauna to feeding guilds which may be an additional

useful qualitative descriptor for benthic communities.

Posidonia’ sinuosa seagrass beds

P. sinuosa occurs from Shark Bay to the Great

Australian Bight (Larkum & den Hartog 1989).

However, the infauna of Posidonia sinuosa has only

been sampled at one other location in southern

Australia -— Albany in SW Western Australia

(Hutchings et al. 1991). The density of the Albany

infauna was twice that recorded from Upper Spencer

Gulf in this study. This difference between the two

faunas was largely the result of higher densities of

molluscs, crustaceans and ascidians at Albany (Table

5). Conversely, Upper Spencer Gulf has a considerably

increased species richness, the result of an increased

number of polychaete species in Upper Spencer Gulf

(Table 5). By comparison of the species lists in

Hutchings e2 al. (1991) and Appendix 1 of this paper

it is clear that there is. little similarity in the species

composition of the infauna between these two localities.

Therefore, as for P. australis, the intauna of P sinuosa

appears to vary substantially between locations.

Despite the different species composition there was

a considerable similarity in feeding types between

Upper Spencer Gulf and Albany (Table 6). Deposit

feeders, carnivores including scavengers, and

TaBLe 5. Posidonia sinuosu: distribution of the major phyla (S = number of species, d = overall mean density),

site Polychnetes Molluscs Crustaceans Echinoderms Astidians Minor Toral

Albany (Hutchin ge et al_ 1991}

$ 24 18* 7 16 1 106

a 7 259 6a 54 40 - 1797

Spencer Gulf (this study)

$ 37 3 4 2 5 4 192

a7 630 95 38 u u 300

* amphipods not identified to species

= densities {number

Gulf = 2,052 m’),

of individuals m~) calculated from total abundances and total area sampled (Albany = 1.2 m?, Spencer

Tame 6 Posidonia sinuosa: distribution of the fauna amongst feeding categories at Albany and Spencer Gulf (number of

species (%))-

location herbivore Suspension deposit camivore. omnivore

Albarty (from Hutchings et a/,, 1991)

polychaetes 3 (7) (3) 18 (39) 19 (41) -

molluscs (2 (50) 4 (17) 4 (17) 417) -

crustaceans - 1 vo m(s)) 6 (35) 9 (53)

ascidiang i4 (100) . - -

echinoderms - 1 (14) 3 (26) 2 (26) 2 (26)

totals 15. (14) 26 (24) 25 (23) 31 (29) b ()

Spencer Gulf (unis study)

polychaetes I4 (Ja) 16 (16) 38 (39) 28 (29) 1a)

molluscs 9 (28) 12 (38) 319) § (25) -

crustaccans (5) 22 (50) 20 (45)

ascidians = 5 (00) - : =

echinederms 2 17) 1 (8) 3 (25) 6 (50) -

tolals 25 (13) 36 (19) 66 (35) 62 (33) Luh

4 FP A. HUTCHINGS, T J. WARD, J. H. WATERHOUSE & 1. WALKER

Suspension feeders were jhe dominant feeding types

in P. sinwosa beds in both locations.

Concinsions

While species diversity in the subtidal seagrass beds

is higher in Upper Spencer Gulf thas in Albsny, it as

lower than in the seagrass beds of NSW. The density

of individuals is far lower in Upper Spencer Gulf than

elsewhere. and this may reflect the different

temperature and salinity regimes in Upper Spencer

Gulf compared to Albany and the aine locations along

the NSW coast, Collett et ai. (1984) concluded that

hydrological conditions were Important determinants

in the compostiion of the infauns of seagrass beds, and

more important than latitudinal effects. This study and

the study of Albany (Hutchings ef al, 1991) indicate

thal the conclusions of Collett er al. (1984) may also

hold for the broader geographical region of southern

Australia. Hydnmlogical conditions may also be

responsible for the difference between the Northerp

Spencer Gulf intertidal infuuna (Ainslic et af. 1989)

and the intertidal infauna of Upper Spencer Golf

observed in this study.

Acknowledgments

Field collectlons were funded by the International

Lead Zinc Research Organization, New York. Dr K.

Tiller at the CSIRO Division of Soils, Adelaide kindly

provided laborajory space. Vic Neverauskas, Rod

Grove-Jones, Nigel Holmes, Kim Horne, and Annette

Green assisted with field and laboratory work. Jurgen

Sieg of the University of Osnabriick identified the

tanaids. We appreciate assistance from the following

people from The Australian Museum with the

identifications: Helen Stoddart, Roger Springthorpe

(crustaceans), Frank Rowe (echinoderms), Geol?

Holloway (insects), Dug Hoese (fish), Jan Loch and

Winston Ponder (molluscs). We also appreciate the

assistance of Peter Davie (The Queensland Museum)

who idemified some of the decapods and Gary Poore

(The Museum of Victoria) whe identiled the isopods.

References

AINsuie, R.-C., JouNston, De A. & Orrcen, BE. W. (1989)

idal communities af Negthemn Spenter Gulf, South

Australia. Trans. R. Soc. 3. Anst. 13, 69-83,

BuLLock, D. A. (1975) The general water circulation of

Spencer Gulf, South Australia, in the period February to

May. bid. 99, 43-53.

CoLtett, L. C., HuTewinas, P. A., Ginss, P 3. & Cotums,

A.J. (1984) A comparative study of the macry-benthic fauna

of Posidonie australis seagrass meadows in New South

Wales, Aquatic Bot. 3B, 11-134.

Exar, G. £. (1990) The use of the size styucvare uf benthic

macrofaynal comfiunities to elucidate faunal biomass and

secondary production. J, exp. Mar. Biol, Ecol, 137, 5-214.

Howarp, R K., Bpoar. G, 1, & Hutcrmas, P. A. (1989)

Faunsl sssembluges of seagrass beds. pp 536-564, In

A. W. D. Larkum, A.l McComb, & S.A, Shephend, (Ede)

“Seagrasses ; A Treatise on the Bivlogy of Seagrasses with

apecial reference io the Australasian Region” (Elsevier,

North Holland}.

Hutcomes, PA. & Recver H. F (1974) The faume of

Careel Bay, with commeats on the ecology of mangrove

and seaerass communities Aust. Zool. IB, 99-123.

. FOR, Wetts, D, I. Watker, & G, A. Kenpeice,

(1991) Sengrass, sediment and inf4una — 2 comparison of

Fosidonia australis, Posidonia sinuove and hibolis

eatarctica in Princess Royal Harbour, south-western

Australia. 0, Distribution, composinon x abundance of

amacrofauna, pp, 611-633. fn F. E. Wells, D, 1. Walker, H.

Kirkman, & R. Letherbridge (Eds) “Proceedings of the

Third International Marine Biological Workshop: The

Marine Flora and Pauna of Albany, Westem Australia”. Vol.

2. (Western Australian Museum, Perth.)

Larkum. A. W. D. & ovw Harroc, C_ (1989) Evolution and

Biogeography of seagrass beds. pp.: 12-156, fe A, W, D.

Larkum, A.J. McComb, & S. A. Shepherd, (Eds)

“Seagrasses 4 Treatise on the Biology of Seagrasses with

spectal reference to Ihe Australasian Region”. (Elsevier, Nort

Holiand),

Nunes, R, A. & Lanwon, G. W, (1986) Physical propeny

Uistributions and seasonal trends in Spencer Gulf, South

Austmulia: an inverse estuary, Aust, J) Mar, Freshw, Res.

37, 34-53,

STEFANSON, R. (IS77) Spencer Gulf 9 review of the

oceanography, marine biology aud the implications for

development. pp. 13-16. Third Australian Conference oo

Caastal and Ocean Engineering. (Institution of Enganeers

Australia, Melbourne).

Watken, D. 1., Hurcaas, P A & Weis FE. (1991)

Seagrass, sxhunent and infauna- a rison of Fosidonis

australis, Posidania sinuose and Amphibolis antarctica in

Princess Royal Harbour, south-western Austrulia. [.

Seagrass biomass, productivity and contribution 1

sediments. pp. 597-610. Jn F. E, Wells, D- T. Walker, H.

Kirkman, & R. Letherbridge (Eds) “Proceedings of the

Third Enietnational Marine Biological Workshop: The

Marine Flora and Fauna of Albany, Western Australia’ Vol.

2. (Western Austrafian Museum, Perth.)

Warp, T, 3, (1987) Temporal variation of metals in the

seagrass Posidovic australis aid its potential as a sentinel

accumulator near a lead smelter. Mar, Biol. 95, 315-321.

& Youns P. C (981) Trace metal contammation of

shallow marie scdiments near’ lead smelter, Spencer

Gulf, South Australia. Aust. J. Mar: Fresh, Res, 32, 45-56.

&___ (1982) Effects of sediment trace metals and

Particle aiz¢ on lhe community structure of epiberthic

seagrass fauna near a lead smelter, South Australia, Mar,

Ecol. Prog. Ser. 9, 137-146,

WELLS, F. E,, Rose, R, A, & LANG, S. (1985) An analysis

of benthic marine invertebrate communities in subtidal

Seagrass aml yand habitats in Shark Bay, Western Australia

Rec. West. Aust. Mus. 12, 47-56,

INFAUNA OF MARINE SEDIMENTS AND SEAGRASS BEDS OF UPPER SPENCER GULF 9

Appendix 1 Summary of all data

Species Family Habitats

intertidal Zostera Posidonia Posidonia Sm bare 10m bare sum

australis sinuosa

Harmothoe sp. 1 Polynoidae 5 7 3 1 16

Harmothoe sp. 2 Polynoidae 1 1

Harmothoe sp. 3 Polynoidae 2 1 3

Paralepidonotus ampulliferus Polynoidae 1 1

Sigalion sp. Sigalionidae i 2 6 3 2

? Sthenelais sp. Sigalionidae 1 1

Chrysopetalum sp. Chrysopetalidae 1 1

Eumida sp. Phyllodocidae 21 18 15 5 59

Paranaitis sp. Phyllodocidae 1 4 5

Preryosyllis sp. | Phyllodocidae 2 2

Phyllodoce sp. A Phyllodocidae 49 y 4 6 6

Eteone n.sp. Phyllodocidae 9 5 1 L 16

Genetosyllis sp. Phyllodocidae | 1 2

Phyllodacid sp. 5 Phylladocidae I 4 3 8

Podarke tmicroantennata Hesionidae l 2 3

Padarke angustifrons Hesionidae ul 9 20

Typosyllis sp. 1 Syllidae 1 1 17 3 22

Exogone sp, Syllidae 2 2 4

Odantosyllis sp. 1 Syllidae. 2 2

Odontosyliis sp. 2 Syllidae 2 1 3

Pionosyllis ehlersiaeformis — Syllidae 10 14 2 2 28

Syilinae sp. 2 Syllidae 2 7 9

Typosyllis sp, 2 Syllidae 52 20 1 1 74

Ehlersia sp. Syllidae 15 22 1 39

Syllinae sp. 8 Syllidae 1 1

Ceratonereis mirabilis Nereididae 1 6 14 3 4 4 32

Neanthes sp. | Nereididae 1 1 7 9

Neanthes bassi Nereididae 10 15 3 2 13 43

Nereis bifida Nereididae 31 577 29 7 1 645

Neanthes kerguelensis Nereididae 6 9 36 3 74

Platynereis sp. Nereididae 65 5 2 1 73

Neanthes sp, 3 Nereididae 1 7 8

Neanthes vaalii Nereididae 45 i 1 1 49

Ceratocephela n.sp. \ Nereididae 1 L

Neanthes sp, 2 Nercididae 1 1

Ceratonereis sp. 1 Nereididae 248 8 34 I4 69 Il 384

Ceratonereis aequisetis Nereididae 4 4

Nereid sp. 20 Nereididae 2 1 3

Gymnonereis sp. 1 Nereididae 2 2

Nephtys gravieri Nephytidae 109 99 13 15 19 67 542

Glycera americana Glyceridae 3 2 5 5 3 18

Glycinde sp. | Glyceridae 31 12 9 4 3 59

Goniada maculata Goniadidae 1 |

Onuphid sp. | Onuphidae 1 B 21 1 36

Nematonereis unicornis Eunicidae 103 8 25 7 253

Eunicid juy sp. 1 Eunicidae I 1

Marphysa sp. 1 Eunicidae 1 1 2 3 34 41

Marphysa sp. 2 Eunicidae 2 oe) 1 2 2 22

Lysidice sp. 1 Eunicidae ! 5 6

Eunice sp. 1 Eunicidae 5 81 i9 4 109

Eunice sp. 3 Eunicidae 41 72 51 35 199

Eunice sp. 4 Eunicidae 7 6 2 IS

Eunice sp. 6 Eunicidae I 14 39 8 62

Eunice sp. 9 Eunicidae 3 3

Eunicid sp, U Eunicidae 1 1

Eunice sp. 5 Funicidae 5 2 6 B

Augeneria verdis Lumbrineridac 131 18 48 26 16 239

Lumbrineris sp. 1 Lumbrineridae 8 a 4 4 2 19

Arabella sp. | Arabellidae 9 8 ll 3 31

Schistomeringos sp. 1 Dorvilleidae 7 I 8

Schistomeringos loveni Dorvilleidae 19 16 6 16 7 64

Leitoscoloplos sp, 1 Orbiniidae 5 1 1 7

Phylo sp. 1 Orbiniidae 1 5 6

Naineris grubei australis Orbiniidae 4 6 10

10 PF. A. HUTCHINGS, T. J. WARD, J. H. WATERHOUSE & L. WALKER

Appendix 1 Summary of all data (continued)

Species Family Habitats

intertidal Zostera Posidonia Posidonia Sm bare 10m bare sum

australis —_ sinuosa

Scoloplos simplex Orbiniidae 1 74 17 4 4 l 101

Leitosceloplos normalis Orbiniidae 31 414 21 B 1 380

Family Orbiniidae Orbiniidac 17 1 1 9

Aonides oxycephela Spionidae 1 !

Spio sp. B Spionidae 2 1 3

Prionospio sp. 2 Spionidae 1 2 1 4

Polydora sp. 1 Spionidae 2 2

Pseudopolydora sp. 1 Spionidae 5 ] 4 10

Boccardia sp. 3 Spionidae | 1

Boccardia sp, 2 Spionidae 1 1

Aquilaspio multipinnulata Spionidae 7 1 9 ! 2 20

Aquilaspio aucklandica Spionidae 3 2 3 8

Spio pacifica Spionidae 1 1

Scolelepis sp. 2 Spionidae. 5 17 5 27

Pseudopalydora sp. 4 Spionidae 1 5 6

Malacoceros sp. | Spionidae 10 28 38 4 90

Scolelepis sp. 1 Spionidae 2 4 6

Spionidae sp. 2 Spionidae 1 ] 2

Spionidae sp. 3 Spionidae L ! 2

Boccardia sp. | Spionidae 1 1

Aricidea sp. 2 Pilargiidae 1 1 3 1 6

Magelona sp..1 Magelonidae 2 5 1 1 9

sp. 2 Cirratulidae 1 1

Caulleriella sp, 1 Cirranulidae 4 1 5

Caullertella sp. 2 Cirratulidae 1 20 4 31 6 72

Caulleriella sp. 3 Cirratulidae 7 35 43 14 1 100

Caulleriella sp. 4 Cirratulidae 20 3] 38 21 3 3

Cirriformia sp. 1 Cirratulidae 43: 43 28 4 9 127

Cirriformia sp. 2 Cirratulidae 29 43 3 5 2 92

Cirratulid sp. 1 Cirratulidae 3 1 4 1 9

Dipilacirrus sp. Flabelligeridac 92 3 4 1 2 102

Scalibregma inflatum Scalibregmatidae 5 2 ll 2 20

Hyboscolex dichranochaetus Scalibregmatidae 1 1

Armandia intermedia Opheliidae 10 51 5 8 74

Heteromastus filiformis Capitellidae 26 5 31

Capitella. ‘‘capitata*’ Capitellidae 53 608 661

Barantolla. sp, Capitellidae 88 57 88 46 6 285

Mediomastus n. sp. Capitetlidae 19 19 36 7 7 RB

?Scyphaproctus sp. 1 Capitellidae 1 L 5 1 8

Scyphoproctus sp. 2 Capitellidae 7 2 7 I 17

Notomastus sp. 1 Capitellidae 2. 96 51 25 2 186

Notomastus torquatus Capitellidae 4 4 29 16 63

Leiochrides sp. 1 Capitellidae. 3 1 4

Poechilochaetous serpens Poecilochaetidae Ll 2 8 ll

Asychis sp. Maldanidac 5 2 iS 19 1

Petaloproctus sp, Maldanidae 1 10 2 10 I 24

Praxillella sp. Maldanidae 1 2 2 5

Maldanid sp. 3 Maldanidae 3 3

Owenta fusiformis Oweniidae 5 3 8

Pectinaria sp. 1 Pectinuridae t 3 4

Isolda pulchella Ampharetidae 1 1 2

Ampharetid sp. 1 Ampharetidae ! 3 38 7 49

Ampharetid sp, 3 Ampharetidae 1 1

Lysilla pacifica Terebellidae i 4 7 26

Polycirrus sp. 1 Terebellidae 1 1 2

Polycirrus tesselatus Terebellidae 4 5 26 1 3 39

Polycirrus cf. nephrosus Terebellidae 1 1

?Decathelepus Terebellidae 1 1

Thelepus plagiostoma Terebellidae 1 2 3

Thelepus extensus Terebellidae 1 1

Streblosoma sp. Terebellidae 33 24 57

Lanussa exelysis Terebellidac 6 18 1 25

Lysilla laciniata Terebellidae 2 1 1 4

Pista australis Terebellidae 18 16 1l 6 122 63

Pista violacea Terebellidac 440 6 2 3 2 473

INFAUNA OF MARINE SEDIMENTS AND SEAGRASS BEDS OF UPPER SPENCER GULF Il

Appendix { Summary of all data (continued)

Species Family Habitats

intertidal Zastera Posidonia Posidonia Sm bare 10m bare sum

australis sinuosa

Neoleprea sp. Terebellidae 4 8 3 ah

Loimia ingens Terebellidae 2 2

Lanassa acellata Terebellidae 7 2 9

Eypolymnia koorangt Terebellidae 24 26 3 1 2 56

Nicolea amnis Terebellidac 2 7 1 10

Lanicides fascia Terebellidae 1 i

Amphitritinae sp. 3 Terebellidae 1 1 2

Amphitritinae sp. 4 Terebellidae 2 2

Terebellides stroemii Trichobranchidae 7 6 5 1 19

Trichobranchus sp. Trichobranchidae 1 4 4 1 10

Sabellid sp. 1 Sabellidae 37 1 3 91

Exchone variabilis Sabellidae 2 5 2 4 B

Laonome sp. 1 Sabellidace 2 2.

Sabellid sp. 3 Sabellidae 2 6 10 18

Sabellid sp. 4 Sabellidae ! I

Galeolaria sp. Serpulidae 3 3

Serpula sp. 2 Serpulidae 1 1

Serpula sp. | Serpulidae 1 1

Hydroides sp. 1 Serpulidae 10 4 7 4 l 26

total polychaetes 498 2590 1779 1292 1029 497 7685

Haliotis sp. Haliotidae 1 1

Amblychilepas nigrita Fissurellidae 16 1 17

Amblychilepas cf. oblonga _—Fissurellidae 1 1

Amblychilepas sp, Fissurellidae 1 1

Emarginula sp. Fissurellidae 1 1

Collisella sp. Lotiidae l 1

Naccula parva Lotiidae 5 10 15

Isoclanculus dunkeri Trochidae 2 3 4 9

Phasianotrachus cf. apicinus Trochidae 3 1 4

Thaliotia sp. Trochidae 1 1

Gena sp. Trochidae 1 1

Micrastrea rutidoloma Turbinidae 1 4 4 9

Phasianella australis Turbinidae 1 1 2

Bembicium vittatum Littorinidae 1 1

Triphorid sp, 1 Triphoridae 1 1 2

Zeacumantus diemenensis Potamididae 1 206 207

Bittium sp. 1 Cerithiidae 2 1 3

Bittium sp, 2 Cerithiidae 4 5

Diala. sp. Dialidae 1 3 L 5

Gazameda iredalei Turritellidae 2 1 3

Sabia conica Hipponicidae 1 1 2

Zeacrypia immensa Calyptraeidae T 7

Sigapatella calyptraeformis Calyptraeidae 3 3

Eunaticina umbilicata Naticidae 2 2 4

Ectosinum zonale Naticidae 1 1

Cominella eburnea Buccinidae 2 I 2 2 7

Cominella sp. Buccinidae 1 1

Nassarius pauperus Nassariidae 1 1

Fiusinus australis Fasciolariidae | 1 2

Deniimitrella cf. lincomensis Columbellidae 2 1 3

Splendrilla hurpularia Turridae 2

Conus anemone Conidae 1 1 2

Scaeoleda verconis Nuculidae 3 3

Solemya cf. australis Solemyidae i 16 17

Barbatia pistachia Arcidae. 2 2

Glycymeris flammea Glycymeridae it L

Glycymeris radians Glycymeridae & 8

Limopsis cf. tenisont Limopsidae 1 5 6

Limopsis sp. Limopsidae 5 5

Myrilus sp. Mytilidae 3 3

Austromytilus penetectus Mytilidae 3 Z 2 57 211 277

Trichomya hirsuta Mytilidae 3 3

Musculus cf. ulmus Mytilidae 2 2 1 4 6 15

Musculus cf. paulucciae Mytilidae 3 2 2 10 17

Modiolus albicostarus Mytilidae 1 1

2 P, A. HUTCHINGS, T. J. WARD, J. H. WATERHOUSE & L. WALKER

Appendix 1 Summary of all data (continued)

Species Family Habitats

intertidal Zostera Posidonia Posidonia 5m bare 10m bare sum

australis — sinuosa

Brachydontes erosus Mytilidae 8 3 1 12

Atrina tasmanica Pinnidae 1 I

Atrina sp. Pinnidae 2 2

Pinna bicolor Pinnidae 5 1 6

Electroma georgiana Pteriidae ul 5 2 3 3 24

Electroma sp. Pteriidae 2 2

Malleus meridianus Mallcidae 3 1 4

Valsella vulsella Malleidae 7 7

Equichlamys bifrons Pectinidae ] 2 3

Lima nimbifer Limidae | 1

Monia ione Anomiidae I 18 i9

Monia zelandica Anomiidae 4 4

Anomia descripta Anomiidae 2 2 4

Saccostrea *‘australis'' Ostreidae 1 3 3 19 26

Wallucina assimilis Lucinidae 4 60 2 5 4 is)

Cavitidens perplexa Lucinidae 4 29 I4 1 1 49

Myrtea bractea Lucinidae 1 1

Venericardia sp, Carditidae 1 1 2

Fulvia tenuicostata Cardiidae l 1

Chama ruderalis Chamiidae i 1

Dasinia histrio Veneridae 1 i

Circe rivularis Veneridae 1 1 3 5

Katelysia rhytiphora Veneridae 2 i 15 28

Katelysia sp. Veneridae | 3 1 7

Placamen flindersi Veneridae 13 3

Timoclea sp. Veneridac 1 1 3 l 6

Mactra sp. Mactridae 1 2 3

Téllina mariae Tellinidae 6 7 20 45 12 1g 172

Tellina deltoidalis Tellinidae 226 8 5 239

Tellina sp. 1 Tellinidae 1 1

Tellina sp, 2 Tellinidae 7 7

Semele monilis Semelidae 3 2

Laternula creccina Laternulidae il 48 2 6 67

Myadora complexa Myochamidae 1 2 4 7

Frenamya patula Pandoridac 1 1

Teredo Teredinidae 2 2

Bivalve 2 I 1

Bivalve 4 2 2

Chiton sp. 1 Ischnochitonidae 1 2 3

Stenochiton longicymba Ischnochitonidae 38 46 84

Stenochiton cf: pilsbryanus —_ \schnochitonidae 10 17 2 1 30

Chiton sp. 2 Ischnochitonidae 1 1

Chiton sp. 3 Ischnochitonidae 2 4 6

Dorid sp. 1 Dorididae 2 2

Dorid sp. 2 Dorididae 1 1

Dorid sp. 3 Dorididae 1 1

Philinopsis lineolata Aglajidae 1 I

Philine sp: Philinidae 1 1 2

Atys sp. Haminoeidae 1 1

Salinator sp. Amphibolidae 2 2

total molluscs 25 677 169 194 185 374 1624

Nebaliidae sp. 1 Leptostracan 19 6 25

Paguroidea sp, 1 3 1 l 4

Grapsidae sp. 2 Grapsidac 1 1

Hemigrapsus spinosus Grapsidae 1 31 2 2 1 37

Halicarcinus rostratus Hy menosomatidae 1 3 4

Halicarcinus ovatus Hy menosomatidae 2 2

Portunus pelagicus Portunidae 1 i

Ebalia intermedia Leucosiidae 3 3 6

Ebalia sp. A Leucosiidae 2 2

Macrobrachium intermedium Palaemonidae 9 2 2 13

Alpheus bidens Alpheidae 4 4

Alpheus euphrosyne

richardson Alpheidae 2 2

Alpheus novaezealandiae Alpheidae 2 2

INFAUNA OF MARINE SEDIMENTS AND SEAGRASS BEDS OF UPPER SPENCER GULF B

Appendix 1 Summary of all data (continued)

Species Family Habitats

intertidal Zostera Posidonia Posidonia Sm bare 10m bare = sum

australis sinuosa

Alpheus sp. Alpheidae 1 1

Metapenaeopsis novaeguineae Penaeidae 1 1

Apseudes australis Apseudidae 513 63 2 7 3 588

Apseudes sp. | Apseudidae 5 3 4 2 14

Apseudes sp. 2 Apseudidae 72 8 3 83

Kalliapseudes sp. Apseudidae 1 1

Tanais dulongi Tanaidae 26 819 1 846

Leptochelia ignota Tanaidac 1 1 2 4

Cymadusa sp, Amphithoidae 2 5 24 3 44

Ampithoidae sp. 1 Ampithoidae 2 2

Ampelisca toora Ampeliscidae 2 1 i 4

Byblis cf. bega Ampeliscidae 1 1

Pratolembos drummondae Aoridae 17 22 3 42

Xenacheira fasciata Aoridae 2 2

Bemlos strigilis Aoridae 1 i]

Ceradocus ramsayi Melitidae 1 1 2

Maera mastersi Melitidae 6 1 2 9

Maera sp. Melitidae 1 2 3

Ceradocus dooliba Melitidae 6 6

Ceradocus rubromaculatus —_ Melitidae 2 2

Ceradocus serratus Melitidae l 2 3

Cottesloe berringar Melitidae 6 1 7

?Parelasmopus s}). Melitidae 12 l 3

Melitidae sp. 1 Melitidae 1 1

Melitidae sp. 2 Melitidae I 1

Melitidae sp. 4 Melitidae 3 3

Leucothoe ?commensalis Leucothidae } L

Leucothoe assimilis Leucothoidae 1 l

Leucothoe sp. 2 Leucothoidae 1 1

Leucothoe sp. 1 Leucothoidae 2 2

Waldeckia sp. 1 Lysianassidae u 1

Waldeckia sp. 2 Lysianassidae 1 1

Waldeckia sp. 3 Lysianassidae 1 !

Parawaldeckia. sp. Lysianassidae 1 !

Tryphosella sp. Lysianassidae 1 1

Ischyroceridae sp. 1 Ischyroceridac 3 5 a

Phoxocephalidae spp. Phoxocephalidae 1 23 5 4 33

Atylus sp. Dexaminidae 4 4

Paradexamine lanacoura Dexaminidae l 1 2

Paradexamine ?linga Dexaminidae L 2 3

Paradexamine moorehousei Dexaminidae l L

Oedicerotid sp. 1 Oedicerotidac 1 ) 2

Eusirid sp. 1 Eusiridae 1 1

Eusirid sp. 2 Eurisidae 2 2

Eusirid sp. 3 Eusiridae l 1

Eusirid sp, 4 Eusiridae L 3 4

Hyale sp. Hyalidae 43 43

Tamituka doowi Platyischnopidae it 1

Natatalana wowine Cirolanidae 6 12 3 21

Platynympha longicaudata Sphaeromatidae 2 291 22 & 323

Cymodoce sp. 1 Sphaeromatidae 1 1

Cymodoce sp. 2 Sphaeromatidae 2 49 18 9 78

Cymodoce bidentata Sphaeromatidae 1 L

Limnoria sp. nov. Limnoridae 5 6

Euidotea peronii Idoteidae 4 4

Crabyzos longicaudatus Tdoteidae | 1

Euidotea bakert Idoteidae 7 7

Leptanthura diemenensis Paranthuridae ; 1 !

Acculathura bassi Paranthuridae 2 L 3

Mesanthura stypandra Anthuridae 1 ]

Amakusanthura olearia Anthuridae 3 l 4

Apanthura cf. isotoma Anthuridae 1 1

Haliophasma sp. 1 Anthuridac 3 3 3 1 10

Callianassa sp. Callianassidae 1 6 T

Gomeza bicornis Corystidae 2 4 6

4 P, A. HUTCHINGS, T. J. WARD, J. H. WATERHOUSE & L. WALKER

Appendix 1 Summary of all data (continued)

Species Family Habitats

intertidal Zostera Posidonia Posidonia 5m bare 10m bare sum

australis sinuosa

Ceratoplax punctata Goneplacidae 4 4

Litocheira bispinosa Goneplacidae 3 1 4

Actumnus setifer Pilumnidae 1 1

total crustaceans 33 1862 240 156 58 38 2387

Ascidiidae sp. 1 Ascidiidae 18 2 20

Ascidiidae sp. 2 Ascidiidac 3 1 4

Ascidiidae sp, 3 Ascidiidae 1 1 16 18

Ascidiidae sp. 4 Ascidiidae 5 1 6

Ascidiidae sp. 5 Ascidiidae 1 1

Ascidiidae sp. 6 Ascidiidae 2 2

Microcosmus sp. 1 Pyuridac 3 10 il 2 26

Didemnid sp. 1 Didemnidae 1 1

Pyura stolonifera Pyuridae 1 4 20 6 31

Polycarpa ?pedunculata Ascidiacea 4 4

total ascidians 0 25 2 22 32 22 113

Pentacta ignava Cucumariidae 1 I

Cucumella mutans Phyllophoridae 1 I

Thyone sp. Phyllophoridae 1 1

Scoliorhapis sp. ?nov. Chiridotidae 1 5 5 1 2

Trochodota shepherdi Chiridotidae 1 3 4

Leptosynapta dolabrifera Synaptidae 74 42 33 l 150

Order Synaptid Synaptidae 3 3

Thymio sycia Holothuriidae | L

Goniocidaris tubaria Cidaridae t 1

Temnopleurus michaelseni — Temmopleuridae i 3 4 18

Tosia australis Goniasteridae 1 3 4

Patiriella exigua Asterinidae 1 1

Amphipholis squamata Amphiuridae 5 2 3 1 i

Amphiura elandiformis Amphiuridae 1 1 23 25

Amphiura parviscutata Amphiuridae 10 4 2 20 46

Amphiura (Fellaria) sp. Amphiuridae 2 1 3

Amphiura trisacantha Amphiuridae 3 9 1 3

Amphiura sp. Amphiuridae I 10 15 1 27

Ophiocentrus pilosus Amphiuridae 3 I 4

Ophioconis opacum hiodermatidae 1 1

total echinoderms 1 80 74 78 28 66 327

Syngnathus phillipi Syngnathidae 3 3

Favonigobius lateralis Gobiidae 4 4

Pseudogobius alorum Gobiidae 2 2

Callogobius mucosus Gobiidae 5 5

Heteroclinus sp, 1 ]

Ophiclinops varius l i

Gymmapistes marmoratus Scorpaenidae 4 4

Platycephalidae 1 1

Vanacampus sp. i 1

Nemertean Nemertean 6 5 1 3 2 7

Enteropneust 5 5

Pycnogonid 2 2

Hirudinea 1 1

Sipunculan 1 10 4] 19 4 aN 96

insects 16 16

Porifera sp. 1 2

Porifera sp. 2 98 98

Porifera sp. 3 1 1

total minor taxa 17 128 55 22 18 20 260

Total 574 5362 2329 1764 1350 1017 12396

SEVENTEEN NEW SPECIES OF AUSTRALIAN BUPRESTIDAE

(INSECTA: COLEOPTERA) AND A HOST PLANT OF

CASTIARINA UPTONI (BARKER).

BY S. BARKER*

Summary

Sixteen new species of Castiarina are described: C. aeruginosa sp. nov., C. antia sp. nov.,

C. anthrene sp. nov. C. chrysothoracica sp. nov., C. crucianella sp. nov., C. hasenpuschi sp. nov.,

C. indigesta sp. nov., C. luteofusca sp. nov., C. markhanloni sp. nov., C. melasma sp. nov.,

C. mimesis sp. nov., C. nebula sp. nov., C. rayclarkei sp. nov., C. tenebrosa sp. nov., C. woodi sp.

nov. and C. xystra sp. nov.; and one new species of Themognatha, T. gordonburnsi sp. nov.

Dicrastylis georgei Munir is identified as a host plant of C. uptoni (Barker).

KEY WORDS: Coleoptera, Buprestidae, new species, Castiarina, Themognatha.

Transactions of the Royal Society of S. Aust. (1993), 1I7(1), 15-26.

SEVENTEEN NEW SPECIES OF AUSTRALIAN BUPRESTIDAE

(INSECTA: COLEOPTERA) AND A HOST PLANT OF CASTIARINA UPTONI (Barker).

by S. BARKER*

Summary

BARKER, S. (1993) Seventeen new species of Australian Buprestidae (Insecta: Coleoptera) and a host plant of

Castiarina uptoni (Barker). Trans. R. Soc. S. Aust. W7(1), 15-26 4 June, 1993.

Sixteen new species of Castiarina are described: C. aeruginosa sp. nov., C. antia sp. nov., C. anthrene sp.

nov. C. chrysothoracica sp. noy., C. crucianella sp. nov., C. hasenpuschi sp. nov., C. indigesta sp. noy., C. luteofisca

sp. nov., C. markhanloni sp. nov., C. melasma sp. nov., C. mimesis sp. nov., C. nebula sp. nov., C. rayclarkei

sp. nov., C. tenebrosa sp. nov., C. woodi sp. nov. and C, xystra sp. nov.; and one new species of Themognatha,

T. gordonburnsi sp. nov. Dicrastylis georgei Munir is identified as a host plant of C. uptoni (Barker).

Key Woros: Coleoptera, Buprestidae, new species, Castiarina, Themognatha.,

Introduction

The search for new species of Buprestidae occurring

irregularly in isolated areas has been continued, with

some success, by a group of dedicated amateur

entomologists. Included in this new material are

specimens of species rare in collections and some new

species which are described herein as are some new

species from older series which have been made

available to me.

The major gap in knowledge of this family in

Australia is information on their larval biology and food

plants. Castiarina uptoni (Barker) occurs in very

isolated arid areas of the country. The larval food plant

of the population which occurs in the Ashburton River

district in W.A. has been identified as Dicrastylis

georgei Munir.

Gardner (1989a) in a generic revision of the tribe

Stigmoderini pointed out that Polychroma Dejean held

date priority over Castiarina LaPorte & Gory, but had

not been used for over 130 years. She subsequently

applied to the ICZN to have the name Castiarina

conserved (Gardner 1989b) and that action has now

been taken (Opinion 1628, Bull. Zool. Nomencl. 48

(1) March 1991, pp. 74-75).

Materials and Methods

Collection date and locality information listed for

each specimen is a copy of all data written on each

individual label. Male genitalia were prepared and

displayed by the method described by Barker (1987)

referred to in the text and if previously published,

reference is given to the publication date. The

specimens illustrated in all species but two are the

holotype. Two allotypes are illustrated and the fact

noted in the appropriate remarks section. Measure-

ments given are mean total body length and width with

standard error, except where there are insufficient

* Department of Zoology, University of Adelaide, GPO Box

498, Adelaide, S. Aust. 5001.

specimens available to make the last calculation, when

only mean is given. Acronyms used in the text for

museum and private collections following the four letter

system of Watt (1979) are: ANIC Australian National

Insect Collection, Canberra; NMVA National Museum

of Victoria, Melbourne; SAMA South Australian

Museum, Adelaide; WAMA Western Australian

Museum, Perth; RCBA Mr R. Clarke, Byron Bay;

JHIQ Mr J. Hasenpusch, Innisfail; MHSA Mr

T. M. S. Hanlon, Sydney; MPWA Mr M. Powell,

Melville; RMNA Mr R, Mayo, Wallsend; ASSA, Mr

A. Sundholm, Sydney; GWQA, Mr G. Wood,

Atherton.

Castiarina crucianella sp. nov.

FIGS 1C, 2B

Holotype. &, 136 km NE Paynes Find, W.A.,

31.ix,1984, Jones & Powell, WAMA.

Allotype. 9, same data as holotype, WAMA.

Paratypes. W.A.: 200, 399, same data as

holotype, SAMA, MPWA.

Colour. Head, antennae and pronotum blue-grey.

Scutellum blue. Elytra orange with following blue

markings: narrow basal margin; premedial fascia not

reaching margin, reduced to elongate oblique mark in

some specimens; post-medial fascia reaching margin;

apical mark, all marks connected along suture. Ventral

surface and legs blue. Hairs silver.

Shape and sculpture. Head closely punctured, median

sulcus broad, muzzle short. Antennae, antennomeres:

1-3 obconic; 4-11 toothed. Pronotum closely punctured,

narrow basal fovea extending forwards to middle as

glabrous line, basal notches represented by glabrous

area on each side more marginal than medial; apical

margin straight, basal margin bisinuate; laterally

parallel-sided at base, rounded to widest pre-medially,

rounded and narrowed to apex. Scutellum scutiform,

glabrous, excavate, Elytra punctate-striate, intervals

convex, smooth, laterally angled out from base,

rounded at humeral callus, medially concave, rounded

post-medially and narrowed to bispinose apex; sharp

Fig. 1. Photomicrographs of male genitalia of the following Castiarina spp.: A. C. militaris (Carter); B. C. xystra sp.nov.:

C. C. crucianella sp.nov.; D. C. aeruginosa sp.nov., E. C. sanguinolenta (C & G); F.C. mitnesis sp.nov.; G. C. inconspicua

(Saunders); H, C. markhanloni sp.nov,; 1. C. antia sp.noy.; J. C. guttifera (Obenberger); K. C. tenebrosa sp.nov.; L.

C, hasenpuschi sp.noy., M, ©. Inieofusca sp.noy.; N.C. wood sp.noy.; O. C. nebula sp.noy.; P, C. anelis (Saunders):

Q. C. indigesta sp.noy.; R. C. rollei (Kerremans); 8. C. rayclarkei sp.nov.; T. C. chrysothoracica sp.nay. and Themognatha:

VU. T gordonburnsi sp-nov,

marginal spine, smaller sutural spine, margin rounded

and indented between spines, apices diverging slightly.

Ventral surface with shallow punctures, hairy, hairs

medium length, edges of abdominal segments glabrous.

S7; tuncate, medially indented in both sexes. Size.

Males, 4.7 * 13.1 mm (3). Females 5.2 * 13.4 mm (4).

Male genitalia. (Fig. \C) Parameres diverging from

basal piece, more so post-medially. rounded to apex.

Median lobe pointed, sides obtusely angled away.

Apophysis of basal piece medium width, rounded

apically.

Remarks. Resembles C. crux (Saunders) and C. nota

Barker. Differs from those species by being larger, with

bright blue markings and different male genitalia

(Barker 1990: Figs IK, IL). Name derived from crux

L., cross.

SEVENTEEN NEW SPECIES OF AUSTRALIAN BUPRESTIDAE

Castiarina aeruginosa sp. nov.

FIGS 1D, 2A

Holotype. ot, 17 km E Mt Carbine, Qld, 23.1.1991,

J, Hasenpusch & §, Barker, SAMA [ 21,241.

Allotype, 9 Brumby Gulley, Mareeba-Kuranda Rd,

Qld, 4.11981, 8. Barker, SAMA Lt 21,242.

Puratypes. Qld: lo LO. Mt Molloy, 26.x1).1985,

A Walford-Huggins, SAMA, W.A.. lor, 1 Q, 12 km

E Broome, 5.iv.1986, D. Knowles, MPWA,

Calour, Head coppery-yreen. Antennae purple.

Pronotum and sculletum coppery-green. Elytra pale

yellow with following green markings: narrow basal

margin, Variable post-medial spot at margin on each

elyiron; apical mark. Ventral surface; sternum

coppery-green, ubdomen pale yellow: Legs purple.

Hairs silver,

Shape and seulpture, Head closely punctured, median

Sulcus broad, muzzle short. Antennae, antennomeres;

1-3 obconic; 4-1 toothed, Pronotum closely punctured

laterally. less so medially, basal fovea, basal notches

more marginal than medial; apical margin projecting

medially, basal margin barely bisinuate; laterally

angled inwards trom base, rounded to Widest pre-

medially, rounded and narrowed to apex. Scutellum

scutiform, glabrous, flat. Elytura punctate-striate,

intervals convex, shallowly punctured; laterally angled

out frou base, rounded at humeral callus, medially

concave, rounded post-medially and narrowed to

bigpinese upex; small marginal spine. very small

sutural Spine, margin rounded and indented between

spines, apices diverging slightly, apical margin

subserrate. Ventral surface with shallow punctures

moderately hairy, hairs short, edges of abdominal

segments glabrous. S7: males truncate; females

rounded, slightly indented medially.

i’ ea

a, 5mm

Fig. 2, Habitus illustrations of the following Castiarina

specics A. €' -aerupinesa sp.nov. holotype; B.C

crucianella 6p,nov; holotype.

Size. Males, 4.3 % 11.2 min (4). Females, 5.)

13.5 mm (3).

Male genitalia. (Pig. 1D). Short and broad. Parameres

angled away from basal piece, more su post-mecdtally,

rounded to apex, Median lobe sharp, sides obtusely

angled away, Apophysis of basal piece mediune width,

rounded at apex,

Remarks. This species resembles. C. straminea

(Saunders) but distinguished by greenish reflections

of coppery markings and is smaller species, male

genitalia differ (Barker 1986: Fig. 2N) and has northern

distribution in Qld and W.A.. C. strained occurs in

central and southern Qld and torthern N.S.W, Name

derived from aeravo L., yerdigris.

Castiarina chrysotheracica sp. nov.

FIGS IT, 4B

Hoelorype. a, Mt Carbine, Old, 15.7,)991, R. Clarke,

SAMA £ 21,243.

Allorype, 9 . same data as holotype, SAMA I 2),244.

Paratypes. Qld: 299. Mt Carbine, #1.1992,

R. Clarke, RCBA: Let, Mt Garnet, 94.1992,

R. Clarke, MHSA; 1c", same data ax holotype, RCBA;

20 07, Mt Carbine, 16.1.1991, R. Clarke, ROBA; Ic,

Mt Carbine, 8.1.1992, R. Clarke. RMNA,

Colour. Head and antennae green with yellow

reflections, Pronotum green with volden reflections.

Scutellum yreen with yellow reflections. Elytra yellow

with the following markings: blue-green narrow. basil

margin and medial V-shaped mark surrounding

scutellum; blue post-medial fascia reaching margin:

blue apical mark, last two marks narrowly connected

along suture. Ventral surface: sterum green with yellow

reflections; male abdomen testaceous; female abdomen

green except apex S7 testaceous. Legs green. Huirs

silver.

Shape.and sculpture. Head clasely punctured, median

sulcus broad, muzzle short. Antennae, antennomeres:

1-3 obeonic; 4-11 toothed. Pronotum closely punctured.

narrow basal fovea extending forwards to middle as

glabrous line, basal notches more marginal than

medial; apical margin slightly projecting medially,

basal margin almost straight; laterally angled outwards

from base, rounded pre-medially. tapered to apex,

Scutellum scutiform, glabrous, excavate. Elytra

Punctate-striate, intervals convex, punctured; laterally

angled out from base, rounded at humeral callus,

medially concave, rounded post-medially and narrowed

in bispinose apex: sharp marginal spine, smaller sutural

spine, margin rounded and indented between spines.

dpices. diverging, apical margin subserrute. Ventral

surface with shallow punctures, maderately hairy, hairs

short, edges of abdominal segments glabrous, $7.

rounded in both sexes.

Size. Males, 14.3 + 0.24 » 3.5 4 O12 mom (A),

Femiles, 13.5 * 5.1 mmm (3).

18 S. BARKER

Fig. 3. Habitus illustrations of the following Castiarina

species: A. C. rayclarkei sp.nov. holotype; B. C. indigesta

sp.nov. holotype.

Male genitalia. (Fig. 1T) Parameres slightly angled

outwards from basal piece, slightly rounded medially,

rounded at apex. Median lobe sharp, sides obtusely

angled away. Apophysis of basal piece short and broad,

rounded apically.

Remarks. Colour and pattern of male resemble those

of C. garnettensis (Barker) but that species has

spineless apex and male genitalia are different (Barker

1989: Fig. 1G). Name derived from chrysos Gr., gold

and thorax Gr., chest.

Castiarina melasma sp. nov.

FIG. 4D

Holotype. 9 , Milmerran, Qld, 1.xii.1990, R. Clarke,

SAMA I 21,245.

Paratypes. Qld: 29 9Q, Milmerran,

R. Clarke, RCBA,

Colour. Head, antennae, pronotum and scutellum green

with yellow reflections. Elytra yellow with black apical

mark. Ventral surface and legs green. Hairs silver.

Shape and sculpture. Head closely punctured, median

sulcus narrow, muzzle short. Antennae, antennomeres:

1-3 obconic; 4-11 toothed. Pronotum closely punctured,

basal fovea extending to middle as impressed line, basal

notches on each side more marginal than medial; apical

margin projecting medially, basal margin barely

bisinuate; laterally parallel-sided at base, angled

inwards then rounded to widest pre-medially, rounded

and narrowed to apex. Scutellum narrow, scutiform,

glabrous, excavate. Elytra punctate-striate, intervals

convex, flat, punctured; laterally angled outwards from

base, rounded at humeral callus, medially concave,

rounded post-medially and narrowed to bispinose apex;

small, sharp marginal spine, very small sutural spine,

margin rounded and indented between spines, apices

hardly diverging, apical margin subserrate. Ventral

surface with shallow punctures, moderately hairy,

sternal hairs medium length, abdominal hairs very

short, edges of abdominal segments glabrous. S7: male

unknown; female truncate and indented medially.

Size. Females, 13.7 x 5.3 mm (3).

7.xii.1990,

Remarks. Structure, colour and pattern distinct. Name

derived from melasma Gr., a black spot.

Castiarina indigesta sp. nov.

FIGS 1Q, 3B

Holotype. & , Kuranda, Qld, 2.ii.1992, J. Hasenpusch,

SAMA I 21,246.

Allotype. 9, Davies Creek, Mareeba, Qld, 18.i.1991,

S. Barker, SAMA I 21,247.

Paratypes. Qld: 200, 19, Kuranda, i.1948,

G. Brooks, ANIC; 50° 0", 1 9, Kuranda, F. P. Dodd,

SAMA; 1c’, Kuranda, 10.i.1980, G. Wood, RMNA;

19, Mt Molloy, ii.1987, G. Wood, RMNA; lo". Mt

SEVENTEEN NEW SPECIES OF AUSTRALIAN BUPRESTIDAE 19

Fig. 4. Habitus illustrations of the following Castiarina

species: A. C. xystra sp.nov. holotype; B.C.

chrysothoracica sp.noy. holotype; C. C. antia sp.nov,

allotype; D. C. melasma sp.nov. holotype.

Garnet, 19.1.1989, S. Lamond, MHSA; Lo", Herberton,

1.1989, S. Lamond, MHSA; 20° 0", 19, Mt Garnet,

13.1.1991, R, Clarke, RCBA; 10°, 19, Mt Garnet,

7.1.1992, J. Hasenpusch, JHIA; to, 19, Mt Garnet,

9.1.1992, R. Clarke, RMNA; Lo, 19, Mt Garnet,

10,i.1992, R. Clarke, RCBA; Io", 19, Mt Garnet,

10.1.1992, R. Clarke, MHSA; I[o', Davies Creek,

18.1.1992, J. Hasenpusch, JHIA; 10, Kuranda,

22.1.1992, J. Hasenpusch, JHIA; 19, Windsor

Tableland, 2.ii1.1992, J. Hasenpusch, JHIA.

Colour. Head black with blue reflections. Antennae

blue-green. Pronotum black medially (narrowly at

apex, broadly at base) laterally red-brown, Scutellum

black. Elytra reddish-brown with following markings:

narrow dark blue basal margin; black sutural mark

commencing medially, reaching apex expanded as

black apical mark. Ventral surface and legs dark blue.

Abdominal segments in some male specimens all

brown. Hairs silver.

Shape and sculpture. Head closely punctured, median

sulcus, muzzle short. Antennae, antennomeres: 1-3

obconic; 4-11 toothed. Pronotum shallowly punctured,

deep basal fovea extending forwards to apical margin

as glabrous impressed line, basal notches more

marginal than medial; apical margin projecting

medially, basal margin bisinuate; laterally angled

outwards from base, rounded to widest part pre-

medially, tapered to apex. Scutellum cordiform, few

punctures, glabrous, flat. Elytra punctate-striate,

intervals convex, wrinkled and punctured; laterally

angled out from base, rounded at humeral callus,

medially concave, rounded post-medially and narrowed

to truncate spineless apex; apices diverging, apical

margin subserrate. Ventral surface with shallow

punctures, sparse short hair, edges of abdominal

segments glabrous. S7 rounded in both sexes.

Size. Males, 16.8 + 0.32 x 6.4 + 0.10 mm (19).

Females, 19.1 + 0.26 X 7.6 + 0.17 mm (10).

Male genitalia. (Fig, 1Q) Parameres slightly angled

outwards from basal piece, rounded post-medially and

narrowed to apex. Median lobe sharp, sides obtusely

angled away. Apophysis of basal piece short, medium

width, rounded apically.

Remarks. This species was confused with C. analis

(Saunders) a smaller species occurring further south.

C. analis has a prominent red margin around the elytra

and pronotum, also their male genitalia are distinct

(Fig, IP) Name derived from indigestus L., confused.

Castiarina xystra sp. nov.

FIGS. 1B, 4A

Holotype. & , Black Mt, A.C.T., iii.1931, T. G., ANIC.

Colour. Head light blue apically, dark blue basally.

Antennae blue-green, Pronotum dark blue medially,

red laterally. Scutellum blue. Elytra red with following

blue markings: narrow basal margin, medially

expanded into large rounded mark; sinuate fascia

covering humeral callus, thin red strip on each side

separating last two marks; post-medial fascia expanded

anteriorly and posteriorly along suture; apical mark.

Ventral surface: sternum predominantly blue medially,

presternum red laterally with blue spot each side close

to margin, presternal process red, meso- and

metasternum red medially; third coxae blue basally,

red apically, abdomen predominantly red, bases of

segents 5, 6, 7, 8 mainly blue. Legs blue. Hairs silver.

Shape and sculpture. Head closely punctured, median

sulcus deep, muzzle short. Antennae, antennomeres:

1-3 obconic; 4-1] toothed. Pronotum closely punctured,

basal fovea extending forwards to middle as impressed

line, basal notches on each side more marginal than

medial; apical margin projecting medially, basal

margin almost straight; laterally angled outwards from

base, rounded post-medially to apex. Scutellum

cordiform, punctate, excavate. Elytra punctate-striate,

intervals convex and punctured; laterally angled

outwards from base, rounded at humeral callus,

20 5, BARKER

medially Concave, rounded post-medially and numowed

to spineless apex; apices hardly diverging. Ventral

surface With shallow punctures, moderately hairy, hairs

long on stemun, medium length on abdomen, edges

of abdominal segments glabrous, S7: male truncate:

female unknown,

Size, Male, 15,6 % 60mm ()

Male venittalia, (Pig. 1B) Parameres angled outwards

trom basal picee, rounded ul apex. Median lobe sharp,

sides woutely angled away, Apophysis of basal piece

narrowed and rounded apically.

Remarks. The elytral pattem is similar wo that found

in C\ militariy (Carter) but ihe basal colours ditfer

being red in C. vysiwa and yellow in ©. militaris.

C.. tystva is a larger species and male genitalia ure

different (Fiz YA), Name derived from xysrra 1...

scraper,

Castiarina rayclarket sp. nov.

FIGS 1S, 3A

Heloivpe. (t, Acaciw Plateau, NS.W,, 16.1,1992,

R. Clarke, SAMA J 21.248.

Allotype. @, Avdeia Plateau, N.S.W.,

R. Clarke, SAMA T 21,249.

Paratypes. Old: 19, Warwick, 26.17.1992, R. Clarke,

MHSA4, NSW: 19. Acacia Plateau, 711.1992.

R. Clarke, RCBA,

Colour Head and antennae bright green. Pronotum:

bright gteen medially and along anterior margin and

base; red laterally. Elytra yellow with following black

markings wilh green reflections: narrow mark along

suture from middJe to apex covering both spines.

Ventral surface and legs bright green. Huirs silver.

Shape and sculpture, Head shallowly punctured,

glabrous, median sulcus, muzzle mediam. length,

Antennae, antennomeres: +3 obconic: 4-f toothed.

Pronotum shallowly punctured, glabrous, basal fovea;

apical margin projecting medially, basal margin

bisinuate, [aterally parallel-sided at base, rounded

medially to apex, Povea-on each side at basal angles.

Scutellum scutiform, glabrous, flat. Elytra puncrare-

striate, inleevals corivex and smooth; laterally angled

outwards from base, rounded at humeral callus,

ticdiatly parallel-sided, rounded post-medially and

narrowed W) bispingse apex, small marginal spine,

lager sutural spine, apices diverging. Ventral surtace

with shallow punctures, sparse short hair. edges of

abdominal segments glabrous. 57: rounded in both

SEXES.

Male genitalia, (Fig, WS) Parameres angled outwards

from basal piece. rounded near apex. Median lobe

sharp, sides acutely angled away. Apophysis of basul

piece medium width, rounded apically.

Size. Male, 184 * 67 mm (i). Females, 20.6 *

3.0 mm (3),

V1.1992,

Remarks. Vhis species shows morphological

similarities to C. rollei (Kerremmans). It diflers in

colour, size and male genitalia (Pig. IR) from that

species, Named after Mr Ro Clarke, Byron Bay.

Castiarina antia sp. now,

FIGS IL, 4c

Holotype, Of. Auairuding, W.A,, 23. 4i1,1997,

M. Powell & D, Knowles, SAMA I 21,250,

Allorype. GY. 38 km N Binnu, W.A., 13. xi. 1988,

M. Peterson, WAMA,

Colow. Head, antennae, pronatumn, scutellum ventral

surtace and legs purple-bronze, Elytra yellow with

following dark blue markings: holotype with narrow

basal margin, pre-medial fascia represented by medtal

spot on each elytron (allotype has broad fascia reaching

margin); broad post-medial fascia teaching maryan;

mark covering apex, Ventral surface and legs purple-

bronze, Hairs silver

Shape and sculpture. Head closely punctured, medrin

sulcus broad, muzzle very short, hairy. Antennae

compressed, antennomeres: L3 obconic; 4'4-toothed;

5-10 touthed. Pronotum closely punctured, basal fovea

extending forwards to middle as impressed line: apicul

margin projecting medially, basal margin bisinuate;

laterally angled inwards tront base, rounded to widest

pre-medially, rounded to apex, taterally hairy,

Scutellum seutiform, glabrous. flat. Elytra punctate-

sinate, iitervals Convex, punctured: laterally angled

out from base, rounded at humeral callus, medially

concave, rounded post-medially, rounded abruptly to

bispinose apex: small blunt marginal spine, minute

sutural spine, margin rounded and indented between

spines, apices diverging; Ventral surtace with shallow

punctures, hairy. hairs long, edges of abdominal

segments glabrous. $7: males truncate, medially

indented; females rounded.

Male genitalia. (Fig. (1) Short. Parameres parallel-

sided at base, rounded outwards. pre-medially, rounded

to apex. Median lobe blunt, sides acutely angled ayay.

Apophysis of basal piece narrow, rounded apically

Size. Male, 3.3 10.2 mm (1). Female. 3.3 © 10:7 mm

(1).

Remarks, Only Custiarina species Unat has reduced

number of abtendomeres. The specimen illustrated is

the allolype (Fig. 41). the first specimen collected.

Name derived from «aria L.. forelock.

Castiarina anthrene sp, nov.

FIG. 6E

Holarype. 9, 7 km W Nalbarra HS (28.398. 117,36).

W.A,., 24-30 111.1981, T. FE Houston, WAMA,

Paratype W.A.: | 9 same data as holotype, SAMA.

Celouw, Head, antennae black with blue reflections.

Pronotum black medially, blue laterally, Scntellum

SEVENTEEN NEW SPECIES OF AUSTRALIAN BUPRESTIDAL al

Nig. 5, Habitus illustrations of the following Caytiarina

species: A, OC) luteefiasca sp.nov, holotype; B. C, weed)

sp.noy. holowpe. CC fenebrosa sp-noy, holotype.

black, Elytra yellow with following black nuarkings:

narrow basal margin; pre-media fascia not reaching

margin anteriorly, extending to margin posteriorly;

post-tuedial fascia reachiny margin; apical mark, last

three marks connected along suture. Ventral surface

and legs dark blue. Hairs silver.

Shape und seulprire. Head closely punctured, median

sulcus, very Short muzzle. Antennae compressed,

antennomeres: 1-3 obeonic, 444 toothed; 5-LL toothed.

Pronotum closely punctured, basal fovea extending

forwards to middle as glabrous line, basal notches

represented by glabrous area on each side more

murginal than medial, connected by glabrous posterior

margin; apical margin projecting medially, basal

margin alinest straight; laterally parallel-sided at base,

rounded fo widest medially, rounded to apex. Scutellum

scutiform, glabrous. excayate, Elytra costate, intervals

3, 5, 9 raised at basal half wrinkled and punctured;

laterally angled out from base, rounded at humeral

callus, medially concave, faintly rounded post-

medially, wpered, potinded to bispinose apex; small

sharp marginal and sutural spines, margin rounded and

indented between. apices slightly diverging. Ventral

surface with Shallow punctures, moderately hairy. hairs

long, edges of abdominal segments glabrous. $7: mule

unknown; fetiales truncate, medially indented.

Size. Females, 1.0 * 3.6 mm (2).

Remarks. This is a very distinctive elongate species

showing the typical modifications found in wasp/bee

mimics. Name derived from anthrene, Gr., wasp.

Castiarina markhanloni sp. nov,

FIGS 1H, 6C

Holotype, o , Round Hill Reserve, N.S.W., 10.1992,

T. M, 8. Hanlon, SAMA 1 21,251,

Allotype. 9 , same data as holotype. SAMA J 21.252,

Paratypes. N.SW.; Boro. 109 9, same data as

holotype. MHSA.

Colour, Head, antennae, pronotum bronze-green with

purple reflections. Scutellum purple: Elytra pale yellow

with following dark blue markings with purple and blue

reflechons: narrow basal margin, pre-medial fascia

ends expanded anteriorly over humeral callas and

posteriorly touching margin; post-medial fascia

touching margin; mark covering apex, all marks

connected along suture. Veniral surface and legs

purpie-bronze with purple reflections. Hairs silver.

Shape and sculpture. Head closely punctured, shallow

median sulcus, short muzzle. Antennae, antennomeres;

13 obconic; 44 toothed; 5-11 toothed. Pronotum

closely punctured, punctations small medially larger

laterally, basal fovea extending forwards to middle as

glabrous line then to apical margin as impressed line;

apical margin projecting medially, basal margin

bisinuates laterally rounded from base, widest pre-

medially, rounded to apex, laterally hairy. Scutellum

small, scutiform, glabrous, Nat. Elytra punctate-striate,

intervals corivex, wrinkled, punctured; laterally angled

outwards from base, rounded al humeral callus,

medially concave. rounded post-medially and narrowed

to bispinose apex; small blunt marginil spine, sutural

spine minute, margin in most specimens almost

truncate and indented between spines. apices diverging.

Ventral surface with shallow punctures. long dense

hair. edges of abdominal segments glabrous. $7:

truncate both sexes.

Size. Males, 9.7 + O18 * 3.5 + 0.05 mm (14)-

Females, 10.2 + 0.22 * 3.9 + 0.09 mm (IN).

Male genitalia. (Fig. |H) Short. Parameres parallei-

sided from basal piece, angled outwards, pre-medially

rounded then tapered, rounded at apex. Median lobe

blunt, sides acutely angled away then widened.

Apophysis of basal prece medium width, rounded

apically.

Remarks. This species could be confused with

C. incvonspicua (Saunders). Main differences are the

smaller scutellum and the strongly rounded pronotum.

The male genitalia show some similarity in form

22 S. BARKER

Fig. 6, Habitus illustrations of the following Castiarina

species: A. C. nebula sp.nov. holotype; B. C. hasenpuschi

sp.nov, holotype; C. C. markhanloni sp.nov. holotype; D.

C. mimesis sp.nov. holotype; E. C. anthrene sp.nov.

holotype.

although those of C. markhanloni are narrow and the

apophysis of the basal piece is more tapered (Fig. 1G).

Named after Mr T. M. S. Hanlon of Sydney.

Castiarina mimesis sp. nov,

FIGS IF, 6D

Holotype. o , Badjalling, W.A., 7.xi.1970, 8, Barker,

SAMA 121,253.

Allotype. 9 , same data as holotype, SAMA 121,254.

Paratypes. W.A.: IS@o, 599, same data as

holotype SAMA; 2.9 9, South Tammin Flora Reserve,

8.x1.1970, S. Barker, SAMA; 10, 70 km S Perth,

Albany Highway, 19.xi.1970, S. Barker, SAMA; 20°",

59 9, Northam, C. G. Jessup, SAMA; 49 9, no data

SAMA; 19, 34 km N Gin Gin, I1.xi.1990, M. Powell,

MPWA,

Colour. Head, antennae, pronotum bright green.

Scutellum blue-green. Elytra orange with following

blue markings: narrow basal margin; vitta on each

elytron from humeral callus to pre-apex; line along

suture meeting vittae at pre-apex. In some specimens

vittae shortened at basal end leaving narrow mark over

each humeral callus. Ventral surface and legs bright

green. Hairs silver.

Shape and sculpture. Head closely punctured, medium

sulcus, muzzle short. Antennae, antennomeres: 1-3

obconic; 4/4 toothed; 5-11 toothed. Pronotum closely

punctured, basal fovea extending forwards to apical

margin as impressed line; apical margin projecting

medially, basal margin almost straight; laterally

parallel-sided from base, rounded post-medially to

apex. Scutellum scutiform, glabrous, excavate, Elytra

punctate-striate, intervals convex, heavily punctured;

laterally angled outwards from base, rounded at

humeral callus, medially concave, rounded post-

medially and tapered to bispinose apex; very small

marginal and sutural spines, margin rounded and

indented between spines, apices hardly diverging.

Ventral surface with shallow punctures, hairy, hairs

long, edges of abdominal segments glabrous. $7:

truncate both sexes.

Size. Males, 8.9 + 0.19 x 2.8 + 0.07 mm (19).

Females, 8.9 + 0.21 x 2.9 + 0.08 mm (18).

Male genitalia. Parameres angled outwards from basal

piece, rounded post-medially then increasing in width,

notched apically. Median lobe blunt, sides acutely

angled away. Apophysis of basal piece broad, rounded

apically.

Remarks. This species was confused with C. sang-

uinolenta (C & G ). Their elytral colour and markings

are very similar but male genitalia are very different

(Fig. IE) and they are easily distinguished by

differences in structure of their elytral spines. In C.

sanguinolenta these are sharp and equal, in C. mimesis

they are small and sutural spines are indented. They

occur together and are part of a Muellerian mimicry

complex, The name is derived from mimesis L.,

imitation.

Castiarina tenebrosa sp. nov.

FIGS 1K, 5C

Holotype. o, 4 km W Paluma, Qld, 7.i.1986,

E. E. Adams, SAMA I 21,255.

Allotype. 2, 4 km W = Paluma, Qld, 4.i.1986,

A, Sundholm, SAMA I 21,256.

Paratypes. Qld: 29 9 , same data as holotype, ANIC;

lor, 1 Q, same data as allotype, ASSA; 2, oo", 4km

W Paluma, 6/7.i1.1986, A. Sundholm, ASSA.

Colour. Head, antennae and pronotum black with green

reflections. Scutellum green. Elytra yellow with black

markings coalesced forming basal, pre-medial and sub-

apical spots, first two coalesced to form vitta also spot

on humeral callus covering margin. Ventral surface

green with yellow reflections. Legs dark blue. Hairs

silver.

Shape and sculpture. Head shallowly punctured, broad

median sulcus, short muzzle. Antennae, antennomeres:

1-3 obconic; 4'%-toothed; 5-11 toothed. Pronotum

shallowly punctured, small basal fovea; apical margin

straight, basal margin bisinuate; laterally parallel-sided

SEVENTEEN NEW SPECIES OF AUSTRALIAN BUPRESTIDAE 23

at base, rounded to widest pre-medially, narrowed to

apex, Scutellum cordiform, glabrous, flat. Elytra

punctate-striate, intervals convex, puncvared; laterally

angled outwards from base, rounded at husneral callus,

medially concave, rounded post-medially and narrowed

to bispinose apex; shatp marginal spine, small, sharp

sutural spine, margin straight between spines, apices

diverging. Ventral surface with shallow punctures,

moderately hairy, hairs medium length, edges of

atiominal segments glabrous. $7: rounded in both

sexes,

Size, Males, 11.9 + 012 x 4.2 + 006 mm (6),

Females, [6 * 4.4 mm (2)

Male genitalia. (Fig, 1K) Parameres angled outwards

fram basal piece, rounded post-medially, parallel-

sided, rounded tw apex. Median lobe sharp, sides

oblusely angled away. Basal piece wide, rounded

apically.

Remarks, This species was confused with C. guitifera

{Qhenberger) which has only been collected at

Kuramia, Qld, The male genitalia differ (Fig. LF) Name

derived from tenebrosus L., dark.

Castiarina hasenpuschi sp. a0v.

FIGS IL, 6B

Holorype. o . Mt Lewis, Qid, 61.1991, 1. Hasenpusch,

SAMA J 21,257,

Allatype, 9. sare dataas holotype, SAMA I 21,258

Furatypes, Qld: Lor, 29 9, same data as holulype,

JHIQ.

Colour, Head black with blue reflections, Antennae

black with blue-green reflections. Pronotum black with

purple reflections.. Scutelluis blue, Elytra yellow wah

following black markings: two fasciae and apical mark

coalesced leaving a pre-nedial, post-medial and pre-

apical yellow spot on each elytron and one on margin

at humeral callus. Ventral surface preen, Legs blue.

Hairs silver:

Shape end sculpture. Head closely punctured, median

sulcus, short muzzle. Antennae, antennomeres: 1-4

obaeanic, 5-11 toothed. Pronotum closely punctured,

narrow basal fovea; apical margin projecting medially,

basal margin bisinuate; laterally rounded from base,

widest pre-medially, rounded and narrowed to apex.

Scutellum cordiform, punctured, Mat. Elytra purictate-

striate, intervals convex, punctured; laterally angled

Gut front base, rounded al humeral callus, medially

concave, Tounded post-medially and narrowed to

bispinose apex; sharp marginal spine, small, sharp

sulural spine, margin straight between spines, aptoes

diverging, Ventral surface with shallow punctures,

moderately hairy, hair short, edges of abdominal

segments glabrous,

Males: legs 2 and 3 with reduced pulvilll on tarsomeres

1-3, replaced with single median spine, $7: male

trunctate, indented medially; female truncate, slightly

indented medially.

Size. Males, 9.8 * 3.8 mm (2). Females, 1.4 x 4.0

mm (3).

Male genitalia (Fig. 1L). Paraineres short and wedge-

shaped, Median lobe sharp, sides obtusely angled away,

Apophysis of bass] piece medium width, rounded

zpically,

Remarks, Structurally this species closest to members

of C. sexplagiata group except that elytra are smooth

whereas they are roughened with punctures in most

members of group, Named afier Mi J. Haseripusch,

Tanisfail.

Castlarina liteafusea sp. nov,

FIGS 1M, SA

Holotype &, Mt Lewis, Qld, 11.1992. G. A. Wood,

SAMA I 21,259.

Allorype, &, Mt Lewis, Qk, 6.11992, G. A. Wood,

SAMA I 21,260.

Paratype, 1o, Qhd: NQ., F P. Dodd, ANIC.

Colowr. Head green. Antennae, aptennomeres: 1-4

green; 5-li blue-green. Pronotum: bronze medially;

green apically. Scutellum green. Elytra yellow whh

black matkings coalesced leaving yellow basal and

medial spots on each elytron, these coalesced forming

angled vitta, round yellow apical spot and yellaw mark

on lateral surface of humeral callus. Ventral surface

green. Legs dark blue. Hairs silver.

Shape and sculpture. Head shallowly punctured, broad

median sulcus, short muzzle. Antennae, aliienmnomeres:

1-3 obconic; 4-11 wothed. Pronotum shallowly

punctured, small basal fovea; apical margin projecting

medially, basa] margin bisinuate, latenilly parallel-

sided ar base, angled inwards, rounded to apex, widest

pre-medially. Scutellun: scutiform, glabrous, flat.

Elytra punctate-strate, intervals convex, punctured;

laterally angled outwards from base, rounded at

humeral callus, medially concave. rounded post-

medially and tapered to bispinoase apex; sharp marginal

spine, smaller sharp sytural spine, margin rounded

between spines, Ventral surface with shallow

ponciures, few short hairs, edges of abdominal

segments glabrous. $7: trunctate in both sexes.

Size. Male, 12.4 x 4.7 mm (1). Females, 12.7 *

48 mm (2),

Male genitalia (Fig. IM), Parameres angled outwards

from, basal piece, rounded to apex. Median lobe

pointed, sides acutely angled away. Apophysis of hasal

piece medium width, rounded apicully,

Remarks. Largest of black and yellow C, producta

(Saunders) group mimics and with C. tenebrosa and

following new species, only ones that have yellow vitae

oo elytra. Name derived from /uteus L., yellow and

fuscus L,, black,

Castiarina woodi sp. nov.

FIGS IN, 5B

Hoterype. oF, Mi Lewis, Qld, 1.11992, G. A. Wood,

SAMA I 21,261

a4 S, BARKER

Alloype, 9. same data as holotype. SAMA 21,262.

Colowr, Head and antennae blue-green. Pronotum

green with bronze reflections. Scutcilum blue-green.

Elytra yellow with folkxwing dark green marks;

matkings coalesced to form yellow basal and medial

spols also coalesced forming long vitta covering

humeral callus and margin; large round pre-apical spot,

Ventral surface green. Legs blue-green, Hairs silver.

Shape and sculpture: Head shallowly punctured, broad

tmehan sulcus, short muzzle. Antennae, antennomeres:

1-3 obconic; 4-Ii toothed. Pronetum shallowly

punctured, deep basal foves; apical margin projecting

medially, basal margin bisinuatc, laterally parallel

sided at base, rounded to apex, wides! pre-medially,

Scutellum scutiform, glabrous, flat. Elytra punctare-

striate, intervals cortver, punctured; Jaterally angled

outwards from base, rounded at humeral callus,

medially concave, rounded post-meiially to unispinose

upex; sharp marginal spine, sutural spine represented

by very small notch, margin straight betweert spine and

fotch, apices diverging. Ventral surface with shallow

punctures, moderately hairy, hairs long and denser in

medial strip, edges of abdominal segments glabrous.

S7: truncate in both sexes.

Size. Male, 1.4 x 4.5 mm {l). Female, 11.9 «4.5 mm

(1).

Male genitalia, (Fig. IN) Pacameres more or less

paraliel-sided from basal piece, rounded to apex.

Median Jobe pointed, sides obtusely angled away.

Apophrysis of basal piece short, narrowed and rounded

to apex,

Remarks. Elytral markings of this species arc similar

tothese of C, luteofusea but distinctive male genitalia

and clytral spines distiguish this species. Named after

Mr & A. Wood, Atherton,

Castiarina nebula sp. nvy.

FIGS 10, 6A

Solerype, of, Mt Lewis, Gid, 6.11991, 3, Hasenpusch,

SAMA I 21,263

Atiaype. 9 , same data as holotype, SA MA 121,264,

Perarypes. Qhd: 20-0, Kuranda, 1.19.1990, 5,71, 1990,

G. A. Wood, GWAQ; 1¢, Kuranda, Qld, 2.1.1978,

A, & M. Walford-Huggins, SAMA; 29 9, Kuranda,

S.0.1990, J. Hasenpusch, JHIQ; 1¢, Kuranda,

141.1990, G, Wood, GWAQ,

Colour Head and amennae green. Pronotum black

with blue of green reflections, Scutellum green. Elyira

yellow with following black markings: fasclae and

marks coalesced leaving six yellow spots, Largest pre-

medial, smallest post-medial and intermedial on

margin at humeral callus. Veniral surface green. Legs

blue. Halrs silver.

Shape and sculprure, Head shallowly punctured,

median sulcus, medium teagth muzzle Antennae,

antenaomeres: t-3 oboome: 4/-loothed; 5-IL inethed.,

Pronotum shallowly punctured, very small basal fovea:

apical margin straight, basal margin bisinuate; laterally

angled inwands from base, then rounded from near base

to apex, widest pre-mecdially. Scutelhim scutiform, few

punerures, glabrous, flat. Elytra punctate-striate,

intervals convex, smooth; laterally angled outwatds

from base. rounded at humeral cullus, medially

concave, rounded post-medially and tapered 10

bispinose apex; sharp marginal spine, smull sharp

sutural spine, margin rounded and indented between

spines, apices diverging. Ventral surface with shallow

punctures, few short hairs, abdoniinal segments

glaheous. S7: truncate both sexes, Males: legs 2 and

3 with cach pulvillus om tarsomeres L-3 replaced by

median spine,

Size. Males, 10.9 * 3.8 mum (3). Females, IH x

4,2 mm {4},

Male genitalia (Fig, JO). Parameres widened from

base, rounded at apex. Median Jobe sharp, sides

obtusely angled away. Apophysis of basal piece

medium width, rounded at apex,

Remarks. Elyiral markings of this species are similar

to those of © actasignata (Carter, 1919). However

C. octosignato difters stucturally as elytral spines are

differem shape, Male C. octestenata Unknown. Name

detived from nefufosus L., dark.

Themognatha gordonburnsi sp. nov:

FIGS WW, 7

Holetype, ct. 69 km N Galena Bridge, WA,

29.ix.1992_ M. Golding & M_ Powell, WAMA.

Allotye. 2, 100 km N Murchison River, W.A,,

14.i%.1980, G. G. Burns, NM'VA,

Faratypes, WA.; 20-9, same datd as allvtype,

14,ix.1980, 17.ix.1980, G. G. Burns, NMYA: Sor cr,

same uatt as holotype, SAMA & MPWA,

Colour, Head and antennae black. Pronotum black with

yellow literal murgins, Scutellam black. Elyira

predominantly brown with yellow fateral tmargins and

following black markings: narrow basal margin;

females with faint pre-apical vitta over each humeral

callus-and faint markings on suture, post-medial tascin

aot reaching margin; spade-shaped pre-apical mark

extending over apex. Ventral surface: male predom-

inantly black pre-sternuns with yellow mark on meta-

slettal coxae; abdominal sclerites yellaw wih

testaceous edges; fernale black with yellow mark oa

meta-sternal caxae nd on lateral edges of all visible

abdominal segments. Legs black. Haars silver.

Shape ard sculpture. Head punctured, hairy, apex

medium length, Antennae, antennomeres: 1-3 oboonic,

4 semi-toothed; 5-11) toothed. Pronotum punctured,

apical margin projecting medially, basal margin faintly

sinuous, laterally rounded from base to apex, bulbous

pr:-medially. Scutellum almost circular, flat, without

punenues Elytra punctate-striate, intervals convex and

SEVENTEEN NEW SPECIES OF AUSTRALIAN BUPRESTIDAG 2S

5mm

Fig. 7 Habitus illustration of Themoynatha gordoaburnsi

spunoy, allorype

smooth; laterally angled outwards from base, rounded

at humeral callus, medially more or less parallel-sided,

rounded post-medially to spineless apex. Ventral

surface punctured, hairy. S7: trunctate, medially

indented in mule: rounded in female.

Male genitelia. (Fig. 1U). Parameres angled outwards

from basal piece, rounded at apex. Median Jobe blunt,

sides acutely angled away. Apophysis of basal piece

medium width, rounded apically.

Size. Males, 20.6 + 0.35 x 8&4 + 013 mm (10).

Females, 24.3 * 9:6 mm (3).

Remarks. Although thé locality data on the holotype

and allotype appear different they refer fo the same

area. The allotype has a narrow median yellow band

on the pronotum, not reaching base or apical margin.

T. burnsi is a spring emerging species, adults have been

collected on the Mowers of Grevillea sp, and Bursaria

spinosa, On the basis of its structure and male gemitalia,

it can be grouped with 7 oleata (Blackburn) u blue

and red species which emerges in late summer and

autumn and is usually associated with mallee Mowers.

The specimen illustrated ts the ullotype (Fig. 7), the

first specimen.collected, Named to commemorate my

friend and colleague the late Mr Gordon Burns,

Mornington,

A host plant of C, uptoni (Barker)

Although the adults of many Castiarina species are

commonly collected, the pre-adult stages and host

plants of even the commonest species are mostly

unknown. Castiarina upiuni (Barker, 1979) was

described from dead adults found inside tubular

highway: marker posts, north af Barrow Creek, N.T.

Presumably they had been dropped inte the cavities

by predaccous insects that had been feeding on them,

possibly asilid flies which commonly prey on

buprestids (Barker & Inns 1976). Live specimens were

subsequently collected in the same general area but

their food plants were not recorded. Several years ago

Herbert Demarz collected live specimens of C. uploni

on the leaves of Dicrasrylis georgei Munir, a very low

shrub growing in an ifterdune situation on Yanrey

Station, 4-31 km south of Barradale roadhouse on the

Great Northern Highway, W.A. On 10 August 1990,

I visited the locality with Mr Demarz and we examined

the same plants in that area, but found no adult beetles

on the leaves, Many of the stems were dead and

detached from the individual plants. Examination of

the live part of cach cut stem showed frass and when

dissected, every one was found to be galleried and to

conlain a living beetle. Most were adult, but a

proportion were newly pupated or in various stages

of post-pupal metamorphosis, OF fourteen specimens

dissected out, twelve were C. upteni and two were

C. quadrifasciata (Saunders, 1869). The twa species

belong to different species groups on basis. of structure

of their male genitalia, but are similar in hody colour

and pattern and thus form a Mullerian mimicry group.

When I visited the same area on 20 August 1990, miuny

C. uptoni adults were found on the leaves of the host

plant as well as two specimens of C. quadrifasciata,

As D. georgei does not occur in the N.T., beetles from

the type locality must have another host plant which

could be a second species of Dicrastylis, possibly

D, gilesii F, Muell. which occurs in that area (Jessop

1981).

Acknowledgments

1 thank the following for their assistance: Dr

K. Walker, NMVA; Mr T. A. Weir. ANIC: Ms

H. Vanderwoude, Department of Zoology, University

of Adelaide. 1 thank the following collectors for the

loan of specimens: Mr E, E. Adams, Edungalba; Mr

26 S. BARKER

As D. georgei does not occur in the N.T., beetles from

the type locality must have another host plant which

could be a second species of Dicrastylis, possibly

D. gilesii F. Muell. which occurs in that area (Jessop

1981).

Acknowledgments

I thank the following for their assistance: Dr

K. Walker, NMVA; Mr T. A. Weir, ANIC; Ms

H. Vanderwoude, Department of Zoology, University

of Adelaide. I thank the following collectors for the

loan of specimens: Mr E. E. Adams, Edungalba; Mr

R. Clarke, Byron Bay; Mr H. Demarz, Wanneroo; Mr

T. M. S. Hanlon, Ride; Mr J. Hasenpusch, Innisfail;

Mr A. Walford-Huggins, Yeppoon; Mr R. Mayo,

Wallend; Mr M. Peterson, Perth; Mr M. Powell,

Attadale, Mr R. I. Storey, Mareeba; Mr A. Sundholm,

Elizabeth Bay; Mr S. Watkins, Caparra; Mr G. A.

Wood, Atherton.

References

BARKER, S. (1979) New species and .a catalogue of

Stigmodera (Castiarina) (Coleoptera: Buprestidae). Trans.

R. Soc. S. Aust. 103, 1-23.

(1986) Stigmodera (Castiarina) (Coleoptera:

Buprestidae): taxonomy, new species and a checklist. Ibid.

10, 1-36.

(1987) Eighteen new species of Stigmodera

(Castiarina) (Coleoptera: Buprestidae) bid. M1, 133-146.

_______ (1989) Contributions to the taxonomy of Australian

Buprestidae (Coleoptera): new species of Astraeus and

Stigmodera (Castiarina) and a key to Astraeus (s.s.). Ibid.

113, 185-194.

(1990) New species of Castiarina (Coleoptera;

Buprestidae) and a redescription of C. diversa (Kerremans)

and C. elderi (Blackburn). Jbid. 114, 169-178.

& Inns, R. (1976) Predation on Stigmodera

(Themognatha) tibialis by a fly. WA. Naturalist. 13, 147-148.

Carter, H. J. (1919) Proc. Linn. Soc. N.S.W. 44, 137-173.

GARDNER, J. A. (1989a) Revision of the Genera of the Tribe

Stigmoderini (Coleoptera: Buprestidae) with a discussion

of phylogenetic relations. Invertebr. Taxon. 3, 291-361.

(1989b) Castiarina Gory & LaPorte, 1837 (Insecta,

Coleoptera): proposed conservation. Bull. Zool. Nomencl.

46(3), 170-172.

Jessop, J. (Ed.) (1981) ‘Flora of Central Australia? (A. H. &

A. W. Reed Pty Ltd, Sydney).

SAUNDERS, E. (1869) Descriptions of fifty new species of the

genus Stigmodera. J. Linn. Soc. 9, 460-483.

Wart, J. C. (1979) Abbreviations for entomological

collections. N.Z. Zool. 6, 519-520.

ADULT AND LARVAL STAGES OF PARAUSTROSTRONGYLUS RATTI

(NEMATODA: TRICHOSTRONGYLOIDEA) FROM RATTUS FUSCIPES

BY IAN BEVERIDGE* & MARIE-CLAUDE DURETTE-DESSETT

Summary

The morphology of the adult and the third and fourth larval stages of Paraustrostrongylus ratti from

the native rodent rattus fuscipes is described, with particular reference to the synlophe, as well as

the mechanism of attachment of the nematode to intestinal villi. The synlophe of the fourth larval

stage has an oblique axis of orientation, in contrast to the frontal orientation in the adult stage, and

most closely resembles the synlophe of species of Dessetostrongylus parasitic in dasyurid

marsupials. Ontogenetic data therefore suggest that Paraustrongylus evolved from an ancestor

resembling Dessetostrongylus.

KEY WORDS: Nematodes, Trichostrongyloidea, Paraustrostrongylus, ontogenesis, larvae,

morphology, rodents.

Transaartions of the Royal Society af S. Aust (1993), 17(L), 2736

ADULT AND LARVAL STAGES OF PARAUSTROSTRONGYLUS RATTI

(NEMATODA: TRICHOSTRONGYLOIDEA) FROM RATTUS FUSCIPES

by LIAN BEVERIDGE* & MARIE-CLAUDE DURETTE-DESSET+

Summary

Bevermeoe, 1, & Durerre-Desser, M,-C, (1993) Adult and larval stages af Paraustrostrong ylus raiti (Nematoda:

Trichoatrongyloldea) from Rattus fuscipes, Trans, R, Soc. $8, Aust, U7), 27-36 4 June, 1993,

The morphology of the adult and the third and fourth larval stages wf Paraustrostrongylus rari from the native

rodent Rastus fiscipes is described, with particular reference to the syntophe, as well as the tnechanis¢n of artechiment

of the nematode to intestinal villi. The synlophe of te fourth larval stage hasan ublique axis «vf orientation,

in contrast to the frontal orientation in the adult stage, and most closely resembles the synlophe of species of

Dessetostrangylus parasitic in dasyurnud mursepials. Ontogenctic data therefore suggest that Purausirongyus evolved

from an-ancestor resembling Dessetostrengylis.

Key Worps: Nematodes, Trichostrongyloidea, Paraustrostrongylus, ontogenesis, larvae, morphology, rodents.

Introduction

The trichostrongyloid nematode subfamily

Herpetostrongylinae occurs in the small intestines of

Australian marsupials and is one of the few

tichostrongylold groups in which there is apparently

a close evolutionyry parallel between hosts and

parasites (Durette-Desset 1982, 1985; Humphery-Smith

1983; Beveridge 1986). Three distinct lineages have

been recognised among the cight component genera

in Marsupials, each sharing a probable comron

ancestry with Woolleya, a contemporary genus which

occurs in dasyurid marsupials. Woolleva shares fearures

in common with the genus Viaraaia, belanging to the

family Viannaiidae, which is present in South

American marsupials and rodents (Humphery-Smith

1983).

Two of the herpetostrongyline genera,

Austrostroneylus and Parausrrostrongwus, occurring

in diprotedon! marsupials, with one species in the

marsupial mole, Norerveres typhlops, and one in a

rodent, Ratius fixectpes, are of particwlar morphological

interest due to the development of paired lateral

cuticular inflations, unique within the Trichostrangy-

toidea, termed “floats” by Durette-Desset (1979). The

evolutionary development of these floats was

investigated in Austreastronyylux and in a related penus,

Sutarostroneylus, (Beveridge & Durette-Desset 1986)

and species with intermediale or primitive

morphological features, that is either with a single float

or without floats, were identified. This study suggested

that Sutarostrongylus, parasitic in Thylogale spp.,

exhibited a number of primitive characters, ancestral

to those seen in Ausirosirongylus, but provided no

* De

Melbourme, Parkville, Vic, 3052.

' Laboratoire de biologie peresilaire, protstolovie et

hélminthologie Muséum national d"Histiare naturelle, él,

rue Buffon, Paris. France.

mmem of Veterinary Science, University of

additional insights into the possible relationships

between a postulated Mbolleya-like ancestor and

Sutarostrongylus. Desyetostrongylas is one possible

intermediary between Rbollepa and Sutarostrongylus

(ste Beveridge & Durette-Desset 1986) as it has a

synlophe. or camplement of body ridges, identical with

that of Siarestrongylus except for the fact that the axis

of oriettatian of the synlophe is objique in

Dessetostrongylus but frontal in Sutarastrangylus,

Humphery-Smith (1983) by contrast, placed greater

emphasis on the frontal onentation of the syntophe of

Austrestroneylus and derived it direcdy from an

ancestral stale resembling that found in Maolleye

sprenii.

Cassone er al, (1986), studying new species of

Rbolleya and Parricialinag from dasyurid marsupials

confirmed the dire Hbolleya - Austrostrongylus

relationship identified by Humphery-Smith (1983), but

considered that Dessetostrengylus was a sister group

10 Austrostrongylas.

All studies on the evolution of the Herpetostrongy-

linae to date have relied on the comparative

morphology of the adult nematodes and particularly

on differences in the anatomy and orentation of the

syniophe. The ontogenesis of larval stages is an

important source of phylogenetic information mn the

Trichostrongyloidea (Durette-Desset 1985), but has not

been exploited in the case of the Herpetostrongylinae

ather than in the case of Beveridgiella pearseni (see

Humphrey-Smith 1980), because no life cycles are

known.

We decided to investigate the morphology of the

various life-cycle stages of Paransrrostrangylus rat,

the anly member of the genus to occur in a eutherian

mammal (Obendorf 1979) to anempt tm obrain

additional information on the evolution of the synlophe

of the peous. Due to its abundance in Rartus fiuscipes

and the eage with which Infected rats could be obtained

and kept in the laboratory. # rutti was considered to

2x I. BEVERIDGE & M.-C. DURETTE-DESSET

z

7

ay

Ba

mel

jatar

Taw!

Ss

ae

Figs 1-15. Paraustrostrongylys ratti Obendorf: Adult. 1, anterior end, lateral view, dorsal aspect on left hand side; 2, anterior

end, dorsal view, 3, apical view of mouth opening and lips; 4 optical transverse section through hexagonal buccal capsule,

with dorsal tooth, 5, transverse optical section through anterior end of oesophagus: 6, anterior region, left lateral view;

arrows indicate origins of ridges; 7, anterior region, right lateral view: arrows indicate origins of tidges: 8, bursa, lateral

view; numerals indicate ray numbers according to Durette-Desset (1983); 9, bursa, ventral view; 10, gubernaculum, lateral

view; ll, spicule tips, ventral view; 12, spicule tip, lateral view; 13, genital cone, ventral view of papilla 0; 14, genital

cone, lateral view showing papillae 0 and 7; 15, fernale tail, lateral view. Scale Hines 0.01 mm: figs 1,2, 10, 1-14 to sane

scale; figs 3-5 to sarne scale; figs 6.7 to same scale; figs 8,9 to same scale. Legend: a, amphid; d, deirid; e, excretory

pore; 1, lip; p, posterior atrophic uterus; s, submedian papilla; sp, sphincter; 1, dorsal tooth; ve, vestibule:

LIFE CYCLE STAGES OF P RATT! 29

be mom suitsble fot investigation than species

occurring in marsupials, The morphological data

presemted here also provide (he basis for subsequent

ultrastructral studies.

Methods

Naturally infectex! rats, Rattus fuscipes (Waterhouse),

were trapped at Blackwood, Victoria (37°29’S,

144°19"E), killed im the laboratory and the small

intestine was divided into segments and opened in

warm 0.09% saline The itnestinal segments were

Placed im an incubator for two hours to allow

Neinstodes to migrare ins the saline. Nematodes were

then washed in saline and fixed in hot 70% ethanol,

Small numbers of nematodes were fixed in 2.5%

glutaraldehyde in phosphate buffer at 4°C.

Adult, fourth and parasitic third-stage nematedes

were cleared in Jactophenol and examined, using

Nomarski interference contrast microscopy. Transverse

sections of jhe trody of male and female nematodes

were cul using a cataract sealpel, mounted in

tactophenol for examinauon and orented using the

methods of Durette-Dessei (1971). Apical views of the

anleriar extremity were made by similar means,

Specimens fixed in glutaraldehyde were embedded in

resin, Sections cul ata thickness of |j.m were stained

with toluidine blue and were used to confirm

mormphelogiwal featires seen in hand-cut section,

Additional specimens were dehydrated in a praded

ethanol series, dried in a critical point drier, coated

with pold and examined with a Siemens Autoscan

scanning electron microscope.

Ridges of the synlophe were numbered in an

anticlockwise fashion beginaing with the lefi-ventral

ridge, in order to demonstrate homologies between

stages. The numbering system for the ursal rays and

papillae: follows that of Duretle-Desset (1985),

Faeces from naturally infected rats were collected,

mixed with an equal quantity of charcoal and cultured

on moist filter paper in Petri dishes at laboratory

temperature. Frve and eight days later, larvae emerging

from the faccal-charcoal mixcure were collected in

distilled water and concentrated by sedimentation.

Third-stage larvae were examined live in water as

well as after having been immobilised by heating. Some

larvac were killed in hot 70% ethano) and cleared In

glycerol by transferring to a mixture of 70% ethanol

and glycerol and allowing the cthanol to evaporate.

Measurements were made either with an ocular

micrometer or from drawings made using 4 drawing

tube and are presented in the text in millimetres as the

range followed by the mean in parenthescs.

Morphological terminology for the synlophe follows

that of Durerie-Desset (1985), A drawings are oriented

with the dorval aspect uppermost and the left hand side

of the nematode body towards the left margin of the

page,

Parasitic third-stage larvae of Aoorani were

distinguished from the synhospiralic nematodes

Nippustrongylus magntes and Odilia bainae by feanures

of the tail and cephalic extremity, based on material

obtained from monospecific experimental infections

with each of these two species. To obtain a

monospecific infection of adult F ratri, one naturally

infected R. fuscipes was killed, all P rare’ in the small

intestine were sorted while being maintained in warm

saline, and were transferred by enlerotomy to a

laborgtory-reared R, fuscipes under general

anaesthesia. Four weeks Later, the recipient rat was

killed, the intestine removed and. 10% buffered formal

saline injected rapidly into it to fix nematodes in: site,

Fragments of intestine with worms attached

subsequently were dissected and prepared for scanming

electron microscopy us deseribed above, Additional

segments were dehydrated; embedded in paraffin,

serially sectioned at a thickness of 10 um, and the

sections stained with haematoxylin and eosin fur

histological examinaticus.

Results

Adult mile snd female nematodes, fourth-stage

larvae and two parasitic third-stage larvae were

examine! morphologically undes the light microscope,

33 well as third-stage larvae cultured from faeces, Light

microscopic observations were compared with

scanning electron micrographs of the adults.

Poraustrastronegylus nati Obendarf, 1979

FIGS 1-38

Description: Adult: small nenxstodes, red when live,

spirally coiled in 3-5 tight coils, ventral sucface

innermost, capable of uncoiling and beconung straight

when maintained in warm isotonic solutions. Cephalic

extremity with prominent cephalic vesicle, 20-30 fine

transverse ridges on vesicle (Fig. 1). Mouth opening,

surrounded by four sub-median papillac and two

amphids, no fabial paplae visible; six rounded lips

project into mouth opening. Buccal capsule

prominently developed, heavily scleratised, circular

to sub-hexagonal in transverse section (Fig. 4), walls,

arched inwards, increase in thickness towards base,

lumen increases in diameter twards base; prominent

tooth projects inwards from dorsal Jobe of oesophagus;

sub-ventral teeth absent, esophagus elongate, slender,

clavate, widened at anterior extremity; excretory pore

yanable in position, usually anterior, occastonally

posterivr, to vesophago-intestinal junction; deirid tiny,

domed, al level of exuretory pore; nerve ring in mid-

oesophageal region, visible in few specimens only.

30 1. BEVERIDGE & M.-C. DURETTE-DESSET

“My

Figs 16-23, Paraustrostrongylus ratti Obendort: adult. 16-22, transverse sections of body, 16-19, male, 2.3 mm long; 16,

at posterior end of cephalic vesicle, 0.06 from anterior extremity; I7, in oesophageal region, 0.30 mm from anterior extremity;

18, in posterior half of body, 1.60 mm from anterior extremity; 19 in cloacal region, 0.10 mm from posterior extremity;

20-22, female, 2.6 mm long; 20, oesophageal region; 21, 0.50 from anterior extremity; 22, 1,90 mm from anterior extremity;

23, posterior end of male, ventral view, showing termination of ventral ridges (arrows) and termination of floats (f).

Legend: d, dorsal; J, left; r, right; v, ventral; ridges are numbered (1-7) in an anticlockwise direction from the left ventral

tidge. Scale line: 0,01 mm.

LIFE CYCLE STAGES OF PRATT Hv

Body covered with numerous fine transverse

annularions (Fig. 36); two lateral, fluid-filled cavities

(= floats) present on either side of body, extend from

unmediately posterior to vesicle to posterior region of

nematode. Synlophe composed of three ventral ridges

(1-3) (Pig. 36), oriented From right wo left, diminishing

in size from left to right; right Sloat with pwo dorsal

(5,6) (Fig. 35) and single ventral (4) ridge directed

towards left dorsal, ridges b-3 commente posterior to

vesicle; left floal, with single ndge (7) (Fig. 35)

directed perpendicular Lo body, commences posterior

to mid-oesophagus; right float with two dorsal (5,6)

and one yentcal (4) ridge; ridec 6 commences postenor

lo vesicle, followed by 5 then 4 in mid-oesophageal

region.

Male (measurements of 10 specimens). Total lenge

LYB-2.66 (2.30); maxitnutn widih (without floats)

,050-0.070 (0057); cephalic vesicle 0055-0065

(0054) long) oesuplragus 0,27-0.30 (0,28) lung, nerve

ning -cire@ 0.15 from anterior extremity; excretory pore

0.21029 (0.25) from antenor extremity; deitid

0.22-0,30 (0.26) from anterior extremity: spicules

0.23-0.32 (0.27) long; gubernaculum 0,025-1045

(0.037) long, Synlophe; ventral ridges ard right float

femiitiate near anterior extremity of spicules: teft Moat

continues tn leve) of gubermaculum,. Bursa symmetri-

cal, lobes indistinct, dorsal labe not separated fron

lateral lobes; rays 2 to 6 of bursa grouped im patter

of 3-2 (sensu Durette-Desset 1983) Ur 1-2-2 allowing

for highly divergent ray 2; ray 2 slender, short,

divergeit, nol reaching margin of bursa; rays 3 to 5

more robust, directed postero-faterally, of appreni-

mately equivalent size, not quite reaching margin of

bursa, my 6 shares common origin with ray 5, robust,

blunt, directed posteriorly, reaches margin of bursa;

dorsal trunk separate (rom lateral trunk; ray 8 uriges

from dorsal trunk, slender. does not reach margin of

bursa; rays 9 slivhily asymmetrical, shor, slender,

arise Close to origin of ray 10; ray LO stout, divides

hear extremity inte + branches; outer pair of branches

more rubust; fmal branches do not reach margin of

bursa. Spicules simple, elongate: alate; anterior

exthemitles irregularly knobbed, distal ups jouted: each

spicule with fine, spiniform ventro-lateral branch

arising in distal 1/6-1/7 of spicule; spicule tips

surrounded by expanded sclerutived [ange in \lorso-

ventral view; each main branch of spicule terminates

in two fine spiniform projections within flange;

pubernaculum elongate, rectangular in dorso-ventral

view, composed of two layers; genital cone heavily

selerotised, complex, cumical in shape, c. 1020 long,

base D020 wide; papilla 0 at tp of ventral lip of genital

cone; paired papillae 7 on dorsal lip of cone.

Female: (measurements cf (0 specimens), Total length

2.48-2.95 (2.70): maximum width (without floats)

0.060-0.070 (0.066), with floats circa (110; cephalic

vesicle (060-0070 (0,063) long, oesophagus 026-032

(Q.29) long) nérve fling circa O18 from anterior

extremity; excretory pore 0.21-0.27 (1123) fram anterior

extremity; deind 0.22-0.26 (0.24) from ameriog

extremity; bail OO8-O.U (0.10) lung; vulva to posterior

extremity 014-021 (0.17); egg 0,065-0.080 (0,074) by

6.035-0.045 (0.038). Synlophe: ventral ridges extend

to vulva; floats disappear in region of uterus, approx,

0.44 from tail, Tail extremely long, conical; vulva

immediately anterior to uterus, opening to exterior on

slight prominence; female genital system monodelphic

though with postendr uterus patent and ovary persisting

in vestigial form: vagina, vestibule and sphincter crea

0.06 long; infundibulum short, circa O04 long,

prodelphic, leads to elongate uterus containing 1-4 eggs;

eges thin-shelled, ellipsoidal.

Fourth stage larva: Small nematodes, spirally coiled

in 34 cous, ventral surface innermost, cephalic vesicle

absent; mouth opening surrounded by four submedian

papillae and two wmphids; lips absent, Hupcal capsule

sub-cylindrical, heavily sclerotised. teeth absent.

Oesophagus elongate, clavate, nerve ring in mid-

vesophagesl region; excretory pore in region of

oesophago-intestinal junction. Synlophe: body floats

absent; five ridges; three ventral ridges (1-3), anenied

from right to left. diminishing in size fron Jefi to right,

ventral ridge (4) on right hand side, oriented from right

to left; single dorsal ridge (5) on right hand side,

onented dorsal from right fo lefi; orientation of

synlophe oblique from right ventral to left dorsal at

about 60° to saginal axis. Tatl elongarm, conical.

Male (measurements of five specimens), Total length

1,09-1,76 (1.44), maximum width 0.033-0.046 (0.040),

oesophagus 0.21-0.28 (0.26), tail 0.049-0,085 (0.066),

Female (measurements of five specimens). Total length

1.47-2.00 (1.80), maximum width 0.030-0,052 (0.044),

oesophagus 9,25-0.29 (0.27), tail 0,052-0,143 (0,082),

Specimens with developed genitalia show distinct

posterior uterus, recurving into short ovary (Fig, 32),

Third stage larva» Parasitic: Two parasite third stage:

larvae were recorded, but detailed morphological

comparisons were possible from one only. Small

nematode, 0.76 long, spirally coiled in three coals;

buccal capsule cylindrical, very lightly sclerotised;

oesophagus slender, clavate, 0.18 bong: excretory pore

in posterior pesnphageal region, 0.12 from antenor

extremity; tail elongate, conical, with dorsal and ventral

projection. Syniophe composed of rwo pairs of alse

beginning on lateral aspects of body, perpendicular to

body surface; towarde mid-region of body, alae

gradually shift in position to dorsal and ventral; m md

hody region, the left ventral parr are larger with one

iu almost a mid-ventral position, and second ridge to

32

I. BEVERIDGE & M.-C, DURETTE-DESSET

Figs 24-32. Paraustrostrongylus rarti Obendorf: larval stages. 24-27 third. larval stage cultured in vitro; 24, entire larva,

lateral view; 25, cephalic extremity, lateral view; 26, tail, lateral view; 27, transverse section in mid body region; 28,

transverse section of parasitic third-stage larva from small intestine; 29-32, fourth-stage larva; 29, anterior end, lateral

view; 30, buccal capsule, lateral view; 31, transverse section of larva in mid-body .region, arrow indicates oricntation

of synlophe; 32, tail of female fourth stage larva with posterior branch of genital system. Scale lines: fig. 24, 0.1 mm;

figs 25-32,.0.01 mm; figs 25-26 to same scale. Legend: p, phasmid; u, posterior uterus and 0, oyary; ridges are numbered

in an anticlockwise direction from the Jefi-ventral ridge.

LIFE CYCLE STAGES OF P RaTiyY as

cme sick: of i; two dorsal alac smaller, one almost

doisal Lo position, Other lo one side of it.

freedivine: (measurcinents of five specimens).

Mender, elongate larvae, 0.50-0.55 (0.53) long,

maxima width 0.016002) (0.018); baceal capsule

cylindrical, ¢. 0.005 long, 0.002 wide, continuous with

selerotised internal lining of anterior oesophagus:

oesophagus slender, 044-016 (0,15) long; nerve ring

CLOTS-0.083 (0.082) from anterior end; excretory pore

CHOR 1.099 (0,096) fromanterior end; intestinal cells

filled with granules, number of cclly not ascertained:

genilal primorhium avoid, 0.008-0.013 (0.010) by

TODS-O008 C007), 128-0.33 (0,31) from anterior

extreme, tail clomgale, conical, 0.065-0,089 (0,075)

lon, with dorsal and ventral spike close to tip; larva

with four longitudinal alae, two on cach side of body.

in Jateral position, whois perpendicular to holy wall,

ditachment ta the intestine! nurse

Adult nematodes are coiled ventrally around

pitestenal villi (Pigs 33, 37) usually with the tail near

the distal Hip of the villus; when fixed. the coils of the

body are mameained (Fig. 34). At the site of

altachntent, nenalodes compress the willl (Figs 37, 38)

and, although they generally retract from the site of

attachment when lixed, the impressions of the ventral

ridges remain in the intestinal epithelium (Fig, 38).

Changes in the epithelium at the site of attachment

include cuboidal to squamous cpithelial cells,

clongahon (= flattening) of nucler associated with the

change 0 ao siyuumous cell type, loss of cytoplasmic

differentianon and the loss of the brush border of

muicavillt, Although sormetimely squamous, no defects

were detected in the epithelium, No marked

iiflanimatory changes were detectel but there were

significant numbers of mononuclear cells, macrophages

und lymphocytes, present in the Bani propria together

with a small number of cosinaphils

Discussion

Morphology of the adult

The description of the adult provided here

supplements the original description by Obendort

(979), which was found to be accurate in all essentials.

Obendort (1979) however, did not provide an apical

view of the untenior extremity and provided only a

single, anonenied drawing of the sy nlopbe of the adult.

In the present examination, the synlophie 1s described

in detail, including the oripins and terminations of the

body ridges, Duretre-Desset (1979) and Beveridge &

Durelte-Desset (1986) have shown that the number of

bendy ridges changes in the posterior region of the body

in several species of Axytresrrongyluy and

Paraustrastrangylus, but comparable data were lacking

for P. rat, This study demonstrated that in P rarti,

the principal nelges arise tm the oesophageal region and

persist to the bevel of the spicules in the male and to

the level of the vulva in the femile. In the male, the

bef, but nol the righe Moat extends almost to the level

of the bursa, as iv does also in P rrichosarl and in A.

safovialas, the latter being d spoctes which possesses

only one float F retti also resembles A. safestatus in

having three rather than the four ventral longitudinal

ridges present momost other members ol tfese gener,

The three principal ventral ridges arise Close to the

cephalic yesiele, while the ridges on the floats arise

somewhat more posteriorly, with the ridge on the left

float arising mid-way between the vesicle and the

exeretory pore and the two dorsal ridges on the right

Nout arise in the anterior oesophageal region, The

ventral ridge arises midway between the cephalic

vesicle and the exeretory pore. The onginy and

lerminations of ridges haye been repomed in few

confamilial species, but have been shown to be of

considerable taxonomig use at the species level in

genera such as Nerriaredirus (see Lichrenfels & Pilitt

1983). Preliminary observations by Beveridge &

Durette-Desset (1986) on species of Adstrostronyylus

suggest that this may be the case in the Merpetostrongy

fae, but features have been described in too few

species to allow any firm conclusions to be diawa,

An imeresting feature of the morphology of / farri,

noted by Obendor! (1979) was the presence of 4

vestigial posteriog branch to the female genial system.

His observations were confirmed inthis study, and the

same structure was also seen in the Jourth larval stage.

The genera Austrostrongylus and Paraustrostrongylay

wre considered to be very closely related but can be

separated on the basis of the presence of a sclerotised

genital cone in: Paraustrostrongylis, the absence of

ventral teeth in Pirastrostrongylus and the position

of ray 2 (Beveridge & Durette-Desset 1986), Species

of Paraustrostrong\las are invariably, monodelphic

whale most species of Austrostrongylues are didelphic,

The evolution of monodelphy has oceurred repentedly

in the Trichostrongyloidea (see Duretie-Desser 1985)

and the vestigial posterior uterus. in P rari provides

an obvious connection between the mortxtelphic and

didelphig dorms seen in these two elosely related

genera, [n other trichostrongyloid genera such as

Neohelignonedla, the posterior uterine branch imuy

persist in the adult nemiutode, but does so only as a

small collection of cells, posterior to the vulva (Durette-

Desset & Cassone 1987), rather than the almost fully

tormed bul diminutive posterior branch seen in P mare.

Anachment ta villi

Results presented here indicate that? rani attaches

to intestinal villi by coiling spirally qroond them, as

in certain other tachosirongylod neraiodes (Duretie

Desset 1985), The ventral body ridges clearly press

ime the intestinal epittelium, and may theref ire assist

34 I, BEVERIDGE & M.-C. DURETTE-DESSET

the nematode in maintaining its attachment to the villus.

The ventral surfaces of both body floats are also in

close apposition to. the epithelium and effectively

increase the surface area of the nematode in contact

with the intestinal epithelium. The ventral ridge of the

right float (4) and to some extent the latero-dorsal ridge

(4) of the right float (5) also cause indentation of the

epithelium, and may therefore also assist in attachment.

Once in place on a villus, dorsal ridges would seem

to have little function in attachment, and one of the

features of P ratti is that it has few dorsal ridges.

However, when observed in warm isotonic solutions,

the nematode is capable of uncoiling completely, and

evidence from the localisation of experimentally

transplanted nematodes indicates that they are capable

of migration within the intestine as is the case with

CP" _.. aT

— ~

a oe ye

Figs 33-38. Paraustrostrongylus ratti Obendorf, 33-36, scanning electron micrographs. 33, entire nematode coiled spirally

around villus in small intestine; 34, entire nematode, 9, showing body coils (4); anterior end to left; 35, dorsal surface

of nematode showing ridges 5 and 6 of dorsal aspect of right body float and ridge 7 on lateral aspect of left float; 36,

ventral surface of posterior region of body showing ridge 7 on left body float and ventral ridges | and 2; 37, longitudinal

section through villus showing posterior part of nematode coiled around yillus, with anterior end extending to left of field;

38, histological section through villus at point of attachment of P. ratri, with nematode retracted, leaving sites of indentation

(arrows) of ridges in epithelium. Scale bars: figs 33, 34, 37, 0.1 mm; figs 35, 36, 38 0.01 mm.

LIFE CYCLE STAGES OF Fl RAITI a8

other trichostrongyloids (see Crofl & Ma 1977), The

dorsal ridges may therefore be of use daring nematode

migrations wijhin the small intestine, when they are

uncoiled and are moving between villi,

Morpitology of Larval surges

The morphology of the buccal capsule of the fourth

stage of P ratii resembles very closely that of the fourth

stage of B. pearseni (sec Humphery-Smith 1980) In

peramelid marsupials and Globocephaloides

trifidospicularis: (see Beveridge (979), a species

parasilic ln macropodid marsupials. The Globocepha-

loidinae, to which the latter genus belongs, was placed

Within the Herpetostrongylidae by Durette-Desset

(1983) bated on features of the bursa and buccal

capsule, Since members of ihe sub-family Jack a

synlophe, their precise alfinities have not been

established, However, the similaritles between. the

fourth stage larvae of G, trifidospiculanis, & pearsuri

and P rat provide additional evidence that the two

subfamilies are related.

Only two parasitic third larval stages of A rated were

found, However, they were identified by the

characteristics of the tail, which was identical to that

of larvae cultured from faeces. Both parasitic larvae

were spirally coiled, as are the fourth stage and adult

The pairs of alae Which were clearly lateral in position

at (he anterior and posterior extremities of the body

were slightly shifted in position in the mid-body region

26 that the larger, left pair were almost ventral in

position, while the smaller, right pair became almost

dorsal in position. This gradual shift in ridge pasitlon

and the hypertrophy and ventral shift of one pair of

alae presumably aids jn attachment, as is the case in

the adult, Two pairs of lateral alac were evident in the

free-living third larval slage but they remained in the

Jateral position throughout the length of the nematode

body. Several other teichostrongyloids have paired

lateral alae tn the third stage larva (Ecker & Schware

1965; Duretie-Desset & Cassone 1987), The apparent

change tn Orientation of the ndges in the mid-body

segion during the initial parasitic phase of the life cycle

appears to assist the nemtapode in sttachiny to villi, bur

(he mechanism by which this might occur is unclear.

Usually there is no change in the synlophe unless a

Moult Occurs, bul lf this ingiance, the change in

position of the alae is vistble in the entire nematode

as well as in sections, Additional observations are

clearly needed co confirm the results reported here.

Evolutionary relationships

The description of the fourth larval stage of F rarsi

feovides additional insight into the evolution of the

Herpetustrongylidac. The type ef information which

can be provided by the larval synlophe has been

discussed by Duretie-Desset (1985), Generally, the

larval synlophe demonstrites primitive features

compared with thar of the corresponding adult and

frequently resembles the adults of other, related genert

thereby allowing phylogenetic reconstructions, In the

ease of P revi, as in other tichostrongyloids, the larval

synlophe differs markedly from that of the adult_ The

larva lacks floats; in the larva, the axis of orientation

is oblique rather than being frontal, as occurs in the

adult and the synlophe has iwo fewer ridges, lacking

those found og the left flout and on the dorsal right

float of the adult, The farval syolophe of P neni most

closely resembles that found in the adults of

Swarostronzylus and Dessetostrongylus but differs

from that of Sutarcstrongylus in baving one. rather than

two dorsal ridges, ane sn onentation since the synlaphe

of Sutarasrronevius has a frontal orientation, simular

to thar af the adult Prati. The oblique orientation

of the synlophe of the larval stage of Ft rare| resembles

that of Desselastrongylus, from which it differs only

in having fewer dorsal ridges. The homology of ridges

between. adult and larva here considered wo be most

likely is that ridges | to 3 of the larva correspond with

ridges 1 to 3 of the adult, while the two additional

ridges of the Jarva (4 and 5) correspond bo two of the

ridges on the nght float of the adult (4 and 5). An

alternative possibility would be that the first four ndges

from left to right correspond to the four yentral ridges

present in rnost species af Austrastrangylas and some

species of Puruustrestrongvlus, wilh the filth ridge

corresponding to one of the ridges on the right float.

This interpretation invalves postulating the loss of #

yentral ridge in adult P rani and the appearance of

two rather than ane new ridge for the right float, The

more parsimonious of the two hypotheses has been

chosen here, 11 is also consistent with the hypothesis

of Humphery-Smith (1983) that species of Wooileva

with three left-ventral, obliquely atiented ridges, were

the likely ancestors of Dessetosirongylus.

The suggested intermediate farms in the proposed

lransition series for the evolutionary lineage hetween

Uholleya and Austrostnongylus/Paraustrastrong wis

have been Heveridgiella (see Durette-Desset 1982;

Humphery-Smith 1983) and Dessetrstranewus (see

Beveridge & Durette-Desset 1986). Potricfaline, an

additional possibility, has a frontally symmetrical

synlophe. while both Beverideielfa and Dessete-

Strong ylug have an oblique orientation to the synlophe.

Patriciatina was considered to have been derived from

Beveridgielia by Humphery-Smith (1983) and Cassone

et al (1986). Beveridgiella has 4 greater number of

dorsal ridges than Dessetestrongyius, and increases in

the number of ridges oocur in # number of evolutlonary

lineages within the Trichostrungyloiden (Durette-

Desset 1985), suggesting that the synlophe in species

af Reveridgicifa is probably derived from a

Dessetostrangylus-like ancestor. This i fact is shown

in the larval stage of 8 pearseil (see Humphery-Smith

36 I, BEVERIDGE & M.-C, DURETTE-DESSET

1980), which has a synlophe close to that of

Dessetostrongylus. The synlophe of the fourth-stage

larva of P rarti differs from that of adult

Dessetostrangylus moorhousei only in lacking an extra

dorsal ridge, Because of the close correspondence

between. their synlophes, it appears likely that

Paraustrostrongylus evolved from an ancestor

vesembling contemporary species of Dessetostrongylus,

thus supporting the hypothesis proposed by Beveridge

& Durette-Desset (1986) and Cassone et af, (1986),

Acknowledgments

Rats were obtained under permits from the Victorian

Department of Conservation and the Environment

(90-053 and 91-095), We wish to thank Christine

Andersen for expert technical assistance, Lilliana

‘Tatarczuch for ibe preparation of the thin sections and

Dr D. M. Spratt for comments onthe manuscript. This

work was supported financially by the Austrahan

Research Council,

References

Bevenipce, 1. (1979) A review of the Globocvephaloidinas

Inglis (Nematoda: Amidostomatidae) from macropadid

marsupials. Aust J. Zool, 27, 151-175,

(1986) Coevolutionary relationships of the helminth

parasites of Australian marsupials. pp 93-117. In

“Coevulutiun and Systematics” A, R. Stone & D. L,

Hawksworth (Eds). The Systematics Association.

(Clarendon Press, Oxford)

& DURETTE-DESSET, M.-C, (1986) New species of

Austrostrongylus Chandler, 1924 (Nematoda,

Trichostrengylvides), from Australian marsupials, with a

redescription of A, minutus Johnston & Mawson, 1938, and

description of a new genus Sutarastrongylus. Bull. Mus.

natn. Hist. nat., Paris 4éme sér. 8, 145-170.

Cassone, J., DURETTE-DesseT, M.-C...& PRESIDENTE, P, J,

A. (1986) Nouvelle hypothése sur I'évolution des

Herpetostrongylinae (Nematada, Trichostrongyloidea)

parasites de Marsupiaux ausiraliens. /bid, 267-283.

Croit, N. A. & Ma, K. (1977) The location of parasites

within their hosts: the influence of surgical manipulation

of the intestine and blood supply on the dispersion of

Nippostrongylus brasiliensis in the rat. Int. J. Parasitol.

7, 21-26,

Durette-DesseT, M.-C. (1971) Essai de classification des

Nématodes Heligmosomes. Correlations avec la

paléobiogeographie des hates. Mem, Mus. natn, Hist. nat,,

Paris, sér A, Zoologie 69, 1-126.

(1979) Compléments morphologiques a letude des

enres Austrostrongylus Chandler, 1924, et

raustrostrongylus Mawson, 1973 (Nematoda,

Trivhostrongylidae). Bull, Mus. natn, Hist. nat., Faris,

d2me sér 1, 1023-1030,

—____ (1982) Relations hdtes-parasites chez les

Trichostrongyloides. Mém. Mus. nam. Hist, nat., Paris

ser. A,, Zoalagie 123, 93-100.

_—_—. (1983) CIA Keys to the Nematode Parasites of

Vertebrates, No, 10 dn R, C, Anderson & A. G, Chabaud

(Eds). Keys to genero of the superfamily

Trichostrongyloidea. Commonwealth Agricultural

Bureaux, Farnham Royal, pp.6s.

(1985) Trichostrongyloid nematodes and their

vertebrate hosts: reconstruction of the phylogeny of

parasitic group. Adu Parasitol. 24, 239-306.

& Cassone, J. (1987) Sur deux Nematodes

Trichostrongyloides parasites d'un Muridé africain. TT -

Chronologie des cycles, description des stades larvaires et

des immatures, Ann. Parasitol. Hum, Comp. 62, 133-158,

Ecxerr, J. & SCHWARZ, R. (1965) Zur Struktur der Cuticula

invasionsfhaiger Larven einiger Nematnden, Zeit, f

Parasitenk. 26, 6-142.

HuMrPHERY-SMITH, J. (1980) Beveridgiella n. gen.,

Dessétostrongylus 0. gen. (Nematoda, Trichostrongyloidea}

parasites de Marsupiaux australicns: Bull, Mus. natn. Hise.

nat., Paris, déme sér. 2, 999-1012,

(1983) An hypothesis om the ¢volution of

Herpetostrongylinae (Trichostrongyloidea: Nematoda) in

Australian marsupials, and their relationships with

Viannaidae, parasites of South American margupials, Anse.

J. Zool. M, 931-942.

LicuTenre.s, J. R. & Piuirr, P. A. (1983) Curicular ridge

patterns of Nematedirys (Nematoda: Trichostrongyloidea)

parasitic in domestic ruminants of North America. with

a key to species, Proc. Helminthol. Soc, Wash. 50, 261-274.

ORFNDORF, D, L. (1979) The helminth parasites of Rattus

fuscipes (Waterhouse) from Victoria, including description

of two new parasite species. Aust. J. Zool: 27, 867-879.

A NEW SPECIES OF GRILLOTIA AND PSEUDOGRILLOTIA

(CESTODA: TRYPANORHYNCHA) FROM AUSTRALIAN SHARKS, AND

DEFINITION OF THE FAMILY GRILLOTITDAE DOLLFUS, 1969

BY R. A, CAMPBELL* & I, BEVERIDGEF

Summary

Two new species of trypanorhynch cestodes Grillotia amblyrhynchos sp. nov. and Pseudogrillotia

spratti sp. nov., are described from the spiral valves of carcharhinid sharks in Australian waters,

G. amblyrhynchos is distinguished from congeners by possessing seven hooks per principal row,

four intercalary hooks that merge with a wide longitudinal band of small hooks on the external

tentacular surface, and a basal armature with microhooks on the external surface. Pseudogrillotia

spratti sp. nov. is the fourth species in the genus and differs from congeners in the absence of a

basal swelling, lack of a pars post-bulbosa, 9-10 hooks per principal row, absence of a band of

hooks from all but the basal region of the tentacle and a single row of intercalary hooks. The

subfamily Grillotiinae Dollfus, 1942 and the family Pseudogrillotiidae Dollfus, 1969 are discussed.

The subgenus Paragrillotia and family Pseudogrillotiidae are rejected. Grillotiidae Dollfus, 1969 is

formally defined for the first time and the three genera, Grillotia Guiart, 1927, Progrillotia Dollfus,

1969 and Pseudogrillotia Dollfus, 1969 are admitted with revised diagnoses.

KEY WORDS: Grillotia, Pseudogrillotia, cestode, new species, Australia.

‘Dransactions of the Royal Sactery of S Awss 993), WIE, 37-46

NEW SPECIES OF GRILLOTIA AND PSEUDOGRILLOTIA

(CESTODA: 'TRYPANORHYNCHA) FROM AUSTRALIAN SHARKS, AND

DEFINITION OF THE FAMILY GRILLOTIIDAE DOLLFUS, 1969

by R. A. CAMPBELL* & |. BEVERIDGET

Summary

CampPaeLe, R, A, & Bevirmce, |, (1993) New species of Grilloria and Pseudoprillotia (Cestwda;

Trypanorhyncha) from Australian sharks, and definition of the family Grillotiidae Dollfus, 1969. Trans. X. Soc,

§. Aust. 117(1), 37-46, 4 June, 1993,

Two new species of trypanorhynch cestodes Grillotia amblyrhynchos sp. novo and Pyeudogrillutia spratti sp.

nov., ate described from the spiral valyes of carcharhinid sharky in Australian waters. G. amblyrhynchas ix

distinguished from congeners by possessing seven hooks per principal row, four intercalary hooks that merge

with a wide longitudinal band of sinall hooks on the external tentacular surface, and a basal armature with microhooks

on the external surface. Pseudogrillona spratd sp. nov. is the fourth species in the genus and differs from congeners

ip the absence of a basal swelling, lack of a pars post-bulbosa, 9-10 hooks per principal row, absence of a band

af hooks from al! but the basal region of the tentacle and 2 single row of intercalary hooks. The subfamily Grillotiinae

Dollfus, 1942 and the family Pseudogrillotiidae: Dollfus, 1969 are discussed, The subgenus Prragrilloria and family

Pseudogrillotiidae are rejected. Grillotiidse Dollfius, 1969 is formally defined for the first tume and the three genera,

Grilloda Guiart, (927, Pragrillotia Dollfus, 1969 and Psewdogrilloria Dollfus, 1969 are admitted with revised

diagnoses.

Key Worps, Grillotia, Pseudogrillotia, cestode, new species, Australia,

Introduction

Few records of Griflotia or its relatives exist from

studies of the cestode parasites of Australian fishes.

Three Grillofia species have been described from

larvae taken from Australian teleosts, one by Shahsarorn

& Lester (1982) and two by Sakanari (1989), The genus

Pseudogrillotia was not listed as occurring in the

Australian region in a checklist of the parasites of fishes

by Beumer ef a@/, (1982). A new species of Prendo

grillotia trom a carcharinid shark reported herein

therefore constitutes the first record of this taxon in

the Australian region and a tiew species of Grillotia

represents the first description of an adult of this genus

from the region,

Materials und Methods

Cestode specimens were fixed in 10% formalin and

lransferred to 70% ethanol for storage. Tentacles were

dissected free and mounted in glycerine jelly or balsam

to facilitate examination.. Whole mounted cestodes were

stained with Celestine Blue, dehydrated in a graded

ethanol series, cleared in clove oil and mounted in

Canada Balsam. Terminology and numbering of

tentacular hooks follows Dollfus (1942),

Measurements are given in micrometers as the range

followed by the mean in parentheses, unless otherwise

* Department of Biology, University of Massachuserts

Dartmouth. North Dartmouth, Mussachusctts 02747 U.S.A,

¢ Pppartanent of Veterinary Science, University of

Melbourne, Parkville, Vic 3052,

indicated, The number of measurements is indicated

as (n). All specimens were measured, Type specimens

have been deposited in the South Australian Museum.

Adelaide (SAM).

Drawings were made with a drawing attachment on

an Olympus BH microscope. [n the figures, vitelllne

follicles ure shown only along the lateral margiris of

proglotides for the sake of clayity.

Grillatia amblyrhynchos sp. nov.

FIGS L9

Types; Holotype: from spiral valve uf Carcharhinus

amblyrkyachos (Blecker, 1856}, Townsville,

Queensland, 4,x7i,1985, coll, B, G. Robertson, SAM

V4213; paratype: | specimen, same dare, SAM

HC233372.

Description: Moderate sized worms, wtal length up

to 30 mun. Scolex (n=2) long and broad, 3.9, 4.2 mm

long, 1.18 mm wide at level of bulbs. Two broadly oval,

patelliform bothridia, 710-900 (780) long by 1070-1180

(1120) wide, notched on posterior border, margins only

slighdly thickened and curved medizlly, posterior and

lateral borders frec; adherent surface spinose with a

barrow modian fissure c. 266 long, paralleled by four

smaller fissures on either side al regular intervals

(Fig. 1}. Pars vaginalis 2.56, 2.74 mm long: Tentacle

sheaths spiral; prebulbar organs distimct. Bulbs

140-1180 long by 232-264 in diameter; retractor muscle

originsies at midlength of bulb. Pare post-bulbosa

lacking, Ratio of pars bulbosa to pars vaginalis 1:2.3

R. A, CAMPBELL & I, BEVERIDGE

mt ee

WF Aye yeaa BRO a 0. SONNY

KY

or , aa

a ae a

7 7

Nant

4 ay

Shey \

P Wy

/ i 4

VIA

Ue tN

OMI JOOS Db) SSS

4. Grillotia amblyrhynchos sp. nov; 1, scolex; 2, immature proglottis showing icstes; 3, mature proglottis, vitellaria

shown only at margins; 4, bulb and prebulbar organs. Scale lines: 1-3, 1 mm; 4, 0.1 mm,

Figs 1-

NEW SPECIES OF GRILLOTIA AND PSEUDOGRILLOTIA 39

to 1:2.4. Scolex ratio (pbo:py:phulb) 1:3.3:1.4-

Tentacles short, length 1.40-1.46 min, tapering. tending

to collapse; dianieter al base 110-114, diameter at mid-

tentacle 95, diameter at tip 34, Armawwre

heieroscambhous (sertsu Keveridge & Campbell 1989),

heteromorphous, with band of small hooks in

metabasal region; hooks hollow, Basal armature

consists of numerous milcrohooks on extemal face

between first four principal rows of hooks (Fig. 6);

intemal face of basal armature consists of large books,

reduced im size fram those of metabasal region.

Principal hook rows alternate, consist of ascending

half-spirals of seven hooks each; rows begin on imernal

surface, merge with band of small hooks on external

surface of tentacle (Fig, 8). Hooks of first proximal

row (base of tentacle} reduced in size and number;

hooks I(1') of first proximal row well separated but

with transverse bases. Hooks 1(1") in all other rows,

separated by distinct space, bases oriented

longitudinally, rose-thom shaped, gradually increasing

to maximum size at mid-tentacle then decreasing in

size toward tip of tentacle, length 36-63 (52), base

94-46 (41), with distinct heed 4-1 (9) and me 4-13 (10),

height 23-38 (32) (n=10). Hooks 2(2") rose-thorn

shaped, similar to 1¢1 °) but smaller, length 36-49 (42),

base 27-38 (31), heel 8 tne 9, height 19-29 (23), Hooks

3(3") erect, falciform, with extended heel 4-6 (5), toe

absent, hook length 4-44 (39), base 19-23 (22), heaphi

22-30 (27), Hooks 4/4") and 5(5") falciform, erect

slighily smaller, heel feduced, we backing: 444) length

29-42 (38), base 10-15 (13), heel 4, feng 17-23 (24);

5(5") Jength 23-25 (24), hase 11 heel 2, beight (7-24

(20), Hooks 6(6°) and 7(7°) spiniforn, smaller mn

proximal rows 1-4, length 10-15, but with distinct toe

4-6; toe absent from Ihese hooks distal to row 4; 6(6")

length 17-21 (19), hase 10, beight U-19 (15); 7(7') lengtir

15-21 (36}, base 8-10 (9), heawbt 13-18 (16). Hooks 6(6')

ara 7(7") of principal rows much reduced and merge

With band of hooks of extemal surface of tentacle.

Viewed From bathridial or antibothridial surfaces of

mmetahasal region, single row of 3-4 iuercalariny hooks

between principal rows beginning with hooks 444’)

of 3(5'). Second Intercalary row of two hooks may

be presenl between principal rows in basal region of

tentacle. Intercalary hooks spiniformn, length 1-19 (15),

base 5-6 (5), beighn 6-'6 (10). Intercalary honk rows

merge with irregularly arranged band of hooks on

extemal surface, Band of hooks opposite proximal rows

1-10 small, wocinate, length 8-13 (10), base 4-6 (5),

heignt 4, toe 4. Remainder of hooks forming band

distal ra row 10 spiniform, length Le2t (8), base 4-4

(6), toe absent, heel 2, heaght 1-19 (17),

First evidence of segmentation about 680 fron

scolex, segments narrow, gradually increasing in length

with maturity, becoming twice as long as wide when

mature. Mature segments acraspedote, 1.92-2.36 11m

(2.22 mm) by L40-1.50 ou (L4S mim}, terminal

segmonts ¢. 2.50 mm by 1.44 mm Genital pores

lateral, irregularly alternate, post-equatorial, 60-64

percent of segment length from anterior margin in

mature segments, Cirrus lor possibly an

hernaphnoditic) sac pyniform, 258-400 by 133-140,

cortaining sinuous sperm duct. No armature visible.

External seminal vesicle not observed. Testes

subspherical, 200 by 180, numerous, about 376-532

(420) in mature segments, occupying all avatlable space

medial to csmoregulatory canals including postovanan

space. Vagina narrow, surrounded by gland cells,

closely parallels posterior border of cirms sac, forming

a bulbous dilatation ventral to proximal pole of cirrus

sac. then tums posteriorly at Toidlime as chlated rube

and cxicods to ovarian isthmus to form narrow

fertilization dict, Ovary 144-176 by 656-728,

subterminal, consisting of two small, transversely

elongated asymmetrical lobes joined by ahort isthmus,

poral Jobe smaller, Jobes subdivided into numerous

lobules. Mehlis' gland immediately posterior to ovarian

isthmus, c. 144 in diameter, Vitellarium follicular,

forming layer encircling osmoreguiatory canals and

Teprodactive organs. Uterns simple, median tbe.

terminatiog in anterior one-fifih of segment. Ventral

osmoregulatury canal largest, diameter 38; dorsal

osmoregulatory canal extremely narro\y, sinuous,

diameter 2.

Brymology: The species is named afer its host, C.

amblyrhyncitas.

Remarks; Presently, (here are 20 valid species of

Grillatia (see Sakanari 1989).

G. amblyrhynchor sp. nov. has characlers of the

subgenus Peragnfloria which Dollfus (19694) created

to accommodate species of Grillotia in which the band

of hooks could not be distinguished from tie adjacent

intercalary rows (i.e. the inverealary rows merged wilh

the band of hooks). Only two species have been

aliributed to the subgenus Puraprillotla, G,

(Paragrilloria) simmonsi Dollfus, 1969 and G,

(Paragriliaia) rowel Campbell, 1977. Catra &

Gavarrino (1999) provided a redescription of

Rirntchobarkrlio situle Linton, 900 and showed that

G. (F.} simmonsi Dolltus, 1969 was a junior synonym

of it, They proposed the new combination G. stmilts

but tid not discuss Dollfis’ (1969a) placement of the

species, now recogmzed as G, similis, in tho subgenus

Paragritlotia, Conceptually, the creation of

Poragrilloria as a subgroup of Grillotia is useful except

for the fact that @, simmons: does not fit the subgeneric

definition. Griflata (P.) rowei was the first species

described that met the subgeneric definition (Campbell

1977). G. amiblyritynehos sp, nov. is a second species

with Uhis subgeneric character, The problem is not

resolved by retuining G. simimenst as type of the

subgenus Faragrivune Peragetiada, then, should be

ciiminared on the basis of its chosen type species.

R. A. CAMPBELL & i. BEVERIDGE

Ute it bah ELE

> Caw calG AG

bs a a a pa S ue ae SSS

DP (ec CO BaF So =

c4 » (Qa

oe ms Yew ts

SP Gort.

Of Zi VE

fi

1 aS: f ise Ue

hy ds fe

wn

gion, external face; 9, hooks | through 7 of principal row. Scale lines:

rhynchos sp. nov.; tentacular armature: 5, basal region, internal face; 6, basal region, external face;

vion, bothridial face; 8, metabasal re

$ 5-9, Grillotia amblyi

5-8, 0.1 mm; 9, 0,05 mm.

7, metabasal re;

Fig

NEW SPECIES OF GRILLOTIA AND PSEUDOGRILLOTIA 4

Both G. simifis and G rowel have six hooks per

principal row untike G. amPlyrinwiches sp, nov. which

has seven hooks per principal row. Nome of the

remaining species of Grillotia has seven books per

principal pow (Sakanari 1989). In G, similis und G.

anbiyrtynchos there are four hooks pet intervalary

row. There are two of threc interedlary hoaks per row

in G, row? but the hooks have transversely elongated

bases whereas those of G: amblyriynches are oval and

have a longitudinal orientation. In G. similis, the first

two intercalary hooks in each row (a and b) are twice

the length of intercalary hooks c and d whereas in G,

amblyriyiches the intercalary hooks are subequal, The

proup of microhonks on the extemal face of the basal

armature of G. amblyrhynchos is absent in G, similis

and G, rewei. The presence of microhooks in the basal

armature is typical of 13 species of Grillotia,

Differences in the numbers of hooks across the bands

uf these species is worthy of note, In G, similis only

@ single hook file remains between the Lntercalary rows

and poncipal rows on the external Face (See Fig, &

Cara & Gavarrino 1999), In Fig. 19 of Dollfus (1969s)

the band of hooks of G. simumwnsi is reduced to 1-3

hooks in width if one disregards all the principal hooks

and |ntercalary hooks Therefore, the charactenstic

feature for the subgenus Faragrilioria of having the

intercalary rows Metge timpercepubly with the “band”

of hooks is misleading because there is only a single

file of hooks rather than # band exclusive of the

intercalary and principal rows. In fact, this armature

pattern demonstrates an intermediate pattern of

symmewy bewween afypical betetoacanths and

pocciloacanths with a single chainette: In G. rowel =

single large hook and two small hooks are present on

each side of the external face between the opposing

Principal rows and in G. amblyrkynachos a band 6-8

spiniform hooks in width is present, The mature

provketides of G, similis and G. umblyrlivnchos

possess postovarian testes, a feature Dollfas (1969b)

considered significant in separating Progrillaria (rom

Grilloria. An heemaphrechtic sac was noted in G.

eriniceus by Dollfus (942), and this feature has been

conlinmed by us in additiumal material of the species,

It was also shown to occur m G. similis by Cara &

Guvarrine (1889) Beveridge & Campbell (3988) have

pointed out the presence of an hermaphroditic sac in

Numerous trypanorhynchs and have remarked upon the:

generic and even familial significance of Ure genital

terminalia,

Pseudogeillotia spranti sp. nuv.

FIGS 10-16

Types: Holotype: from spiral valve of Carehusrininues

brachyurus (Guenther, |R7D), Tatham, WNS.W,,

AL xtil984, coll, D.M, Spratt, SAM 4212;

paratypes: four specimens, same data AHC HC23331.

Description; Scolex craspedote, very Jong and slender,

62-84 mm (7.4 mmj n=5) Jong, maxinnunr width

088-116 mm (1.00 mm) an reginn of bulbs; two

patelliform bothridia, 740-980 (830) wide, posterior

margin indented, with a pair of fissures on lateral

margins. Pars bolfridinlis 088-100 mm (0.95 mm);

temacle sheaths regularly sinuous, pars vaginalis

367.1 mm (64mm); bulbs short and wide,

112-4.28 mm (1,21 mm) long by 220-340 (270) wide:

prebulbar organs prominent; retractor muscle origin

undetermined. Pars posibulbosa absent, Velum very

shart, 30-80 (50). Scolex ratio (pb: pv: plulb) 1: 67

13. Tentacles up to 1.2 long, 70-140 (10) in diameter,

basal swelling lacking. Armature hetercacagthous,

heteromorphaus; heoks hollow. Principal rows of

hooks alternate, ascending in half spirals of 9-10) hooks

per row from external surface. Hooks )()) separated

by a wide space, rose-thorn shaped, large, 70-100 (85)

long. base length 60-70 (65), height 40-60 (50, n=10);

hooks. 2(2') rose-thorn shaped, smaller, 60:75 (70)

long, base 40-60 (45), height 55-80 (66). Hooks 3(3°)

ty 9(9') elongate, slender, bases narrow, gradually

diminishing in size, recurved at tip; 3(3') length and

height 65-85 (77), base 15-35 (25); 4(4°) length und

height 70-90 (84), base 15-30 (23); 5(5") length and

height 65-80 (72); 6(6'} length and height 60-80 (65);

8(8") length and height 45-80 (62), 9(9°) length and

helt 48-70 (62); 1009") length and height 44-56 (49).

Single row of six intercalary hooks, 20-32 (23) long,

present between principal rows alyned with principal

hooks 5(5*) to 10010"). In basal region, principal hooks

of bothridial surface larger than those: of antibothridial

surface; ascending series of six hooks in file number

1, lengths 72, 55, 60, 70, 90, 95; hooks in file number

1’ fengihs 100, 90, 85, 90, 95. Internal surface of basal

armature with band of smalt microhooks, 12-28 (16)

jung, extending in closely spaced V-shaped rows along

tentacle but eading at level of fifth principal rows; hand

of amicrohooks diminishes from 12 hooks wide

proxtmally at. first principal rows mp four hooks distally

at fifth principal rows; band of hocks absent distal to

lift principal row of hioks.

Longest speciinen 50 tin, wilh about 65 seements;

first mature segment about cumber 30. Segments

acruspedote, largest mature segment 18 mni by

0,97 mm; genital pore m posterior half of segment,

980% from antenor margin; irregularfy alternate, Testes

nuinerous (>300}, filling all available intervascalar

medullary space, Linning thick pestovarian band,

40-60 (50, n=|0) in diameter; vss deferens visible at

bevel of avanan isiiimus, coils extending just antenior

to level of genital pore then laterally to enter cleus

sec. Hermaphroditic sac weakly developed, abour 230

hy 0; external seminal vesicle present, weakly

developed. Vagina mabiform, diameter about 20, joining

R. A. CAMPBELL & I, BEVERIDGE

42

—

Gas

if Va ot ie

rs Se

a e Ak

“|

\

TS

—

Figs 10-1. Pseudogrillotia spraiti sp. nov.; adult worm: 10, scolex; 11, mature segment, Scale lines: 10, 1 mm; 11, 0.1 mm.

NEW SPECIES OF GRILLOTIA AND PSEUDOGRIELOTIA 43

cirrus within hermaphroditic sac to form

hermaphroditic dict. Vagina curves posteriorly bo

ovarian isthmms, dilating to 40 but not forming distinct

semittal receptacle. Ovary Gmmature, lobes 150 by #0

Mehlis’ gland posterior 03 iwvarian isthmus, 160 by 90;

Uterine duct coils anteriorly nearly to level of

hermaphrodittc sac; ierus tubular, ascends anterlorly

nearly to antenor end of segment, utetime pore absent.

Virelline. follicles form a circum-medullary sleeve

around proglottis, Lateral osmoregulatory canals berely

visible.

Remarks; Dollfas (19699) created fhe perus

Pseudegrillotie and the family Pseudogrillotiidae for

F pleistacartha because of its combination af two

bothridia, craspeiote scolex, metabasal armature with

a band of nucnerous small hooks, Jack of a chainette

and possession of a postlarval stage Carvasal et al.

(1976) added a second species, P hasipinciata, from

C. amblyrhynchos in Hawaii. P sprarti sp. nov. is

similar lo F pleistacantha ia the number of hooks per

Poincipal row and Jack of a basal tentacular swelling

but can be distinguished from P pleistacantha by (1)

Wick of 4 continucus band of small hooks im the

metabasal armature and (2) lack of a pars post-bulbosa

and velum. The new species can be distinguished fron

P. basipwncrata by (1) the lack of a continuous band

of hooks in the mnetabasal armature, (2) lack of a well

developed basal armature, (3) absence of a basal

swelling and (4) more hooks per principal now (9-1

vs. 6).

P peruviune described by Escalante & Carvajal

(1924) is most similar to P sprarti sp. noy. in having

the bancl of hooks restricted tu the basal reglon of the

temacle fut is readily distinguished from the new

species by (1) possession of four intercalary hook rows

between cach pair of principal rows compared with

one intetcalary mow fy FP spraiti, and (2) differences

in the stape of the large basal hooks (Fig. 14),

Carvajal ev al. (1976) emended the family diagnosis

of Pseudogrillotiidae with the description of the first

species based upon an adult worm. The diagnosis is

still in agreement with Bo aprotti and 2 peruvidna

except that, 25 in species of Pterobothnidae, viz. 2

lurmaliensis Carvajal Combord & Campbell . 1976, the

bard af hooks may be absent from chc metabasal

armature. All other chanicters are in Agreement.

Discussion

‘Trypanorhynch cestodes of the subfamily Grillotilixse

Dollfus, 1942 and the family Pseudogrillotiidac

Dolifus. 1969 have been reported from sharks ani

skates in the Mediterranean, North Ajlantic, olf the

west coast of buth North and South Amenvx, Hawaii

and recently from Australian waters. Dollfus (1942)

created the subfamily Grillotiinae within the family

Lacistorhynchidae Guiart, 1927 distinguishing lt from

other poeciloacanihs with two bothridia by (|)

possession of a longitudinal band of hooke instead of

a chainette and (2) presence of intercalary hooks

between the principal rows. He further noted the

possession of two notched patelliform bothridia,

acraspedote scolex and lack of a postlaryal stage

(without blastocyst), Ina sepunile paper, Doilfus (1946)

created the subgenus Prygrillotia to accommoesiate x

new species that lacked posturvarian testes and later

(Doilfus 1969b) elevated Pragrilforie to generic status

with an erended diagnoses, distinguished for

Grilfotia by very long bulbs, no enigmatic prebulbar

organs, absence of postovarian testes and a external

band of hooks that 1s discontinuous but merges with

the interpolated rows of hooks. Dollfus (19693) also

created the subgenus Puragrilloria, and the familly

Pseudogrillotiidue.

Dollfus (19694) emphasized the distinction between

the armatures of Grifloria and Lacistorlynchus by

erecting a new famity Grillotildae because it separated

genera with chainettes (Lacisarhynchns,

Callitetrarhynchus, Flericeps, vide Reveridge &

Campbell 1989) from genera with interculary rows and

4 longitudinal band of heoks on the external surface,

i.e, Grilfotia, Dollfus (1969a) distinction stands m

contradiction to the keys of Yarnaguti (1959) and

Schmidt (1986). We agree with Dollfas (1969a) and

recognize a further distinction of the armature

possessed hy grilfotlids as being suitably

accommodated by the definition of “atypical

heveroacanthous" in which “there are more rows of

hooks-on the external fice than on the internal face

(of the tentacle)”. Our interpretation Of trypanorhynch

armatures includes the Grillotiidae, Pierobothnidae,

Molicolidae, Otobothriidae and Rhmoptericolidac as

atypical hetercacanths thus separating those families

with intercalary rows and longitudinal “bands” of hooks

from theiz former classical inclusion with the

pocciloacanthous families possessing chainettes, In our

opinion the term “poeciloacanth" is best restricted to

familics with chaiettes, as defined by Beveridge &

Campbell (1989), and considered distinct froni those

wilh longitudinal “hands”. Thus, the poeciloacanthaus

(“chainette”) families are Dasyrhynchidae,

Lacistarhynchidse, Hornelliellidae, Mustelicolidae,

Gymnorhynchidae, and Mixodigmatidae (see

Beverndge & Campbell 1989).

Dollfus’ attempts (1969a) to subdivide species of

Grillotia have been prompted by their rather

homogeneous armatures and varied combinations of

other scolex features, He created the subgenera

Grillotie, Progrillotia, are| Peregrillotia, Poragrillotia,

discussed and rejected above, remained 1 subgenus.

Progrillotie Dollfus. 46 was elevated to generic stanis

and o new genus and family, Pseudogrillotia

a4 R_ A. CAMPBELL & |. BEVERIDGE

|

WRENS

} WS

i

s

Figs 12-16, Pseudogrillotia spratti sp, nov.; tentacle armature; 12, metabasal armature, external surface; 13, metabasal armature,

external surface showing origin of principal rows; 14, basal region, internal surface on right side and bothridial surface

at left; 15, basal region, transition between the basal armature and metabasal region on the extemal surface; 16, lower

tiectabasal region, antibothridial surface showing single intercalary row of hooks. Scale lines: 12-16, 0.1mm.

NEW SPECIES OF GRILLOTIA AND PSEUDOGRILLOTTA 45

(Pseudogrillotiidae) were added by Dullfus (1969a).

Schniidt (1986) included jhe genus Grillotia and

Pseudogniloria in his key but omitted Progniitoria and

the family Pseudogrillotidae. We concur with Schrult

(F986) m considenng a craspedote scalex and postlarval

Stage insufficient to warrant separation of

Preudogrillotia into another family. We have found no

Paper in which Dolifus ever formally defined the family

Grillotiidae but he did comrast the Grillotidac as well

as the subfamily Grilloliinae (Dollfus 1969a) in the

justification for creating Psevdegritiotia and the

Pseudogrillovidae.

Despite the incrensing number of species described

and the varlabiliy en the combination of characters now

apparent, Dollfits' treatment of the group has remained

basically unchanged since 1969. The diverse

combinations of characters and taxonomic importance

previously attributed to them must be questioned in

view of species recently described. The various

combinations include patelliform versus non-

patelliform bothridia, craspedote ur acraspedete scolex,

presence or sbsence of prebulbar organs, long or short

bulbs, vanous origins of the retractor muscle, presence

or absence of a postlarval stage between the blastocyst

stage and adult, presence or absence of a basal

armature, presence Or absence of a demarcation

between the intercalary hooks and the longitudinal band

of hooks on the external surface, and ¢ven the

interpretation of the presence of a Jongitudinal “band

of hooks for some species, For the reproductive system

the details of the terminal genitalia are not well known

but a hermaphroditic sac, combined with an accessory,

external and internal seminal vesicles have been shown

10 occur in several species, the ovary may be terminal

or subterminal in position, and postovarian testes are

present in some species and facking in others. As

proposed by Dollfus (1942) the sub-family Grillotiinae

is justified as separate from thie Lacistorhynchinae in

the Jack of a chainette, presence of a longitudinal ‘band™

of hooks in the metubssal armature, and présence of

intercalary rows of hooks in the armature. These same

characters ate implied as supporting elevation Ww family

mink (Dollfus 19694). Complete revision of the family

Grillotiidac should be made based upon examination

af types of all species. The diversity of characters and

combinations is not unique to Grillotia, but many of

these same combinations can be found in the

Gilquiniidse (see Beveridge 1990). We propose that

they be accommodated by including them in a formai

diagnosis of the family and recognised separately from

the Lacistorhynchidae as folinws-

Grillotiidae Dollfus, 1969 emcended

Diagrasis; Scolex acraspedate or craspedote, Two

wite bothndia, cordiform or patelliform, with or

withomt thickened rims, posterior and lateral margins

free. Principal hook rows alternate, Intercalary rows

vf hooks present, irregular band of small hooks

between termination of principal rows in metabagal

armature. Basal armature preseal or absoiit, Hooks

heleramorphous, hollow, Pars vaginalis long, tentacle

aheaths sinuous. Bulbs long or shart, prebulbar organs

present of sbsent. Retractor muscle originates in

posterior half or bulb. Segments acraspedote, apolytic,

Mature segments elangated, Genital pores marginal,

irregularly alternate, Accessory seminal vesicle,

internal seminal vesicle and bermaphomditic duct

present in type species. Hermaphroditic duct, extemal

amd intemal seminal vesicles reported for some species.

Testes medullary, some usually postoyarian, Ovary

usually separated from posterior segment margm. by

a-space. Vitelline follicles form a continuous sleeve

around the anternal organs. Uterus saccate with Lateral

diverticular. Procercoid in copepods, plerocerci and

postlarvae in teleosts, adults in elasmobranchs.

Type genus: Grilloria Guiart, 1927

Diagnosis; Grillotiidac, acraspedote scolex;

hothridia indented on posterior border, longitudinal

“band” of hooks continuous, intercalary rows of houks

may merge with “band”, special basal armature absent,

prebulbur organs present, hennaphroditic sac present,

imternal and external (or accessory) seminal vesicles

present, ovary well forward, postovarian testes present.

Other genera; Progriliotia Doltfus, 1969

Diagnosis. Grillouiidae, acraspedote scolex,

bothridial margins not indented posteriorly, prebulbar

organs lacking, longitudinal “band” of hooks interrupted

Opposite each principal row, special basal armaature may

be present, bulbs very long, prebulbar organs lacking.

testes in longitudinal tows, accessory seminal vesicle

absent, ovary at posterior extremity of segment,

posiovarian testes absent. Type species: Pr. pastinacae

Dollfus, 1946,

Pseudogrillona Dollfus, 1969

Diagnosis: Grillotiidae, craspedote scolex, bothridia

indented posteriorly, prebulbar organs preseril,

longitudinal “band” of hooks conlinwous but may be

restricted io basal region, special basal armature

absent, hermaphraditic duct present, testes nicemose,

ovary well forward of posterior margin, postyaginal

testes present, posularval stage im life cycle. Type

species: P pleistaccentha Dollfus, 1969.

Acknowledgments

Thanks are due to Mr B. G. Roberson aid Dy

DB. M, Spratt for collecting specimens, and to te

Australian Biological Resources Study for financial

Suppor.

46 R. A. CAMPBELL & I, BEVERIDGE

References

Beumer, J. P., ASHBURNER, L. D,, Bursury, M, E., JETTE,

E,, & Latuam, D, J, (1982) A checklist of the parasites

of fishes from Australia and its adjacent Antarctic

territories. Commonw. Inst. Parasitol. Techn. Comm. No.

48 (Commonwealth Agricultural Bureaux, Farnham Royal,

England.)

BEVERIDGE, I. (1990) Revision of the family Gilquiniidae

Dollfus (Cestoda: Trypanorhyncha) from ¢lasmobranch

fishes. Aust. J. Zool. 37, 481-520.

& CampsBeLt, R. A. (1988) Cetorhinicala n.g.,

Shirleyrhynchus n.g. and Stragulorhynchus n.g., three new

genera of trypanorhynch cestodes from elasmobranchs in

Australian waters. Syst. Parasitol. 12, 47-60.

—___ & __ (1989) Chimaerarhynchus ng. and

Patellobothrium n.g., two new genera of trypanorhynch

cestodes with unique poeciloacanthous armatures, and a

reorganisation of the poeciloacanthous trypanorhynch

families, Jbid, 14, 209-225,

Cama, J. & Gavarrino (1989) Grillotia similis (Linton,

1908) comb, n. (Cestoda; Trypanorhyncha) from nurse

aback in the Florida keys. J. Helminthol. Soc. Wash, 57,

15-20.

CAMPBELL, R, A. (1977) New tetraphyllidean and

trypanorhynchan cestodes from deep-sea skates in the

western north Adantic. Proc. Helminthal. Soc.. Wash. 44,

191-7,

CaRVAIAL, J,, CAMPBELL, R. A. & CORNFORD, E. (1976).

Some trypanorhynch cestodes from Hawaiian fishes, with

descriptions of four new species. J. Parasitol, 62, 70-7.

Doirus, R. P. (1942), Etudes critiques sur les tetrarhynques

du Museum de Paris. Arch. Mus, Natl. Hist, Nat. (Paris)

19, 1-466,

(1946) Notes diverses sur les Tétrarhynques. Mém-

Mus. Natl. Hist, Nat., 22, 179-220,

(1969a) Quel ques espéces de cestodes tetrarhynques

de la cote Atlantique des Etats Unis, dont l'une n’etait pas

connue a l'état adulte. J, Fish, Res. Bd Can. 26, 1037-61.

(1969b) De quelques cestodes tétrarhynques

(heteracanthes et pécilacanthes) recoltes chez des poissons

de la Mediterranee. Vie et Milieu, ser. A 20, 491-542.

ESCALANTE, H. & CarvalaL, J. (1984) Larval trypanorhynch

cestodes from Peruvian teleost fishes, with descriptions of

two new species. Stud. Neotrop. Fauna Environm. 19,

185-194.

SAKANARI, J, (1989) Grillotia heroniensis, sp. nov., and G.

overstreeti, sp. nov., (Cestoda; Trypanorhyncha) from Great

Barrier Reef Fishes. Aust. J. Zool. 37, 81-7.

Scumipt, G. D. (1986) ‘CRC handbook of Tapeworm

Identification’ (CRC Press Inc,, Florida.)

SHAHAROM, F, M. & Lester, R. J. G. (1982) Description and

observations on Grillotia branchi n- sp., a larval

trypanorhynch from the branchial arches of the Spanish

mackerel, Scomberomorus commersani. Syst. Parasitol. 4,

1-6.

YAMAGUTI, 8, (1959) Systema Helminthum. IL. The Cestodes

of Vertebrates, (Interscience Publishers, New York).

ABUNDANCE AND LIFE HISTORY OF NATIVE AND INTRODUCED

EARTHWORMS (ANNELIDA: MEGASCOLECIDAE AND LUMBRICIDAE)

IN PASTURE SOILS IN THE MOUNT LOFTY RANGES,

SOUTH AUSTRALIA

BY G. H. BAKER*, V. J. BARRETT*, R. GREY-GARDNER** & J. C. BUCKERFIELD**

Summary

Seasonal abundance of the native earthworm, Gemascolex walkeri (Megascolecidae), and

introduced earthworms, Aporrectodea rosea and Octolasion cyaneum (Lumbricidae), was

monitored in three pastures in the Mt Lofty Ranges, South Australia. Highest numbers of

earthworms were recorded in late winter and early spring. Densities of up to 108, 32 and 96 m”

were recorded for G. walkeri, A. rosea and O. cyaneum respectively. All three species of

earthworms occurred predominantly in the top 10 cm of soil for four to five months per year

(autumn to spring), when soils were wettest. During other seasons, they were found lower in the soil

profile. Distributions within pastures were patchy and could, in some cases, be explained by soil

type (A. rosea) and soil moisture (G. walkeri). Although introduced species of earthworms

generally predominate in pastures in southern Australia, native species such as G. walkeri can

occasionally constitute a substantial proportion of the total earthworm population.

KEY WORDS: Earthworms, Megascolecidae, Lumbricidae, pastures, South Australia,

depth profile, life history.

Transactions of the Royal Society of 5. Anst., (1993), Ut, 47-33,

ABUNDANCE AND LIFE HISTORY OF NATIVE AND INTRODUCED EARTHWORMS

(ANNELIDA; MEGASCOLECIDAE AND LUMBRICIDAE) IN PASTURE SOILS IN THE

MOUNT LOFTY RANGES, SOUTH AUSTRALIA

by G. H. BAKER* V. J. BARRETT* R. GREY-GARDNER* & J, C. BUCKERFIELD™

Summary

Haxer, G. H., BARRETT, V. J, Grey-Garpner, R, & Buckerriecp, J, C. (1993) Abundance and lile history

of native and introduced earthworms (Annelida ; Megascolecidae und Lumbricidae) in pasture soils in the Mount

Lofty Ranges, South Australia Trans, R. Soc. S. Aust. 147(1), 47-53, 4 June, 1993.

Seasonal abundance of the native earthworm, Gemascolex walke'rt (Megascolecidae), and introduced earthworms,

Aporrectadea rosea and Octolasion cyaneum (Lumbricidae), was monitored in three pastures in the Mt Lofty

Ranges, South Australia. Highest humbers of earthworms were fecorded in late winter and eurly spring. Densities

of up to 108, 32. and 96 mr were recorded for G. walkeri, A. rosea and O: cyaneum respectively.

All three species of earthworms occurred predominantly in the top 10cm of soil for four to five months per

year (autumn to spring), when soils were wettest. During other seasons, they were found lower in the soil profile.

Distributions. within pastures were patchy and could, in some cases, be explained by soil type (A. rosea) and

soil moisture (G_ Wwalkeri).

Although introduced species of carthwortns generally predominate in pastures in southern Australis, native

species such us G, walker! canoccasionally constitute a substantial propartion of the total earthworm population,

Key Worbs: Earthworms, Megascolecidae, Lumbricidae, pastures, South Australia, depth profile, lite history.

Introduction

Australia has a rich native earthworm fayna,

dominated by the Megascolecidae (Jamieson 1981).

Howeves, information on the life histories, distributions

and abundances of these native species and the factors

influencing them is scarce (Wood 1974; Jamieson 1931;

Abbott e¢ al, 1985; Baker et al, 1991, 1992a). The

distributions of introduced species in Australia (e.2.

Lumbricidse), are better understood (Abbott 1922,

1985; Abbow & Parker 1980; Tisdull 1985; Kingston

& Temple-Smith 1989; Baker ey af, 1991, 1992a), and

there have been some studies of their life histories and

population dynamics (Barley 1959a;, Tisdall 1985;

Rovira er al, 1987; Kingston 1989; McCredie er al.

1992; Baker e¢ af 1992b). The influence of

earthworms, cither mative or introduced, on soil

structure ated fertility in Australia has received little

study (Barley 1959b,c, Barley & Jennings 1959; Abbott

& Parker 1981), Increased knowledge of the ecology

und Functional importance of the resident communities

of earlhworms in Australian soils will assigt in

developing improved management strategies that may

lead to increased plant production and help reduce soil

degradation (Baker 19893,b).

Baker et ai. (1992b} demonstrated the seasonal

abundances and depth profiles of two introduced

* Divisions of Entumology* and Sails“, CSIRO, P.M. Bag

2, Glen Osmond 5064, South Australia.

species, Aporrectodea trapezoides ami A. caliginosa

(Lumbricidae), in five permanent pastures in South

Australia, A, trapezpides and A. caliginose were active

and abundant in the rvot zone for three to seven

months, from autumn to spring, when soi] moisture

was highest. Mature adults were present from mid-

winter to mid-spring, Patchy distributions of both

species were found within some of the pastures and

were partly explained by variation in soil type.

This paper reports the seasonal abundances, life

histories, lateral and vertical distributions and species

associations of native and intrxiuced earthworm

species that were found together with the mon

abundant 4, trapezaides and A. caliginosa at three of

the pasture sites sampled by Baker et af. (1992b).

Materials and Methods

Three permailent pastures in the Mt Lofty Ranges,

South Australia were sampled for earthworms each

month from March 1990 until November 1990

(Tungkillo) or January 1991 (Birdwood and Parawa).

The climate of the region is Mediterranean, with cool

moist winters and hot dry summers. Mean annual

rainfall yaricd between sites (Tungkillo, 575 mm;

Birdwood, 750 mm and Parawa, 900 mm) as did soil

type (Tungkillo, unitorm sand [Ucl,4]; Birdwood,

yellow and black duplex soils [Dy3.4 and Ddl.1) and

Parawa, yellow duplex soil [Dy2.2}) (Northcote 1979),

The yellow and black duplex soils are Palexeralfs and

the uniform sand is a Quartzipsamment in the LIS Soil

Taxonomy (Soil Survey Stalf 1987). Baker ef ail,

G. H. BAKER, V. }. BARRETT, R. GREY-GARDNER & J. C. HUCKERFIELD

(1992b) provide further information on chemical and

physical properties of the soits af each site.

At each site, soil samples were taken within « 50

x 30 m plot that was divided into five 10 m wide

strips. Each month, three random samples were taken

within each strip at Tungkillo and Birdwood and two

samples were taken within each strip at Parawa. Each

sample was 0.1m? x 30 cm deep. The samples were

separated into three depths, 0-10 cm (within which the

majority of plant roots occurred), 10-20. cm and

20-30 cm. The soil was hand-sorted for carthworms

and cocoons which were preserved in 70% ethanol.

Approximately 400 g of soil were collected From the

0-10 cm layer in one sample from each strip for

gravimetric moisture determination, These soil samples

were collected every month except March 1990, Curves

relating matric suctions of soil water to gravimetric

walet content wete calculated for the soils at each of

the three sites using the filter paper method (Greacen

et al, 1989), This enabled comparisons of soil moisture

avatlabilily between sites.

Species were identified within two weeks of

collection, using the keys in Jamieson (1974) and Sims

& Gerard (1985). Worms were separated into juveniles

(no genital markings), subadults (with genital markings

but no mature clitelhim) and adults (with mature.

clitellum).

Two soil types (yellow and black duplex soils)

occurred within the plot at Birdwood, The distributions

of these soils were mapped by taking 25 regularly

spaced auger samples (each 3 cm wide x 50 cm deep)

within the plot and supplementing these with additional

samples where the soil boundaries were unclear. There

were no obvious spatial variations in soil type wilhin

the other two plots,

The regular sampling at Tungkillo indicated that soil

moisture varied markedly within the plot, being highest

near the centre and éastern edge. This pattern was

possibly related to the drainage characteristics of the

site, the moistest soil belng found in the lowest lying

areas Within the plot. To check this perceived pattern

in soi] moisture, 22 auger samples (10 cr in diameter)

‘were taken at random co-ordinates throughout the plot

during March 199], soil being collected from 0-10,

10-20, 20-30, 30-40 and 40-50 cm below the surface.

Gravimetric soi] moistures were determined for each

depth.

Results

Abundance and Life History

Duta for the seasonal abundance and life history of

three earthworm species that wers commonly collected

a Tungkillo, Birdwood and Parawa are given in Figs

1 amd 2. Similar data for rarer populations of these

and other species found at the three sites are given in

Table 1.

ari

=

f

ia

Numbat per square metre

Number per square metre

Bo ws

3

Numberper square metre

i

s

oz = —— i ome

Mer Apr May Jun Jul Aug) Sep) Gel ow Der day

Manth

Fig. 1.. Numbers (m~2) of juvenile (7 ——_1), sub-adult

( Cruces ) and adult ( —— ) Gemascolex walken’

(Tungkillo) (a), Aporrectodea rosea (Birdwood) (b) and

Octolasian ¢yaneur (Patawa) (c) collected during 1990-91,

Standard errors for the total earthworm numbers (all life cycle

stages combined) are given for each month, * indicates the

presence of cocoons.

Gemascolex waikeri Jamieson (Megascolecidae) was

common at Tungkillo, not found at Parawa and

occurred in only negligible numbers at Birdwood (Fig.

1, Table 1). Aporrectadea rosea (Savigny)

(Lumbricidae) was most common.at Birdwood and was

also found at Tungkillo, Octolasion cyanewn (Savigny)

(Lumbricidae) was found at Parawa but not at the other

two sites. The abundances of G. walker, A. rosea and

cyareum varied between months (Kruskal-Wallis H

= 54.9, p < 0,001 for G, walkeri at Tungkillo; H =

42.3, p < {001 for A. rosea at Birdwood, H = 45,1,

Pp < 0.001 for O cyaneum at Parawa), with highest

numbers found from late winter to early spring (August

to September or Octoher) (Fig. 1). Other species were

penetally recorded in low numbers at the three sites

(Table 1). Gemtascolex lateralis (Spencer)

ABUNDANCE OF EARTHWORMS IN PASTURES 49

(Megascolecidae) was abundant at parawa in May and,

June 1990 (Table 1), but most of the individuals of this

species collected then were found in only one sample

in each month (53% of the total collection in May and

69% in June), There was no significant variation

between months in the abundance of G. lateralis at

Parawa (H = 8&6, p > 0,05).

Numbar per square metre

a Motsture

20

& Ss Ss fe

=

Number per square matte

ho a

hy MbIStUTE

S

an

="\3

%y Moisture

=

o

Number per square metre

oe

S

we

a

0 a _Teseait

Mar Apr May Jun

Jul Avg Sap Geb Noy Dec Jan

Month

Fig. 2, Numbers (m~*) of Gemascolex walkeri (Tungkillo)

(3), Aperrectodea rosea (Birdwood) (b) and Octolasion

eyaneurm (Parawa) (c) collected 0-10 cm ({[[__—_]),

10-20 cm (ea » and 20-30 cm ( MEE ) belo

the soil surface during 1990-91, Standard errors for the total

earthworm numbers (ail depths combined) are given for each

month. Mean gravimetric soil moistures for 0-10 cin deep are

also. indicated for each site (line).

Adults of G. waikeri and A. rosea were mostly

collected in early spring (Fig. 1), Adults of @. cyaneum

were present on most sampling occasions,

Earthworm cocoons were found at all three sites,

during each month fram July to October (up to 39

cocoons per soil sample).

Virtually all the cocoons (> 99%) were found in

the top 10 cm of soil. The identity of these cocoons

is unknown, Most of the cocoons probably belonged

to Aporrectodea trapezoides, the most abundant species

at these sites {Baker er al. 1992b). However, the

breeding seasons of all species can be regarded as

occurring within these months.

Depth Profile

G. walkeri, A, rosea and O. cyateum occurred

mainly in the root zone (0-10 cndepth) for four to five

months during autumn to spring, when soils were

wettest (Fig. 2) (Pearson ty = 0.98, Ty = 0.86 and

Ty = 0.94 for soil moistures and numbers of G.

walkeri (Tungkillo), A. rosea (Birdwood) and QO.

cyaneum (Parawa) respectively at 0-10 cm depth;

p <0.01 in all cases). Earthworms becime more

‘common in the root zone compared with lower depths

during May or June and less. so in October or

Noverober.

a OTunghllo

O Birdwood

4 Parawa

LN Suction (kPa)

n

Apr Way Jan dol fy Sop Nov Ber Jar

Month

Fig. 3. Matric suctions of soil water (kPa) at 0-10 cm depth

at Tungkillo (©) ), Birdwood ( 0 ) and Parawa ( A ).

The surface soil was moister in autumn (May) and

drier in spring (October).at Parawa compared with the

other two sites (Fig. 3). Proportionally more worms

were present in the top 10 cm of soil at Parawa in May

and less in October than at Tungkillo and Birdwood

(Fig. 2, Table {) (comparing the total earthworm

numbers at 0-10 cm and 10-30 cm depth at each site

for all species included in Table 1 and Fig, 2,37 =

70.65, p < 0,001 and x? = 96.91, p < ODO for May

and October respectively),

Distribution Within Plots

The distributions of some species at each of the three

sites were patchy. At Birdwood, A, rosea was most

common in the south-eastern comer of the plot whilst

Gemascolex lateralis and G, stirlingi occurred

predominantly in the north-eastern comer (Fig. 4), The

distribution of A, rosea approximated that of the black

duplex soils at Birdwood (Fig. 4) (x? = 16,25, p <

0901 where the samples with and without A. rosea

50)

TABLE 1.

G, H, BAKER, V. I. BARRETT, R. GREY-GARDNER & J. C. BUCKERFIELD

Numbers of earthworms collected in low numbers at three pasture sites in the Mt Lofty Ranges, South Australia.

Numbers in parentheses aré for earthworms in the top 10 cm of soil, * indicates adults were present and — that no samples

were taken.

Species Monti

M A M I I A 8 oO N D J Total

Aporrectodea rosea

Tungkillo 3 2 26 2 6 4* I 6* 10* - — 60

(7) (6) (4) (1) (6) G)

Microscolex dubius.

Tungkillo 4* 14+ 8* - = 26

(4 (14) (8)

Birdwood 2* 2

(2)

{* 1

Parawa (1)

Gemascolex stirlingt

Tungkillo 4 ‘1 = ~ 5

(2)

Birdwood 1 1 ad ] 6 1 M 1 i] 2 1 20

(5) (1) -

G. lateralis

Tungkillo 1 = a ]

(ly

Birdwood 5 4+ i 4 10 3 J 2 3 33

(1) (4) (10) (3) (I)

Parawa 3 6 §3* 78% 17* 14+ QF 3* 21" 6* 19* 229

(53) (78) (IT) (4) (9) {1)

G. walkert

Birdwood 3* 3 6

_ (3)

Native W

Tungkillo * | 1 J 2 — — 6

; 1) r)) (1) (2)

Native J

Birdwood ti i

within the areas with yellow and black duplex soils

were compared), but the distributions of the two

Gemascolex spp. could not be explained by variation

in soil type. —

At Tungkillo, G. walkeri was most abundant near

the centre and eastern end of the plot (Fig, 5a) (x? =

31.29, p < 0.001 where the frequencies of samples with

0, 1-5, 6-10 and > 10 earthworms inside and outside

the area in Fig. 5a were compared), The soil within

the area where G. walkeri was most common was

moister in late summer than the soil outside it (Table

2). During the wettest months of the year, the same

irend was also apparent. Mean soil moistures for the

samples taken at 0-10 cm depth during the routine

monitoring at Tungkillo from July to October 1990 -

were 17.2% (where G. walkeri was most common) and

13.1% (elsewhere) (t = 2.23, p < 0.05).

G. lateralis was mostly found in the south-eastern

corner of the plot at Parawa (Fig. 5b) (x7 = 62.05,

Pp < 0.001 where the frequencies of samples with 0,

15 and > 5 earthworms inside and outside the shaded

area in Fig. 5b were compared). There was no obvious

variation in habitat within the plor that might explain

this patchiness in G lateralis. However, several

Eucalyptus trees bordered the plot adjacent to the area

where G. lateralis was found. Perhaps these remnants

of natural vegetation ate responsible for local survival

of this native earthworm.

Discussion

The life histories, patterns in seasonal abundance and

depth profiles of G. walkeri, A, rosea and O. cyaneum

in pastures in the Mt Lofty Ranges, South Australia

are broadly similar to those reported previously for

A, trapezoides and A. caliginosa (Baker et al, 1992).

Earthworm activity in the pasture root zone. (0-10 cm

depth) is mostly restricted to winter and early spring

when soils are wettest. Adult earthworms are most

commonly found in late winter-early spring, but for

some species (e.g. O. cyaneuin and G. lateralis), adults

can be found during most months of the year.

The proportions of A_ rosea, O: cyaneum and

Gemascolex spp. populations that survive summer are

not known, but many of these earthworms have been

ABUNDANCE OF EARTHWORMS IN PASTURES S|

G.. lateralis

A, rosea BIRDWOOD

‘ ®

N

ao}-

a absent

20 a 71-5

+ 6-10

a >10

0 10 ra] 30 AG 50

Fig, 4, Distributions of Aporrectadea rosea. Gemascolex lateralis, G. stirlingi and soil typés at Birdwood. Earthworm numbers

are piven as0(0),1-5( Ml), 610 (+}and > 0( a

) per sample, Shaded areas indicate where all or the majority

of eaathworms Were found, Soil rypes were classified as yellow duplex (Dy), black duplex (Dd) and intergrade of Dy and Dd (1).

found in summer > 30 cm below the surface of the

soi] (G, Baker, unpublished data). The increase in

population numbers. that occurs in the surface layers

of the soi] in autumn and easly winter must be

explained by invasion of individuals from lower depths

rather than by breeding, since no cocoons were found

al that time.

Whilst pastures in the Mt Lofty Ranges are

dominated by introduced lumbricids, espevially

Aporrectodea spp. (Baker ef al, 19924), some native

megascolecids, such as G, walkeri, can be locally

abundant and constitute a significant proportion of the

tolai earthworm population. For example, G. wulkeri

constituted 40.1% of the total population collected at

Tungkillo in 1990 (Baker ef al, 1992b), [Most (54.8%)

of the earthworms found at this site were A,

trapezoides. | The contribution that native species make

to soil structure and fertility has yet to be resolved,

Experiments are currently being conducted to evaluate

the role of G. walkers at Tungkillo in tairying surface-

applied lime and nffsetting sail acidity:

A, rosea is one of the most widespread and abundant

earthworm species in South Australia and western

Victoria {Baker e al. 1991, 1992a). Multiple

regressions of the abundance of A. rosea against a

variety of environmental variables have shown that soil

particle size is statistically the most important regressor

for sites in the Mt Lofty Ranges, as is the case for the

other most common introduced species, A. trapezoides

and A, caliginosa (Baker et al, 1992a)., In general, the

abundance of these earthworms increases with % clay

content, Variations in soil particle size helped explain

differences in the abundance of A. trapezoides and A.

caliginesa at several farms including Tungkillo,

Birdwood and Parawa (Baker er al, 1992b). They may

also explain the differences in abundance reported hete

for 4. rosea at Tungkillo and Birdwood. A. rosea was

more abundant in the black duplex soil (19% clay) than

in the yellow duplex soil (7% clay) at Birdwood and

the uniform sand at Tungkillo (also 7% clay), The

absence of A, rosea from Parawa (where clay content

was 14%) is, hawever, not explained by clay content.

32 ti, H, BAKER. V, 1, BARRETT, R. GREY-GARDNER & J. C. BUCKERFIELD

TUNGKILLO

G. walkerl

PARAWA

G. lateralis

eered

im

4s

i

a 10 20 a ay ay

Fig. 5. Distributions of Gemascolex watkeri at Tangkillo and

G. tateralis at Parawa. Eorthworm number. are given as 0

(O), 15 (M), 6-10 (+), 1-20 ( @ ) and > 204 A)

per sample. Shaded areas indicate where the majority of

eanhworms were found.

©} cyaneum was restricted to Parawa, the wettest site

with an annual rainfall of 900 rom, compared with 575

and 750 mm at Tungkillo and Birdwood respectively.

Previous surveys (Baker e7 al. 1991, 19923) have found

O. evaneum at only a few sites in the highest rainfall

regions of South Australia and western Victoria. There

TABLE 2.

Gravimetric moistures (%) for soils collected at Tangkilla in

March 1991

Soils were collected at varying depths inside and outside the

area in which Gemascolex walkeri was most abundant (sec

Fig. 5}. Results of t tests for significant differences between

data are included.

Depth Soil Moastute-

(em) Inside Outside i Prob.

0-10 1,48 0.93 §.20 < 0,001

10-20 2.60 iv? 4.65 < 0.001

20-30 3,83 1799 5.00 < 0001

30-40 474 2.55 3.78 < O01

40-50 642 2.82 2.80 < 0.05

ate a few records of O. cyanewm in Western Australia,

all from the high rainfall region of the south-west

(Abbott 1981), Sims & Gerard (1985) comment that

O, cyaneum prefers moist habitats in Britain.

The distributions of the native species within-the

three sites were patchy, In some cases, possible reasons

for the patchiness can be offered. For example, G.

walker muy be restricted by soil moisture at Tungkillo.

Baker ef al. (1992b) teported minor levels of

parasitism af A. trapezvides and another introduced

earthworm, Microscolex dubius, at Tungkillo during

spring by an undescribed dipteran. G. walkeri was

more abundant than these other two species in the top

10 cm of soi] at the same time, but was not parasilised,

Acknowledgments

This research was funded in part by a grant from

the Australian Wool Research & Development

Corporation, We especially thank D, Hobbs, A. Philp

and V, Rathjen for the use of their land, Penny Carter

for much assistance with the earthworm collections,

Richard Merry for help with the pedology and soil

chemistry and Peter Williams for help with the soil

moistures,

References

Asner, 1. (198!) Twa species of lumbricid garthworm newly

recorded from Western Austratin, Rec, West Aust. Mus.

9, 273-277.

(1982) The distribution of earthworm in the Perih

metropolitan area, Jbid. 10, 11-34,

____ (1985) Distribution of introduced earthworms in the

northern jarrah forest of Western Australia. Aust, J, Soil

Res. 23, 263-270,

& Parker, C. A. (1980) The occurtence of

carthworms. in the wheat-belt of Western Australia in

relation to land use and. rainfall. Jbid. 18, 343-352.

& __ (1981) Interactions between earthworms

and their soil environment. Sail Biol. Biochem. 13, 191-197,

. Ross, J. S. & Parwsn,C, A. (1985) Ecology of the

lame indigenous earthworm Megascolex imparicystis in

relation to agriculture near Lancelin, Western Australia-

JR. Sac. W. Aust, 68, 3-15,

Baker, G. H, (1989a) Role for earthworms in ameliorating

soil acidity pp. 64-65, In K. J. Hutchinson (Ed.) “Biological

Processes and the Fertility of Tem and Mediterranean

Pastures.” (Aust. Wooi Corp. Tech. Publ.).

______ (1989) Potential for introduced earthworms. in

pastures in southern Australia. Ibid. pp, 68-70,

, Barrett, V. J., BUCKERFIELD, J. C., GREY

Garpner, R, & Carrer, P. J. (1991) Distribution and

abundance of earthworms in agncultural soils in South

Australia and western Victoria pp. 30-51. /n Ri G, Faweet

(Ed.) “Workshop on Tillage Systems, Rotations, Nutrition

and Associaked Root Diseases.” (S, Aust. Dept. Agnc. Tech.

Publ).

ABUNDANCE OF EARTHWORMS IN PASTURES 53

, GREY-GARDNER, R, & BUCKERFIELD, J. C. (1992b)

The life history and abundance of the introduced

earthworms Aporrectodeq trapezvides and A. caliginosa

(Annelida: Lumbricidae) in pasture soils in the Mount Lofty

Ranges, South Australia, Aust. J. Ecol. 17, 177-188.

; BUCKERFIELD, J, C., GREY-GARDNER, R., Merry, R.

& Douse, B. (1992a) The abundance and diversity of

earthworms in pasture soils in the Fleurieu Peninsula, South

Australia. Soil Biol. Biochem. (In press).

Bar ey, K. P. (1959a) The influence of earthworms on soil

fertility, I. Earthworm populations found in agricultural

Jand near Adelaide. Aust. J, Agric: Res. 10, 171-178.

(1959b) The influence of earthworms on soil fertility.

Ti. Consumption of soil and organic matter by the

earthworm Allolobophora caliginosa. Ibid. 10, 179-185,

(1959c) Earthworms and soil fertility, IV. The

influence of earthworms on the physical properties of a red-

brown earth. Ibid. 10, 371-376.

& Jennings, A. C. (1959) Earthworms and soil

fertility, O10, The influence of earthworms on the availability

of nitrogen, [bid. 10, 364-370.

GREACEN, E. L., WALKER, G. R. & Cook, P. G. (1989)

Procedure for the filter paper method of measuring soil

ae suction. CSIRO Division of Soils Div. Report No.

08, 7 pp.

JAMIESON, B. G. M. (1974) Earthworms (Oligochaeta:

Megascolecidae) from South Australia. Trans. R, Soc, S.

Aust. 98, 79-112.

—— (1981) Historical biogeography of Australian

Oligochaeta. pp. 887-921. In A. Keast (Bd.) “Ecological

Biogeography of Australia.” Vol. 2 (W. Junk, The Hague)

‘pp. 887-921.

Kinoston, T. J. (1989) Aporrectodea caliginosa and

Ligmbricus rubellus populations under irrigated and dryland

pastures in northern Tasmania. pp. 199-205. Jn P. P. Stahle

(Ed.) Proc. V Australasian Conf Grassland Invertebrate

Ecology (D & D Printing, Melbourne).

& TEMPLE-SMITH, M. G. (1989) Earthworm

populations under Tasmanian pastureland. bid. pp. 192-198.

McCrepiz, T. A., Parker, C. A., & Asport, I. (1992)

Population dynamics of the earthworm Aporrectodea

trapezoides (Annelida: Lumbricidae) in a Western

Australian pasture soil, Biol. Fertil. Soils 12, 285-289.

Norricore, K. H. (1979) “A Factual Key for the Recognition

of Australian Soils.” (Rellim Press, Adelaide),

Rovira, A. D., SmerTem, K. R. J. & Lee, K. E. (1987) Effect

of rotation and conservation tillage on earthworms in a red-

brown earth under wheat. Aust. J. Agric. Res. 38, 829-834.

Sims, R. W. & Gerarp, B. M. (1985) “Earthworms.” Syn,

British Fauna No, 31. (Linnaean Soc. London, London).

Sou. SURVEY Starr (1987) “Keys to Soil Taxonomy” Third

Printing. SMSS Tech Mon. No. 6. (Ithaca, New York),

TisDALL, J. M. (1985) Earthworm activity in irrigated red-

brown earths used for annual crops in Victoria, Aust. J.

Soil Res. 23, 291-299.

Woop, T. G, (1974) The distribution of earthworms

(Megascolecidae) in relation to soils, vegetation and altitude

an he slopes of Mt Kosciusko, Australia. J. Anim. Ecol.

, 87-106.

TRANSACTIONS OF THE

ROYAL SOCIETY

OF SOUTH AUSTRALIA

INCORPORATED

VOL. 117, PART 2

THE EARLY CAMBRIAN VOLCANICS FROM RED CREEK, EASTERN

MT LOFTY RANGES, SOUTH AUSTRALIA

BY C. G. GATEHOUSE*, J. B. JABOT, B. J. CLOUGH*® & A, J. MCCULLOCH*}

Summary

In the Red Creek area of the eastern Mt Lofty Ranges, Early Cambrian lavas, tuffs and

volcaniclastic sediments are interbedded with the top 270 m of the Heatherdale Shale, the top

member of the Normanville Group. Tuffs and volcaniclastic siltstones extend up into the basal 60 m

of the conformably overlying Carrickalinga Head Formation, the basal unit of the Kanmantoo

Group. Several lava flows of trachybasalts, one of which exhibits pillows, occur about 100 m below

the top of the Heatherdale Shale. The volcanics at Red Creek appear to be at a slightly higher

stratigraphic level than the Truro Volcanics, the type section of which is 24 km to the NNW of

Red Creek. It is proposed that the lavas at Red Creek represent a flow of hawaiite composition from

a central volcanic complex, closely analogous to that of the Truro Volcanics but not necessarily

from the same centre or erupted at exactly the same time. Geochemical data indicate that both the

volcanics at Red Creek and the Truro Volcanics belong to the same alkaline Early Cambrian

"within plate" volcanic province, which may be linked genetically with other Cambrian mafic

alkaline provinces such as the Yumali/Coonalpyn area. It is suggested that the closest tectonic

analogy for these provinces is that of a rifted continental margin as proposed for the Tertiary

intraplate yolcanic province of eastern Australia.

KEY WORDS: Early Cambrian volcanics, hawaiite, Mt Lofty Ranges, Normanville Group,

Kanmantoo Group

Jransuctions of the Koya! Society of S. Awst, 0993), 117(2), 57-66.

THE EARLY CAMBRIAN VOLCANICS FROM RED CREEK,

EASTERN MT LOFTY RANGES, SOUTH AUSTRALIA

by C. G, GaTenouse*, J, B. Jacof, B. J. CLouGcH#* & A. J. MCCULLOCH*?

Summary

GateHouse, CG, Jaco, J. B., Clouex, B. I. & MeCuLvocn, A. J. (1993) The Early Cambrian volcanics from

Red Creek, eastern Mi Lofty Ranges, South Australia. Trans. R. Soc. S. Aust. 117(2), 57-66 4 June, 1993,

In the Red Creek area of the eastern Mr Lofty Ranges, Barly Cambrian lavas, tule and volcaniclastic sediments

ame interbedded with the top 270 m of the Heatherdale Shale, the top member of the Normanville Group. Tufts

and volcaniclastic siltscanes extend up ivito the basal 60 m ot the conformably overlying Carrickalinga Head

Formation, the basal unit of the Kenmunioo Group. Several lava tlows of trachybasalts, ane of which exhitits

pillows, occur about 100 m below the top of the Heatherdale Shale: The volcanics a) Red Creek appear to be

al Slightly higher stratigraphic level than che Truro Volcanics, the type section of which is 24 kim in the NNW

af Red Creek, It is proposed that the lavas at Red Creek represent u flow of hawaiite composition from a central

volcanic complex, closely analogous to that of the Truro Volcanics but nol necessarily from the same centre or

erupted at exactly the samne time, Geochemical data indicate that both the volcanics at Red Creek and the Truro

Volcanics belong to the same alkaline Early Cambrian *within plate" volcanic province, which may be linked

genetically with other Cambrian mafic alkaline provinces such a8 the Yurnali/Coonalpyn area, [t is suggested

that the closest tectonic analogy for these provinces is that of a rifted continental margin as proposed for the

‘Vertiary intraplate volcanic province of eastern Australia.

Key Woros: Early Cambrian volcanics, hawailte, Mt Lofty Ranges, Normanville Group, Kanmantoc Group

Introduction

This paper is a progress report on work an the Early

Cambrian volcanics generally known as the Truro

Volcanics of the eastern Mt Lofty Ranges. The Truro

Volcanics are important in chat, although they are

limited both in stratigraphic and geogmphic extem, they

represent the most extensive outcrops of yoteanics in

the Cambrian sequences of the Stansbury

Basin/Kanmantoo Trough area. As described helow,

voleanics in the Red Creek area extend From wuhin

the Heatherdale Shale, the top member of the

Normmanville Group, up into the Carrickalinga Hew

Formation, the basal member of the Kanmantoo Group,

Hence « study of the Truro Volcanics is important in

the determination of the stratotectonic setting of the

Kanmantoo Group transition which has been

interpreted in various ways as set out below,

You der Borch (i980) considered the Kanmantoa

Group sédiments to be the initial phase of fully

developed continental margin sedimentation which

typified the easiern flank of cratonic Australia during

most of the Palneazoic, Tamer & Foden (1990)

suggested that this phase was marked by renewed mafic

igneous activity in the form of the Truro Volcanics-

The basal part of the Carrickalinga Head Formation

ar Carrickalings Head lles north of Normanville where

von der Borch (980) proposed that the Houghton

Antictinal Zone acted a5 a separating feature between

*S.A. Dept of Mines and Energy, P.O. Box 151. Eastwood,

S.Aust, 5063

+ Dept of Applied Geology, Gartrell School, University of

South Australia, The Levels, S,Aust, S095,

shallow-water sediments to the west and continental

slope and rise environments to the cast, Scheibner

(1986, Fig. 4) Implied that the Kanmantoa Group

sediments were at least in part deposited on oceanic

crust and that che “Kanmantoo Trough” was part of a

marginal sea, extending eastwards to the Stavely

volcanic belt of westem Victoria. Parker (1986)

suggested that the Kammantoo Group sediments were

probably deposited in an extensional tectonic regime

on a marginal shelf reflecting tectonic activity to the

west and a. continental marpin to the east. Powell (1990)

regarded the “Kanmantoo Fold Belt’ as representing

the westernmost part of the “Tasman Pold Belt”, He

suggested that the Kanmantno Group represents passive

margin sedimentation.

The term Truro Volcanics was defined by Forbes er

ai. (1972) from a type section in Levi Creek, UL km

north of Truro (Fig. ), They: included two units within

the Truro Volcanics (Fig, 2), with the lower unit

comprising a 240 m thick sequence of interbedded

volcgnictastic sediments, amygdaloidal volcanics and

limestones. The upper unit as described by Forbes et

al. comprises 60m of marble, shaly marble,

metasiltstane and a thin {2 2) harizon of parphyritic

andesite, Forbes et ef, (1972) noted that this sequence

is overlain by the Hestherdale Shale which, near the

base, contains clasts of volcanic rocks. However, the

exact nature of the contact is unclear due to poor

exposure. The Heatherdale Slrale, the txp unit of the

Normanville Group, is overlain by the Carrickalings

Head Formation, the basal wnit of the Kanmantoo

Group. Furbes ef al. (1972) suggested thar che marble

in the upper member of the Truro Volcanics is

3k C. G. GATEHOUSE, J. B. JAGO, B. J. CLOUGH & AJ. McCULLOCH

FNL ABBEWEIT

}™ “4 ia ret

|

at

Mr ster 1 4 1 ag

! Trura e —

if Cay

Sa ca Hill

sedan,

e

AADELMIDE

a" fe

SE Br’ O02 4

Ne

SUBSURFACE DISTRIBUTION S \

Ye

Metadalerités ye

Ye

NA

Bosalt - andésite

Waster’ limit) oF

Murray Basi

fe

KILOMETRES

LOCALITY

\ —

5 he

Sutherlands

Maculia

~

“sea GASTON

AQ _aedon Hill

wt om

ENLARGEMENT j Seen

Fig, 1. Locality Map. The-type section of the Truro Volcanics is 2 km south of Mt Rufus 1.

equivalent to the Fork Tree Limestone which at Sellick

Hill. conformably underlies the Heatherdale Shale

(Daily 1963).

Rocks assigned to the Truro Volcanies by previous

workers occur in outcrop only in the Karinya Syncline,

north-east of Adelaide as shown by Cobb & Farrand

(1984). A recent report which may extend the known

distribution of the Truro Volcanics is that of Polomka

(1988)! who reported float material of possible Truro

Volcanics about 2.km NNE of Moculta, near the base

of what he mapped as Carrickalinga Head Formation.

However, the exact stratigraphic position of these rocks

is doubtful because of the poor outcrop in the arca,

Polomka described the rock as comprising fine to

medium grained phenocryst pseudomorphs of epidate

(2%), within a very fine actinolite (45 %) and epidote

(30%) groundmass.

'Polomka, S. M. (1988) The geology of the Mt Karinya area,

(South Australian Institute of Technology, Departinent of

Applied Geology, Unpublished thesis).

2Cooper, B. 1. & Gatehouse, C. G. (1988) Sedan Hill,

Carrickalinga Head Formation. Ia Gatehouse, C. G,

(compiler) Kanmantoo Field Symposium Excursion Guide.

S. Aust, Dept Mines & Energy Kept Bk 88/35,

3Gatehouse, C, G,, Jago, L B. & Clough, B. I. (1991a) 4

progress report on a measured reference section at Red Creek

for the Kanmantoo Group in the Karinya Syncline. §. Aust.

pert Mines & Energy Rept Bk 91/27.

§MicCullech, A. J. (1990) The gealogy of the Towitta area.

(South Australian Institute of Technology, Department of

«Applied Geology, Unpublished thesis).

5Van der Stelt, B. (1990) ‘The geachemisiry, petrology and

tectonic setting of the Truro Volcanics, (University of

Adelaide, Unpublished B.Sc. (hons) thesis).

5Gaichouse, C. G., McCulloch, A. J., Clough, B. J. &

Sarunic, W, (19910) Mi Rufus 1 Well Completion Report.

3° Aust. Dept Mines & Energy Rept Bk 91/25.

TRankin, L.. R., Clough, B. J. & Gatehouse, C. G. (1991a)

Mafic suites in basement beneath the Murray Basin: new

data for the Early Palaeozoic history of the Tasman orogenic

gprovince. S. Aust. Dept Mines & Energy Rept Bk 91/44

Rankin, L..R.. Clough, B. J,. Farrand, M. G., Bartek 8.

R., Lablack, K. Gatehouse, C. G. & Hough, L . P. (1991b)

Murray Basin’ basement transect project: 1990 well

er i reports, §. dust. Dept Mines & Energy Rept BR

125,

EARLY CAMBRIAN “TRURO VOLCANICS" 59

TRURO VOLCANICS TYPE SECTION

DESCRIPTION

CUMULATIVE

THICKNESS

(metres)

ROCK UNIT

Phyllite:

Conglomerate

and breccla: ... calcitio; of volcanic Iragments, scoriaceous, pebbles of

porphyritic andesite.

HEATHERDALE SHALE

shaly in part, more massive near base with

calcareous siltstone and thin interbeds ot dark volcanic rock

with large feldspar phenocrysts (position of volcanics

uncertain).

FORK TREE

LIMESTONE

Volcanic rock: .. pinkish-grey. Weathered.

Breccla: grey, amygdaloidal voicanic fragments, calcite matrix.

Limestone; ..-- grey, cream - weathering.

Voleanics: ..... green, phyliitic.

Voleanics: ..... shaly.

Exposure poor.

VOLCANICS

Volcanics: on top of calcareous greywacke?

Green volcanics: with minor grey amygdaloidal volcanics, conglomerate at top of

volcanic pebbles.

Limestone: ,... pale graen-grey. Falds plunge approximately 50° to south.

Volcanic rocks: . green apidote, calcite veined.

Volcanic breccia: im thick of 120mm clasts of vesicular volcanics.

Volcanic rocks: . metabasalt?, minor grey amygdaloidal volcanics.

Volcanic rock: . . amygdaloidal, calcita vains; amygdales 7mm long, calcite-filled.

Elongation trends 170°.

Volcanic rock: . . fine grained.

No outcrop.

Green phyllite, gray limestone, chert,

Silty phyllite, bioturbated.

pale grey, medium-grained laminated quarizite.

Fig. 2. Stratigraphic section al type section of Truro Volcanics, {1 km north of Truro (after Forbes er al, 1972),

60 C. G. GATEHOUSE, J. B, JAGO, B. J. CLOUGH & A.J. McCULLOCH

At the extreme eastern edge of the Mt Lofty Ranges, Sutherlands (Forbes ef al. 1972), The best exposure

Early Cambrian volcanics have been recorded at Sedan of Early Cambrian volcanics from the eastern Mt Lofty

Hill (Cooper & Gatehouse 1988”; Gatehouse et al. Ranges are those mapped in Red Creek by Coats &

1990), at Red Creek (Gatehouse et al. 1991a*; Thomson (1959) as feldspar porphyrite. The remainder

McCulloch 1990*; Van der Stelt 1990°), of this paper deals largely with the Red Creek area,

Accomodation Hill (Forbes er al. 1972) and near north-west of Sedan (Fig. 1).

Measured section (m}

|

1

'

es

S

3

KANMANTOO GROUP

Carrickalinga Head Formation

KANMANTOO GROUP

Cartickalinga Head Fmn

NORMANVILLE GROUP

Heatherdtale Shale and Truro Volcanics

wip

3

5

o of

sc

o>

ce

Oe

TT ie

do

2a

=

Fag

o ®

x

acd ©

2

a

=

COWG_LOMERATE

Slistone

Sandstone

Shale

Liensstone

Fippies

Velcanics

Crans Inmination

BHALE

FINE SAND

MEDIUN SAND f-

COARSE SAND

NE IMPAATE

Pillow java Rack sampie ipcoton

SILTY

SADME 92-1044

Fig. 3. Stratigraphic section. at Red Creek.

EARLY CAMBRIAN YOLCANICS 4it

Stratigraphic setting

Detailed measured sections at Sedan Hill (Cooper

& Gatehouse 988) and at Red Creek (Gatehuuse er

al, I99la*) show that volvanic rocks, generaily

referted wo as the Truro Voleanics are interbedded

within the Heatherdale Shale and cogiinue imo the

basal paris of the gradationally overlying Carrickalinga

Head Formation. The Red Creek section has been

logged an some detail (Gatchouse et.al. 1991b%); only

a summary of the section is given here (Fig, 3).

About 300 m of Hextherdale Shale is exposed at Red

Creek; the base is not exposed due io the presence of

the Palmer Fault. As exposed at Red Creek the

Heatherdale Shale is a pale grey larninated phyllite

which becomes darker and less micaceous up-section.

There are minor calcareous siltstone honzons, Within

these units are thin tuffaceous and/or volcanogenic

siltstone horizons which become increasingly abundant

up-section

About 200 m above the base of the measured section

are several basaltic lava flows, one of which exhibits

pillows (Fig. 4). The pillowed flow has an uneven base

and cuts down into an underlying 0.2m thick tuff

horizon; it appears to thin to the south. It contains

several pillow strictures with chilled margins and triple

junctions (Fig. 4). An intrusive igneous body seen

between 145 and 150 m on the measured section may

represent 2 feeder pipe to the lavas higher in the

section, Above the pillow lavas the tuffaccous/

voleanogenic siltstone horizons continuc with reduced

Frequency into the basal 60 m of the gradationally

overlying Carrickalinga Head Formation. Detailed

descriptions uf the complete stratigraphic section will

be given in a later paper.

In the north branch of Red Creek (Fig. 3), close to

the buse of the Carrickalings Head Formation, there

are several beds, and/or blocks, of crystal tuff

comprising almost pure feldspar crystals clearly

winnowed from-enclosing ash. Such beds may have

Fi

Wig. 4+. Pillow lavas al Red Creek. Note the triple junction

al centre left.

formed by current activity at the site of deposition or

by differential sir-fall separation,

Regional stratigraphic interpretation

As noted above, tm Red Creek, lavas and

voleatogesic sediments extend thorugh the exposed

Heatherdale Shale up into the base of the Carrickalinga

Head Formation. However, near the type section (Fig.

2) of the Trure Volcanics 24 kin NNW of Red Creek,

the highest known volcunics are at least 200 m below

the top of the Heatherdale Shale (see imag in Forbes

et al, 1972. and Fig. 2 herein}, Indeed Forbes e ul.

(1972) suggest that the bulk of the volcanics Lit the type

area occurs below an equivalent of the Fork Tree

Limestone, which on Fleuricu Peninsula ties

conformably below the Heatherdale Shale, However,

sinoe this correlation with the Fork Tree Limestone

is made on lithological grounds only, and furthes, that

there are calcareous horizons within the Heatherdale

Shale both on Fleurieu Peninsula and at Red Creek,

then there may be some doubt about the correlation.

It could be argued (hat the marble of the cype area of

the Truro Volcanics, described as Unit C4 by Forbes

etal. (1972), is equivalent to the calcareous horizons

found in the Heatherdale Shale at Red Creek rather

than being equivalent to the Fork Tree Limestone,

However, if the correlation of the Unu C4 marble in

the type section to the Fork Tree Limestone is correct

then it Sugpests that the volcanics at Red Cresk are in

a higher stratigraphic position than those of the type

section and should not be referred in the Tro

Volcanics. In his discussion on the Stansbury Basin,

Gravestock (is press) has included all these volcanics

as Truro Voleanics. A further compheation is the

almost complete Jack of exposure of the basal 150 a3

of the Heatherdale Shale near the type section of the

Truro Voleanics. The recent drilling of Mt Rufus Na,

1 stratigraphic hole 2 fam north of the type section

suggested that there may be several unrecugnised faults

in the area (Gatehouse et al. '991b*). Until the

position is clantied the volcanics described herein sre

simply referred to as the volcanics fram Red Creek,

Elsewhere in the Stansbury Basin, green tuff beds

in the Parara Limestone may be correlated with the

Truro Volcanics of the Karinya Syncline (Gravestock

in press). A tuff bad from within the Heatherdale Shale

at Sellick Hill has been dated at 526+4 Ma (Cooper

«1 al. 1992), this tuff may also be equated with the

Truro Yolcanics or the volcanics at Red Creek o¢ with

neither,

Petrography of volcanics at Red Creek

In hand specimens the voleames in Red Creek have

4 grey fine vrained gtaund mass with distinctive large

a2 C. G, GATEHOUSE, J. B. JAGO, B. J. CLOUGH & A.J. McCULLOCH

(approximately 0.5 mm diameter) phenocrysts of

feldspar (up to approximately 25%) and minor iron

staining. The rocks show variable intensities of tectonic

foliation.

In thin section it appears that the phenocrysts are

predominantly of zoned plagioclase although

approximately 10% of the phenocrysts are of an alkali

feldspar. The groundmass is predominantly of acicular

feldspar laths producing a trachytic texture, The

tectonic fabric of these rocks varies from slight to

intense foliation, the feldspar phenocrysts having

rotated parallel to the foliation, Alteration is pervasive

with sericitisation of the feldspar and chlorite/iron-

oxide replacement of mafic minerals. Sporadic veinlets

of quartz and calcite cross-cut this tock, Petrologically

the lavas classify as trachytic basalt.

Geochemistry of volcanics at Red Creek

Two samples (6728 RS 1632 and 6729 RS 1515) of

massive pillow lava were taken from separate localities

FLEURIEU TRUAO

PENINSULA TYPE

SECTION

Dally 1983, ;

Daily & Milnas 1972 Modified after

Forbes et al, 1972

CAMPANA

CREEK

MEMBER

BLOWHOLE

CREEK

SILTSTONE

MEMBER

MADIGAN CARRICKALINGA

INLET HEAD

MEMBER FORMATION

HEAD

CARRICKALINGA

FORMATION

7

HEATHERDALE HEATHERDALE

SHALE SHALE

— ?

FORK TREE

LIMESTONE

FORK TREE

LIMESTONE

SELLICK HILL

FORMATION

TRURO

VOLCANICS

WANGKONDA

FORMATION

2.

MOUNT

TERRIBLE

FMN

> —

MOUNT

TERRIBLE

FMN

EQUIVALENT ?

near Red Creek and geochemically analysed for a

comprehensive suite of elements (Tables 1 and 2).

Several analytical methods were used, including:

ICP (acid digestion) — major elements

XRF — As, Ba, Bi, Sb, Sn, V, Zr

Atomic Absorption Spectrography — Ag, Cr, Cu, Ni,

Pb, Zn

Fire Assay — Au, Pt, Pd

ICP Mass Spectrography — Ce, Dy, Nd, Er, La, Eu,

Lu, Yb, ¥, Sm, Gd, U, Th, Sr, W, Ta, Mo, Nb, Ga,

Co, Cs, Rb.

The two geochemical analyses (Tables 1 and 2) of

the pillow lava reflect their described lithology, as

basaltic lavas. that have undergone preenschist facies

metamorphism. Elevated loss on ignition (LOD values

(average 11.5%) attest to alteration effects resulting in

hydration. Clearly the present chemical composition

of the lavas at Red Creek is not primary, as the volatile

content is significantly higher than in analogous fresh

rocks. This is to be expected from the presence of

SEDAN RED

HILL CREEK

Altar Gatehouse (this paper)

et af, 1990

CAMPANA

GREEK

MEMBER

BLOWHOLE CREEK

SILTSTONE MBR?

MILENDELLA LST MBA

MADIGAN

INLET

MEMBER

CARRICKALINGA

HEAD

FORMATION

CARRICKALINGA

HEAD

FORMATION

MADIGAN

INLET

MEMBER

HEATHERDALE

SHALE

with

VOLCANICS

HEATHERDALE

VOLCANICS

BASE CUT OFF BY FAULT BASE CUT OFF BY FAULT

Fig. 5. Correlation diagram of the Cambrian sequences of the Red Creek-Truro-Sellick Hill areas

EARLY CAMBRIAN VOLCANICS 62

Taste {. Geochemical analyses and CIPW norms, NB,

CIPW weight % norms are calculated from analyses

recalculated to 100% free of HO and CO,; ce is not

recalculated, Fe data is recalculated nsing a Fe,: Total Fe

cation cation of 0.20.

Major Elements in Percent CIPW Weigh % Norms

6728 RS 6728 RS

163255 163255

SiO, 42.30 4230 ab 1885 19.92

TiO, 242 224 or 1971 13.69

ALO, 1580 1440 an 1847 1531

Fe,0, 875 BOS ne 985 13.29

FeO di W823.

Mad 0.28 626 al 467 321

Meo 264 #332 mt 294 272

Cad 805 «885 il 532 4.95

NasO 378 450 O73

x6 288 (1.99

P.O, 0.78 60.66 Total 100.01 100.0)

HO+

H,0

cb,

Lol 10.80 12.30

Total 9848 0887 DE 4841 46.82

=not analysed, DI=Differentiation Inmlex

hydrous secondary minerals in these lavas. Similar

alteration of mafic lavas in the Victorian greenstone

belts (Crawford & Keays 1978) is.considered to have

caused hydration, along with slight addition of CO,

and Na,O accompanied by leaching of SiO,, CaO,

ALO, and K,O, However, the degree of chemical

change was considered to be minimal, and magmatic

trends Werte clearly visible. To minimise the effect of

hydration dilution and related chemical mobility, plots

using elements considered immobile during alteration

are used and analyses are recalculated to 100% volatile-

free prior to plotting.

The lava at Red Creek plot on the border between

phono-tephrite and tephrite-basanite close to the

basallic-trachyandesite field in the SiO, versus

Na,O+K,0 classification plot of Le Bas er al. (1986;

Fig, 6), In the Nb/Y versus Zr/TiO, classification plot

(Winchester & Floyd 1977; Fig. 7) using elements

considered immobile during alteration, the lavas

classify as alkali basalt.

To further define the alkaline basalt a classification

scheme devised for the Tertiary alkaline volcanics of

eastern Australia (Johnson 1989) based on CIPW

norms was utilised (Fig, 8). Using this classification

scheme the division between sub-alkaline and alkaline

mafic lavas is that alkaline lavas have <10% normative

hypersthene; on this basis the basalts in Red Creek

classify as alkaline and plot within the field of

hawaiites. The presence of considerable levels (9.85

and 13.22%) of normative nepheline suggests these

rocks are silica undersaturated. However, plots using

immobile elements (Fig. 7) suggest these lavas have

not attained silica undersaturation but show that the

lavas at Red Creek and the Truro Volcamics at Mt Rufus

1 (Gatehouse et af. 1991b*) belong to 4a group of

analyses that straddle the boundary between alkali

basalt and silica undersaturated nepheline/basanite

fields.

The analyses agree with field evidenve suggesting

that the lava at Red Creek represents a single thin

submarine Java sequence from a common source, in

that they plot close together on all classification plots.

The lavas plot outside the tectonic discrimination

fields of Pearce & Cann (1973; Fig. 9), although closest

to the intraplate field; and in marked contrast to the

MORE-telated metadolerites that occur in the Murray

Basin basement (Rankin et al. 19912, b74) and ag sills

and dykes in the Mt Lofty Ranges (Liu & Fleming

1990; Rankin ef af. 1991a”). On a MORRB nommalised

spidergram (Fig, 10) the Red Creek lava shows a

distinctive tread Of elemental enrichment relative to

MORB values of the incompatible eletnents from Sr

through to Ni, which is typical of the more silica

TABLE 2. Trace elements in ppm.

6723 RS 6729 RS Detéction

4632 155 lienit (ppm)

Ap <1,00 < 1,00 1

As 64.00 72.00 2

Aw 2,00B 7.008 iB

Ba B80 610 id

Bi 4,00 <4.00 +

Ce 82.00 73.00 Ot

Co 29.00 44,00 I

Cr Ito LOS 4

Cs 1,30 t.20 2

Cu 6.00 $5.00 2

Dy 3.00 7.20 0.1

Er 3.90 3.20 O1

En 2.80 2.50 O.1

Ga 26,00 20.00 1

Ga 9.00 8 60 a4

La 38,00 36,00 a1

Lu 0,60 0.50 0,1

Mo 4.00 3,55 o.s

Nb 76,00 66.00 os

Nd 44.00 42.00 0.1

Ni 52,00 54.00 4

Pe 12.00 $2.00 4

Pa 1.008 <=1.00B LB

Pt <5,00B <5.0)8 5B

Rb 22,0 60.00 0.2

Sb <4.00 4.00 os

Sm 9.40 a OL

Sn <4.00 10.00 4

Sr 240 255 0)

Ta 3.00 2,20 02

Th 4.80 3.60 0.1

U 2.60 2.20 aa

v <5.00 <5,00 s

WwW 3,00 3,00 ?

Y 34.00 29.5 |

Yb 3.70 3,00 OL

Zn 22,00 76.00 3

Zr 380 320 +

o

undersalurated alkaline mafic layas of such provinces

as the Tertiary eastern Australian volcanic province

(Johnson 1989); note the marked contrast of the alkaline

intraplate lavas with MORB related metadolerite from

MB-12 in the Murray Basin basement (Rankin et al.

199la’ and b*).

Other alkaline volcanic areas similar to Red Creek

include the Yumali/Coonalpyn area, the Peebinga-1

16-00

12-BO

a 60

6-40

Naz0 + K20%

320

Tt

4av0 82-40

35-00 BOG 580 7TDO

SI02%

A.,...,,.Hed Creek matic tava

Fig. 6. Red Creek lavas on port of classification plot of Le

Blas et al, (1986). Values calculated to 100% volatiles free.

Zr/Tio;

i]

Rhyalite

—

ae

Ahyodacite/Dacila

04

Tag A, Trachyandesite

eS.

Sub-alkaling Basalt

0.004

Nb/Y¥

12] Mt Rufus’ 1

© Red Creek

Fig. 7. Red Creek and Mt Rufus L lavas on part of

classification plot of Winchester & Floyd (1977).

Cc. G. GATEHOUSE, J. B. JAGO. B, J. CLOUGH & AJ. McCULLOCH

(MBT-1) drillhole into the northwestern termination of

the magnetically defined Mt Stavely Belt (Rankin ez

al. 1991a’, b), and the Truro Volcanics as defined in

Mt Rufus 1 drillhole (Gatehouse et a/. 1991b°: Fig.

10). Of the other volcanics the nearest are the Truro

Volcanics that crop out on the west limb of the Karinya

‘Syncline 22 km NW of Red Creek; the lavas at Red

Creek represent a more evolved lava type than the

Truro Volcanics in their type section, resulting in a

line more distant from’ MORB in Fig. 10. This chemical

difference is seen in the petrology where the lavas at

Red Creek are distinctively porphyritic with

phenocrysts of alkali feldspar.

The Red Creek analyses are compared with a classic

continental intraplate voleanic suite in Fig. 11, where

they are normalised to Karoo type basalts, the lavas

at Red Creek and the other Cambrian alkaline volcanics

noted above exhibit enrichments relative to Karoo basalt

of the elements Sr‘to Ti, with Red Creek being one

of the most chemically evolved suites. This supports

the field relations which suggest that these lavas are

not directly analogous to the thick piles of dominantly

tholeiitic basalt fissure lavas of the Karoo Province,

but rather are more closely analogous to the more

localised central complexes seen in intraplate alkaline

volcanic complexes (Johnson 1989).

In summary, it is proposed that the lava at Red Creek

represents a single flow of hawatite composition from

a central volcanic complex, closely analogous to that

of the Truro Volcanics but not necessarily from ihe

DIFFERENTIATION INDEX

100

felsic focks

80.

benmoreite

60

intermediate

matic

(.¢. alkali, basalt, basanite,

hawaiila, nephelinite)

TOCKS

40

mugearite

hawaiite

0 20 40 60 80

NORMATIVE 100 x (an/(an + ab))

100

a. .,Red Creek mafic lava

Fig, 8, Red Creek lavas on CIPW normative classification

plot for sub-alkaline intraplate lavas (otinson 1989).

EARLY CAMBRIAN VOLCANICS 65

same centre or erupted at exactly the same time,

However, both the laya at Red Creek and the Truro

Volcanics undoubtedly belong to the same Early

Cambrian alkaline ‘within plate’ volcanic province,

which may be genetically linked to other Cambrian-

mafic alkaline provinces such as the Yumali/Coonalpyn

and Peebinga-1 areas (Rankin et a/. 199la’ and b’).

The closest tectonic analogué for these provinces is

that of a rifted continental margin-as proposed for the

Ti / 100

zr Yu

a Red Creek matic volcanics

Fig. 9, Red Creek volcanics on tectonic discrimination plot

of Pearce & Cann (1973). “Within plate” basalts plot in field

D, Morb (ocean floor basalts) in field B, low Ktholeiites

in field A and B, calc-alkaline basalts in fields C and B,

NORMALISED VALUES.

0.1 t

Sr kK Rb Ba Th Nb Ce P Zr Ti

ELEMENTS

¥ Sc Ni Cr

Red Graek volcanics

Mt Rufus 1 lavas

——— MBT-1 lavas

Yumali / Coonalpyn lavas

MBY-12 matadolarie

Fig. 10. MORB normalised spidergram for averages of possible

Cambrian mafic suite from the Murray Basin basement and.

the Cambrian lavas of the Mt Lofty Ranges.

Tertiary intraplate volcanic province of eastern

Australia (Johnson 1989); metadolerites which post-

date the volcanics indicate 4 change with time from

intraplate to MORB type composition (Liu & Fleming

1990) indicating an ensuing period of crustal thinning

and major dyke emplacement, associated rifting and

crustal extension.

Acknowledgments

The authors wish to thank Dr David Gravestock

(S.A. Dept of Mines & Energy) for reviewing this

paper and making some yaluable suggestions.

Logistical support was provided both by the §.A. Dept

of Mines & Energy and the University of South

Australia, This work was supported by a University

of South Australia Research Grant to JBJ, This paper

is published with the permission of the Director

General of the S.A, Dept of Mines & Energy,

References

Coats, R. P, & THomson, B. P. (1959) TRURO map sheet,

Geological Atlas of South Australia, 1:63 260 Series. (Geol,

Surv. S. Aust., Adelaide),

Cops, M. A. & Farranb, M. G. (i984) A new occurrence

of the Truro Volcanics. Quart. Geal. Notes Geol, Surv.

S. Aust. 89, 8-10,

Cooper, J. A., Jenkins, R. J, FE Comeston, W. &

Witutams, I. 8. (1992) lon-probe zircon dating of a mid-

Early Cambrian wf in South Australia. Geol, Soc. Lond.

Journal 149, 185-192.

Crawrorp, A. J, & Keays, R. R. (1978) Cambrian

greenstone belts in Victoria; marginal sea-crust slices in

the Lachlan Fold Belt of southeastern Australia, Earth &

Planet, Sc. Lett. 41, 197-208.

100

NORMALISED VALUES

Red Creek yolcanics

> ‘Mt Rufug 1 lavas

MBT-1 lavas

Yumali / Coonaipyn lavas

MBT-12 metadalerita

Fig. ll. Karoo Basalt normalised spidergram for averages of

possible Cambrian mafic suites from the Murray Basin

basement and the Cambrian lavas of the Mt Lofty Ranges.

66 ©. G, GATEHOUSE, J. B. JAGO, B. J. CLOUGH & A. J. McCULLOCH

Dany, B. (1963) The Fossiliferous Cambrian succession in

Fleurieu Peninsula, South Australia, Rec, 5. Aust. Mus,

14, 579-601,

& Mivnes, A. R. (1972) Revision of the stratigraphic

nomenclature of the Cambrian Kanmantoo Group, South

Australia. J; Geol. Sac. Aust. 19, 197-202.

Forses, B. G., Coats, R. P. & Datiy, B. (1972) Truro

Volcanics, Quart. Geol. Notes, Geol. Surv. S, Aust. 44, 1-5.

GateHouse, C. G, (1988) TEPKO map sheet. Geological

Aulas of South Australia , 1:50,000 Series, shect 6728-II!.

(Geol. Surv, §. Aust., Adelaide),

GateHouse, C. G., Jaco, J. B. & Cooper, B. J. (1990)

Sedimentology and stratigraphy of the Carrickalinga Head

Formation (low stand fan to high stand systems tract),

Kanmantoo Group, South Australia, J J. B. Jago & P. J.

Moore, (Eds) “The evolution of a Late Precambrian-Early

Palaeozoic rift complex: The Adelaide Geosyncline” Geol,

Soc, Aust. Spec. Pub,, 16, 351-368.

Gravestocx , D, I. (in press.). Early and Middle Palaeozoic.

In “Geology of South Australia.” (South Aust. Dept. Mines

& Energy, Adclaide)

Jounson, R. W. (Ed) (1989) “Intraplate volcanism in eastern

Australia and New Zealand”. (Cambridge University Press,

Cambridge).

Le Bas, M. J., Le MAITRE, R. W., STRECKEISEN, A, &

ZANETTIN, B, (1986) A chemical classification of voleanic

rocks based on the total alkali-sili¢a diagram. J. Petrol,

27, 745-750.

Lru, S. F & Fremia, P. D. (1990) Mafic dykes and their

tectonic setting in the southern Adelaide Foldbelt, South

Australia pp.401-413. Jn A. J, Parker, P. C. Rickwood &

D. H. Tucker, (Eds). “Mafie Dykes and Emplacement

Mechanisms”. (Balkema, Rotterdam).

Parker, A. J. (1986) Tectonic development and metallogeny

of the Kanmanton Trough in South Australia. Ore Geol.

Rev. I, 203-212.

PEARCE, J. A. & CANN, J. R, (1973) Tectonic setting of basic

volcanic rocks determined using trace element analyses.

Earth & Planet. Sc. Lett. 19, 290-300.

PoweLL, C. Mc. A. (1990) Gondwanaland context of the

Tasman Fold Belt. Geol. See. Aust. Abstracts, 25, 190-191.

SCHEIBNER, E. (1986) Suspect terranes in the Tasman Fold

Belt System, Easter Australia. Jn D. G. Howell (Ed.)

*“Tectonostratigraphic terranes of the Circum-Pacific

Region", Circum Pacific Council for Energy and Mineral

Resources, Earth Science Series. 1, 493-514.

TuRNER, S. P. & Fopen, J. D. (1990) The nature of mafic

Magmatism through the development of the Adelaide

Geosyncline and the subsequent Delamerian Orogeny,

South Australia pp. 431-435 In A. J. Parker, P. C. Rickwood

& D. H. Tucker (Eds) “Mafic Dykes and Emplacement

Mechanisms”. (Balkema, Rotterdam).

Von DER Borcu, C. C. (1980) Evolution of Late Proterozoic

to Early Palaeozoic Adelaide Foldbelt, Australia.

Comparison with post-Permian rifts and passive margins.

Tectonophysics 70, 115-134.

WINCHESTER, J. A. & Fuoyb, P. A. (1977) Geochemical

discrimination of different magma series and their

differentiation products using immobile elements. Chem.

Geal, 20, 325-343,

BIOLOGY OF ETIELLA BEHRII ZELLER (LEPIDOPTERA: PYRALIDAE):

A PEST OF SEED LUCERNE IN SOUTH AUSTRALIA

BY A. D. AUSTIN, T. C. R. WHITE, D. A. MAELZER & D. G. TAYLOR*

Summary

The general biology of the pyralid moth, Eviella behrii Zeller, a serious but sporadic pest of lucerne

seed crops in South Australia, is documented. Larvae are recorded as feeding on six native and 18

introduced legume species. Head capsule widths of field-collected larvae show that E. behrii, like

other pyralids, has five instars. Light-trap catches indicate the presence of at least four overlapping

generations per year in the Adelaide area, with the majority of moths occurring during three

generations from August to February. In lucerne crops in the Keith area, 150 km south-east of

Adelaide, young and mature green pods suitable for oviposition are available from October to

February. Two peaks in moth catches were recorded in late December and late January,

respectively. Until mid-January, the number of eggs was strongly correlated with the numbers of

moths caught, but thereafter the numbers of eggs and larvae decreased, with the second peak of

moths giving rise to very few larvae. These observations are in agreement with the opinion of

farmers that early lucerne crops are more heavily infested with E. behrii than late crops. Surveys of

vegetation in the Keith area show that native host plants are very scarce, and that volunteer lucerne

plants along road-sides and irrigation banks are the likely source of early infestations in Lucerne

crops. Mortality agents recorded include ten species of parasitoids, two pathogens and a number of

general heteropteran and spider predators. These and other possibilities are discussed as factors

leading to the observed decrease in larval numbers in mid to late January, a time when lucerne crops

still contain a large proportion of green pods suitable for larval development.

KEY WORDS: Etiella behrii, Pyralidae, ecology, life history, lucerne, alfalfa, host plants,

parasitoids, predators, pathogens.

Transactions ef the Reval Society af S. sust. (1993), 117(2), 67-76.

BIOLOGY OF ETIELLA BEHRII ZELLER (LEPIDOPTERA: PYRALIDAE)<

A PEST OF SEED LUCERNE IN SOUTH AUSTRALIA

by A, BD, Austin, T. C. R. WHiTE, D. A. MABLZER & D. G. TayvLor*

Summary

Ausiin, A. D.. Warre. 'T. C. R., Magvzer, D, A. & Taytor, D, G. (1999) Biology of Fella behrtt Zeller

(Lepidopten; Pyralidae): a pest of seed lucerne in South Australia. Trans. R. Soe. 8. Aust. U7(2) 67-76, 4 June, 1993,

The general biology of the pyralid moth, Eriella behrii Zeller, a serious but sporadic pest of lucerne seed crops

in South Australia. is documented. Larvae ure recorded as feeding on six native and 18 introduced Jeguine species

Head capsule widths of field-collected larvae show thal £. behrii, like other pyralids, has five instars. Ligh(-trap

caiches indicale {he presence of at feust four overlapping generations per year in the Adelaide ares, will! the ninjority

of yroths Occurring during three generations from August to February. In lucerne crups in the Keith area, 150

km south-east of Adelaide, young and mature green pods suitable for oviposition are available from October to

February, Two peaks in moth catches were recorded in late December and late January, respectively. Until mnid-

January, the number of eggs was strongly cocrelated with the numbers of moths caught, but thereafter the uuinbers

of eggs and larvae decreased, with the second peak of moths giving rise to very few larvae. These observations

an: in agreement with the opinion of farmers that early lucerne crops are more heuvily infested with £. bebtrit

than late crops, Surveys of vegetation in the Keith area show that native host plants are very scarce, und than

volunteer lucerne plants along road-sides and irrigalion banks are the likely source of curly infestations if Wweerne

crops. Mortality agents recorded include ten species of parasitoids, two pathogens. and a number of general

heteropteran and spider predators, These and other possibilities are discussed as factors leading to the observed

deerease in larval nufubers in mid to lute January, a time when Jucerne crops still contain a large proportion

of green pods suitable for larval development,

Kev Worns. “ella behrii, Pyralidac, ccology, life history, lucerne, alfulfa, host plants, parasitoids, predators,

pathogens.

Introduction

Ole of the most serious pesis which ean limit the

production of lucerne seed in South Australia is Eriella

behrit Zeller (lucerne sced web moth) (Maclzer ef al.

1982u, 1982b; Durhanr 1984; Austin ef al, 1986). It

is One of seven species of Friella, the larvae of which

feed on the seeds of legumes (Whalley 1973), The most

witlespread and economically important species is the

cosmopolitan &. zinckeneller (Treitschke), a serious pest

of Jepunies in many parts of the world, which in

Australia is known only from Queeasland (Abdul-Nasr

& Awadalla 1957; Stone 1965; Singh & Dhooria 1971;

Whalley 1973; Common 1990), The remaining six

species are restricted to Australia, parts of south-east

Asta and some Pacific islands, Only three species, E.

chrysoporelia Meyrick, EB. hobsont (Butler) and E.

behril, are recorded from South Australia; the former

twoonly curely as adults, and only from the northern

and western parts of the State, The multivoltine £.

behrii is a widespread and common pest throughout

Australia, and of economic significance because of its

heavy but sporadic outhreaks in dryland and irrigated

crops of seed lucerne.

* Departmen of Crop Protection, Waite Ayricultural

Reseurch Institute, University of Adelaide, Glen Osmond,

5. Aust. 5064,

Little is known of the general biology or ecology

of £. hehrii, possibly because of its sporadic

occurrence. In most years it does little damage to

lucerne seed crops in S.4., but occasionally it causes

widespread damage. Atsuch times the species is also

reported fo damage lupins, field peas and some clovers,

e.g. in 1954 and 1971 (Austin eral, 1986; DAM and

DGT, unpubl.). £. behrii is thought to live on native

plants along road-sides and in patches of scrubland

during spring and eatly summer before infesting

lucerne seed crops in mid summer, but this has not

been confirmed. The present study was therefore

undertaken to elucidate this pari of its phenology and

10 document its genetal biology. Data have been

included from a pest management program on Jucerne

seed crops (see Maelzer et al. 1982a, 1982h, Durham

1984), information collected in 1959-60 (by DGT), and

life history studies undertaken during 1980-82,

Materials and Methods

Location of study

The project was centred on a property (‘Brecon’) 10

km south of Keith, S.A., inthe middle of a mayor area

of ducerne seed production The area bas 4

Mediterranean-type climate with mostly cold, wet

winters and hot, dry summers (Anon, 1987). The

68 A. D. AUSTIN, T. C. R. WHITE, D. A. MAELZER & D, G. TAYLOR

average rainfall for the area is 471 mm per annum. The

early (spring) dryland lucerne crops are watered

naturally by rainfall, but the majority of the seed crops

are managed to flower in mid-summer and are

irrigated, mostly by flood irrigation rather than by

sprinklers. Work in the Adelaide area, which has a

similar climate to Keith, was carried out in 1959-60

and was centred at the Waite Institute campus of the

University of Adelaide.

Trap catches of adult moths

Adults were caught at a light trap each night in two

separate studies in widely different years: over the

period 1.111.1959 to 18.11.1960 on the Waite Institute

campus (by DGT) and from 1.xit.1981 to 20.11.1982 at

‘Brecon’ (by DAM). The second period of trapping was

part of 4 general program of sampling insects in lucerne

seed crops (Maelzer er al, 1982a, 1982b). The light

trap in 1959-60 was a mercury-vapour type, while that

used in 1981-82 was a 15 watt blacklight type (cf.

Hardstack er al. 1979), located in the middle of a

lucerne area containing both dryland (early) and

irrigated (late) crops,

Sampling eggs and larvae from lucerne crops

Five sampling sites were selected at ‘Brecon’ around

a field of lucerne which was to be irrigated later that

summer. All were dryland areas (i.e. not irrigated) and

were of the commonly grown Australian cultivar,

Hunter River. Each had been closed off to grazing

animals some weeks earlier and contained lucerne

plants with flowers and pods. Samples were collected

from these sites about once per week from the

beginning of December 1981 until mid January 1982

and taken back to the laboratory. Ten small samples

(each of about 50 pods) were also taken from one or

two more distant, dryland sites (up to 30 km away).

including fields of lupins with volunteer lucerne plants,

Each sample was taken by walking through an area

of 4-5 ha and picking stems of lucerne with flowers

and pods from a number of widely spread points until

15-20 stems had been gathered. The stems were

temporarily stored in a plastic bag on ice. In the

laboratory each stem was searched, ftom the oldest

raceme upwards, and each pod on each raceme was

examined for eggs and larvae on the outside, and larvae

inside. The procedure was continued until

approximately 200 pods had been inspected. This

meant searching from 1-12 stems (x = 5.0; $.D. = 1.9)

bearing from 1-1] racemes each (X = 5.3; §.D, = 2,3).

The racemes on any one stem were at any stage from

unopened flowers to fully matured brown (dry) pods.

All stages of flowers and pods were usually present

on a single stem and, wherever possible, stems were

selected that had some well-developed pods. For each

raceme a record was made of the number of E. behrii

eggs and larvae (classed as instars: 1 + 2,3 + 4,

or 5) and the stage of development of the pods (classed

as flowers, pods just forming, juvenile pods, young

pods, mature green pods and mature brown pods —

see Fig. 1). In earlier samples, no stages of E, behrii

had been found on flowers, on pods that were just

Fig. 1. Stages of pod development on lucerne plants: A, pods just forming; B, juvenile pods; C, young pods; D, mature

green pods; E, mature brown (dry) pods,

BIOLOGY OF ETIELLA BEHRII ZELLER (LEPIDOPTERA PYRALIDAE) 69

Fig, 2. Female Efiella behrii ovipositing onto a raceme of mature green lucerne pods.

forming, on juvenile pods or on mature brown pods.

Therefore, in later samples racemes bearing only these

stages were excluded from samples, and only racemes

bearing some young and mature green pods were

examined.

Later in the crop season, when the dryland crop had

nearly all brown pods, similar samples were taken from

an adjacent crop of irrigated lucerne, of the aphid

resistant cultivar WL318, but counts were made of

larvae only. Because of possible differences in numbers

of larvae in young and in mature green pods, the

numbers were expressed as stratified means per 1000

pods.

Rearing techniques and laboratory culture

Techniques for rearing field-collected E. behrii

larvae through to the adult stage were developed so

that a constant supply of each stage was at hand for

life-history studies. Small larvae (Ist and 2nd instars)

collected from lucerne pods were successfully reared

on a medium based on either powdered peas or lucerne

seed (cf. Shorey & Hale 1965), but the agar had to

be boiled for twice the suggested time for the medium

to set. Each larva was placed in a 2.5 cm’ vial

containing 1.5 cm? of medium. Vials were sealed with

hard plastic lids. The pupae were removed from the

vials and placed in a dry container until the adults

emerged. Both reared and field-collected adults were

kept in gauze cages and given access to 10% sucrose

solution. All stages were maintained in a dimly lit room

held at 24-28°C and 75% R.H. in which the generation

time was 25-35 days,

Cooper (1979)! obtained large numbers of E. behrii

eggs from females ovipositing into the crevices of a

darkened wooden box. We had no success with this

method, nor with a variety of other surfaces including

perspex, cardboard, freshly crushed lucerne, and

rearing medium with lucerne seeds embedded in it.

However, we obtained eggs laid onto freshly shelled

peas or fresh mature green lucerne pods. Peas proved

most successful because eggs could be easily and safely

removed from their smooth surface. Eggs were more

difficult to remove from lucerne pods because of their

more complex surface. They were collected with a fine

paint brush and transferred to a dry glass vial, and the

Ist instar larvae were then transferred to a second vial

containing rearing medium. With this method we

initially obtained 80 eggs and reared 90% through to

the adult stage.

Surveys of natural vegetation

Extensive surveys for E. behrii larvae were carried

out in the native vegetation surrounding the irrigated

lucerne growing area south of Keith (for approximately

8 km in each direction) and along road-sides between

Tintinara and Willalooka (35 km north-west of Keith)

in October-November, 1981. Extensive surveys were

also conducted by one of us (DGT) in 1959-60 around

Keith and Tintinara, and this information was extracted

from the unpublished records of the (then) Department

of Entomology, Waite Agricultural Research Institute.

'Cooper, D.J. (1979) “The Pathogens of Heliothis punctigera

Wallengren”. Ph.D, thesis, University of Adelaide, Unpubl.

Ww AOU AUSTIN, TOC. RWI DA MAELZER & DUG. TAYLOR

Paresiteidy and pathogens ef F. behvii

To trap udults of known parasitonds of £. behrii, ten

yellow pan traps fled with soupy water were placed

armund the edges of the inigaled WLAI8 lucerne crop

and ere CXanuned once per week, The traps were rut

Inyin early December 1981 to mid-February 1982. In

addition, 41 late-staze A. behrii larvae (4th and Sth

instars) collected from both the drylind ung irrigated

lucerne crops were reared on artificial deet in the

laboratory to check for possible parasitoids, Similarly.

one of Us (DCT) collected large numbers af &. beheti

faryae From both native and crop plants in S0-Mb io

obtain parasitoids. Also, other workers have bred

parasitoids from £ behrii collected in the Keith area

over the last 40 years. Voucher material of parasitoids

collected in the above manner has been lodged in the

insect collection at the Waite Institute, and has been

used lo collate the duty presented here.

Pathogens of 16 field-collected 4th and Sth instar

larvae that died soon after in the laboratory wore

screened for pathogens by DJ. Cooper at Waite Institute

using the methods of Cooper (1979)!

Results and Observations

Life history stages

At all stages of its life cycle & brheii parallels

descnptions of bolt the behaviour aud the appearance

ole, zinckenella, apart from being slightly smaller in

size (Abul-Nasr & Awadulla 1957; Stone 1965; Singh

& Dhoorit 1971). The eggs are ovoid, averaging 0.6

mim it length (8.0, = 0.03; n = 50) and 0.4 mm in

widih ($.D, = 0,3), colourless when first laid, but Soon

turn pale yellow or orange. A day before hatching the

hiuck heat capsules and prothoracic shields of the

Jatvac are clearly visible hrough the chorion, Eggs

are laid on either young or mature green pods, and

are usually placed between the pod and the calyx or

between the coils of the pod.

The frequency distribution of head capsule widths

of field-collected larvae sugested tive peaks and hence

Nive wstars (Fig. 3). but there was much vatisatien

between peaks, The minge of cach instsr was beter

differentiated hy the minima of a three-point mowing

average peocess (Pig. 3), The mean widths for five

inshirs were then calculated as 0.1%, 0.28, 1.44, 01,70,

om 0.99 mim wall multipliers between means, for

Dyar’s Jaw, of 1,54. 7,59, 1.60 and 141. Five instars

were also confirmed from the rearing of £ behrii in

ihe laboratory. The Ist instar has a pale yellow body

and a very distinetive black head capsule and

prothoracic shicld. Soon atier batching it constructs

a fine fuinel-like silk tube around itself with one end

ullached to the sortie of the pod, i then chews through

the pod and into seed where it remains feeding, The

entrance hole in the pod quickly heals over. the web

tube soon disintegrawes, and no evidence remuing to

indicate that the pod as pnlestes!, The larva docs not

qonnally leave this firs seed until most of ity contents

are devoured. It then atacks the next seed in the pod,

although it is now too lange to get iaside it. By the late

ard to carly 4th instar, We larva chews its way out of

the origmal pod and enters one ar more new pods,

eating some (and occasionally all) of the seeds. within

Commonly, at this stage most seeds are only partly

devoured, ‘fhe larva tow has a bluck head capsule,

semi-circular prothoracic shield and spiracles, The pro-

and mesothorax are golden-brown, the metathorax and

abdomen pale green There are seven dorsal and later

longitudinal pale: golden-hrown stripes on the body.

By the late dil instar the larva attacks several pods,

meshing these and its frass together with silk to form

a protective lent. This webbing is the first readily

observed outward sign that &. behrit has attacked o

erop. By the 3th instar the larva has a pale golden-

brown head with dark brown mandibles, The

prothoracie shield 1s still black bul the rest of the body

ix green wilh a pinkish tinge to the abdomen. The

longitudinal bands are more diffuse and pinkish, This

Howl instar becomes a steadily darker pynk, the green

heing musked completely just prior (o pupation

Pupation takes place a dew centimetres benwath the

Surfitce of Ihe soil, inside a cocoun of silk, which

incorporates particles ol soil.

Adult moths are small (10-15 mm long), greyish and

distinctive becatise of thew typical ‘inoul’ (formed by

the forwurd-directed pulps), the white stripe along the

full length of the leading edge of exch lore wing, und

the transverse orange band near the buse of each fore

wing (Fig. 2), At rest, moths have av ‘alert’ appearance

with the head aod thorux raised and the abdometi and

distal ends of the wings touching the surface. ‘The

antennae of the female are simpoth and undifferentiated

While those of the male have an enlarged basal segment

and i “bush! of scales on the clonguted second segment

The maxillary palps of ihe female diverge at their

second yom fo form an-open ‘V" at the ip-of the wneat,

These palps are parallel in the male moth,

Hast plants and sources. of infesterien

Table | lists all the plant species tor whieh & beled

larvae have been recorded feeding on thelf seed [fi S.A,

Fram 1959 to 1982, Larvae have also been rmeeorded

on many of the listed exotic species in Other sbtles, as

wellas on-soybeun (Glycine mvay (1L.) Merrill), peauues

(Arachis hypegaea L.) and Acacia evelups AL Cun,

ex G, Dan in Western Australia, Narihern Territory

(Comunoa 1990) and Queenstind (Tumer 1980),

all these hosts, lucerne is the dost commonly and

severely attacked, but the species alse suiees sporauie

damage lu lupins, Geld peas and some Govers (Austin

eta, 1986; DAM & DGT, tunpulst

BIOLOGY OF ETIELLA BEHRI ZELLER (LEPIDOPTERA PYRALIDAE) 7

8

Frequency

0.4

0.6

Head width (mm)

‘

|

tHE } ia

.O

Fig. 3. Head capsule widths of larvae of Etella behrii collected randomly from the field at ‘Brecon’ (Keith) (n= 329), expressed

as a frequency distribution (histogram) and three-point moving averages (MB).

The survey of native vegetation in late 1981 revealed

that there is. very little native scrub left in the Keith

area, and what patches do remain are small,

considerably disturbed and have very few native

legumes, both in numbers of species and individual

plants. Far more commonly encountered are volunteer

plants of various exotic weed, crop and pasture legumes

(especially lucerne) along road-sides, in clearings in

scrubland, and in fields carrying other crops such as

Jupins. Lupins are a potential source of recent

significance which have been planted more frequently

in the Keith area since 1980. Lupins are planted from

early winter to early spring, the seed ripening and

drying out up until late November, thus providing an

ideal carly source of food for larvae, the adults of which

emerge early in spring. Samples of 200 pods of green

mature lupin pods taken from crops south of Keith and

near Tintinara in early November, 1981 showed a small

(<1%) infestation of £. behrii larvae.

Seasonal phenology

Observations in the laboratory indicate that females

can lay more than 200 eggs and that eggs hatch within

24 h at 30-35°C, but usually hatch after 4-7 days in

the field in spring-summer, Larval duration in the field

during spring-summer varied from 10-28 days and the

pupal period from 14-21 days. Adults survived 7-21

weeks in the laboratory when held at ambient

temperature and supplied with 10% sucrose solution.

The light-trap catches of adults at the Waite Institute

campus in 14 day intervals for the period 1.iii.1959 to

18.11.1960 are given in Fig. 4. These catches suggest

72 A. D. AUSTIN, T. C. R. WHITE, D, A, MAELZER & D. G, TAYLOR

TABLE 1. Known host plants of Etiella bebrii in South Australia (records collected 1959-82).

Native Legumes

Acacia victoriae Benth (elegant wattle)

Daviesia brevifolia Lindley (leafless bitter-pea)

Eutaxia microphylla (R. Br.) Black (Eutaxia, mallee

bush-pea)

Hardenbergia violacea (Schnee.) Stearn (native lilac)

Pultenaea tenuifolia R. Br. ex Sims

Pultenaea densifolia F. Muell. (dense-leayed bush-pea)

Introduced Legumes

Cytisus scoparius (L.) Link (English broom)

Cytisus proliferus L.f. (tree lucerne)

Genista monspessulana {L,) L. Johnson (Montpellict

broom)

Lupinus cosentinii Guss. (Western Australian blue lupin)

Lupinus angustifolius L. (lupin)

Lupinus uliginosus Schkyhr (birdfoot trefoil, greater loms)

Medicago ciliaris (L.) Krocker

Medicago littoralis Rohde ex. Lois (strand medic)

Medicago polymorpha L. var, polymorpha (burt medic)

Medicago rugosa Desr. (gamma medic)

Medicago sativa L. (lucerne, alfalfa)

Medicago truncatula Gaertner (barrel medic, snai) medic,

caltrop medic)

Pisum sativum U.. (field peas)

Trifolium frapiferum L. (strawberry clover)

Trifolium resupinatum.L, (shaftal clover)

Trifolium subterraneum L. (subterranean clover)

Vicia dasycarpa Ten. (woolly-pod. vetch)

Vicia villosa Roth. (hairy yetch, winter vetch)

at least four overlapping generations each year, with

peak catches in late September (spring), late November

and late December. There is then a generation with

a peak in mid to late February and a hint of other small

generations in March-May.-OQur trap catches for adults

in December 1981 to January 1982 for the Keith area

are in accordance with these earlier data by clearly

detecting two large summer peaks (Fig, 6), even though

they occurred at a slightly later time than the 1959-60

800

600

Ww

x

2

wn

Oo 400

#2

Lu

a

=

2

Zz

200

ty)

peaks. When related to monthly mean temperatures

(Anon. 1987) by iterative calculations, these

observations suggest 4 mean generation time of about

210 day degrees above about 13°C. ,

South Australia has a hot dry summer (Anon, 1987)

and as summer progresses, dryland lucerne should

gradually becomes unfavourable for E. behrii. Our

survey in dryland lucerne in the early sumimer of

1981-82 was planned to quantify this change; and in

A 5 °o N D d F

MONTH OF YEAR

Fig. 4. Light-trap catches of adult Hriella behrii at the Waite Institute campus, Adelaide in 1959-60 accumulated in intervals

of 14 days.

BIOLOGY OF ETTELLA BEHRI] ZELLER (LEPIDOPTERA PYRALIDAE)

Table 2 are given, for each sample (a) the numbers

of juvenile, young and mature green pods , and (b)

the total oumber of eggs and larvae of E. Sehrii in

young and mature green pods (none were found in

juvenile pods)

The total numbers of E. behrii were obviously

greater in mature green pods than in young ones from

61.82 onwards (Table 2}, but the numbers of different

Stages were loo small to allow comparisons between

the (wo sorts of pods, except for eggs and Ist+2nd

instars on 30,xii.81 and 6.1.82. For egys on 30.xii.81

plus 6.1.82, there was a significantly greater number

on young pods (51 on 1250 pods) than on mature green

pods (31 on 1211 pods) (x? = 3.95, p <0.05). For

eggs plus Ist and 2nd imstars, 78 on 1250 young pods

were also significantly more frequent than 47 on 121!

Mature green pods OC = 7.69; p <(0E). The

comparison of these two sets. of data suggest that in

the samples for these two days some young larvae were

faillng to establish in the mature green pods. Obviously

the larger number of all stages of EB. behrii on mature

green pods after 6.1.82 was therefore due to larger

numbers of older larvae, Because thé two types of pods

could be considered as different sampling strata, the

total numbers of £. behrii in each sample were

Subsequently expressed as means per 1000 pods,

corrected for the different. numbers of young and

mature pods, Such means of all stages of E. behrii wre

also given in Table 2, and separate means for eggs and

for instars 1+2, 344 and 5 ate given in Fig, 5.

In Table 2, the decline with time of the numbers of

both juvenile and young pods illustrates the maturation

of the crop, and the data in Table 2 and Fig. 5 indicate

@ concomitant decrease in number's of eggs and Ist +

2nd instars of &. bekrii and an increase in numbers

of older larvae. To try to make more: sense: of these

trends, the. numbers of eggs and of all larvae were

plotted against the numbers of adults of FE. beknii caught

per week at the light trap at Keith (Fig. 6). ‘The dat

Sugges! that the numbers of eggs laid up to and after

the first peak of moths were strongly correlated with

the numbers of moths each week. However, the

13

40

1 cays

B inaiars 1

ST & owsarsse

El invetar &

Nurpbers per 1000 pede

Dec, dan:

Fig, 5. Corrected numbers per 1000 pods of Eriella behrii

eggs, Ist + 2nd instar, 3rd + 4th instar, and Sth instar larvae

in a dryland lucerne crop at ‘Brecon’ (Keith) in relation

to calendar time from 1. xii.1981,

numbers of eggs on 15.1.82 and 19,1,82 were much

lower than expected in relation to the number of moths,

which had increased to a second peak (Fiz, 6).

Unfortunately, no counts of eggs were made after

19,i,82 because attention was then diverted to the main

irrigated sced crop which was in full flower.

The cause of the decline in numbers of eggs ii the

dryland crop on 15.:.82 and 19,1.82 (Figs 5, 6) is not

known, There scemed to be an abundance of suitable

pods for oviposition on these two dates (Table 2), but

it ts possible that the crop had dried out further so that

even the young pods were subtly different from those

on earlier dates and were- unsuitable for oviposition.

So too there was an unexpected decrease in the

numbers of young larvae on 19,1,82 (Fig, 5), possibly

again because the pods were not as suitable for their

establishment as they had been earlier.

The numbers of larvae in irrigated luceme seed crops

were of particular interest to us from the view point

TaBLe 2. The numbers of juvenile (J) , young (¥) and matwre green (M) pods in each sample; and chi: numbers of ate sages

af BE. behrli (Ab) in each category of pads.

Numbers of pods Numbers of Z.b, in Mean & >, per

Date of sample J Y¥ M Y feds M pods 1000 Y+M pods

09,12,81 276 a4 91 its) 4 156

16.12.81 362 637 207 7 6 Ls.4

22.12.81 267 573 330 6 3 10,0

30.12.81 113 650 S42 30 3 52.9

06.01.82 95 600 669 21 3? 615

15.01.82 Vv {87 463 2 24 40.0)

19.0.2 15 238 1165 i 44 32.1

Tutal 1145 3639 3467 7? 171

id 4. D AUSTIN, T CR, WHITE, D. A. MAELZER & DUG TAYLOR

ia

~

3

' " a) a} 19 aii so fo wo

oe, aan, Fen.

Mig. & Number of eegs | A) ond larvae (MM) of Brielle betray

per 1000 pods in samples taken at ‘Brecon’ (Keith) in a

dryland lucerne crop inthe sunwoer of 1981-82; the number

‘of larvae ([2) similarly sampled in an adjacent irrigated

lucerne-crop; and the number of moths (O) caught weekly

ina light-trap in the same itrigated crop, in relation to

calendar time from |.xii.1981.

ol pest management, and the irrigated and unsprayed

WLAl8 crop, which we sampled later in the season,

had keen managed so that it flowered later than the

adjacem dryland crap; and on 19,1.82 it was at about

the same stage of development.as the dryland crop had

been an 9, xi7,.82, The numbers of Jarvae in the irrigated

crop are also giver in Fig. 6, On 15.1.82, they were

about the same as those w the dryland crop on 15,1.82

and 19.i.82, but they similarly declined thereafter (Fig.

6) despite un abundance of seemingly suitable young

and mature green pods for E. behri/. In particular, there

wus no increase if the numbers of larvae afler the

second peak of trapped moths on 19,1,82.

Untirtunately, eggs were not recorded in these samples

from the irrigated tucermne, ner were the larvae

categorised into instars, It is not known, therefore.

where the mortality occurred that caused the decreasing

trend in Jarval numbers, The weather was not obviously

unfavourable for moth survival and oviposition, so the

mortality is likely to have been al the egg and/or larval

stage, Two inain possibilities may be suggested for this

decline. Firstly, it is possible that there was a very Jow

establishment of young Larvae on the aphid resistant

cultivar WL318, as Perrin (1978), for example, found

in some varieties of cowpea attacked by Cydia ptychorr

(Meyrick). The second possibility is that the mortaliry

was duc to predators and pathogens. In particular, the

decrease in larval numbers ocurred at the same time

as a dramatic Increase tn the numbers of Campylonmmur

lida Remer (dimpling bug), which is thought to be

i, isn

f ‘

ea ng

; =

partly predacious, as is C. verbase? (Meyer), a well-

known predatod of mites, aphids and psyillids in Canada

(McMullen & Jong 1970).

Finally, the second peak of moths in mid-January

(Fig, 6) may be expected to be smaller than the first

one because the area of dryland lucerne is muuch greater

than thas of immgaled lucerm: and, as summer

progresses, dryland lucerne is oo longer favourable for

£. bekrii, So one might therefore expect fewer maths

to be caught in a light-trap in February-March, as is

partly illustrated in Fig. 6 for Keith. Other crops

attacked by £. beh, such as lupins, would similarly

cease to produce suitable green pods for young

caterpillars in mid January unless they were irrigated.

Therefore, in most localities there would be a reduction

in numbers of larvae in late January to eurly February

and a subsequent reduction im the numbers of moths

at the next peak catch of maths, as shown in Fig. 4

for Adelaide in mid to late February, The few moths

flying in late February would find even fewer suitable

pods for wriposition at the end of the hot dry summer.

So the species ts likely to be relatively rare in most

localities after February and until the next flush of plant

growth an spring, following Winter rains. During this

imerval the species could either enter an obligate

diapause or it could continue to breed in reduced

numbers (on volunteer lucerne}, The light-trap data

for Adelaide (Fig. 4) strongly support the latter

alternative, because a few moths were caught in late

April w early May. The lack of a diapause was also

dentonstrated directly by ane of us (DGT) by

successfully incubating to adult emergence, larvae

collected! over the winter of 1959,

Parasiteids, predators and pathogens

The parasitoid species which bave been recorded

trom &. beh are given in Table 3 along wath the silage

from which they emerge, whether they are solitary or

gregarious, and their status ay either primary or

hyperparasitoids, The ichneumonid Femefucha oprnea

Kerrick and the braconid Jconella alfalfae (Nixon) have

been recorded as causing between 2% and 1%

mortality 10 field populations in the Keith area in

1939-60 (DGT unpubl.). while the other species listod

have heen only rarely encountered. Yellow pan-traps

placed in the field at ‘Brecon’ in carly December 1982

first caught 7. cychea in early February. The numbers

of this parasitoid appeared to mcrease through February

aod March, but there were too few collected per trap

to quantify diflerences between samples. Phanerotome

sp, was also caught in late February but in very low

numbers,

Of 41 larvae of &. behirii taken from the field in late

January to screen for parnsucids, 25 pupated, and three

were parasitised (lwo by T. cycrea and one by

Penerofoma 3p_j. T. cycred oviposits in) early stage

larvae after the latter emerge from the pod and begin

BIOLOGY OF ETIELLA BEHRIT ZELLER (LEPIDOPTERA) PYRALIDAE) 1S

to build a webbed retreat, while Faneroiama sp.

oViposits into the eggs of its host, Both species then

complete their development after the final host instar

has dropped 1o the soil and pupated. During the 1981-82

study we did not record any other parasitoid species,

even though J. alfalfae has been recovered in moderate

numbers in the past (DGT unpubl,).

We did not directly observe any predation upon eggs

und larvae of E. behrii, but several general predators

wete. seen in lucerne crops, and they may feed on &,

behrii. The predators included C. livida (see abave)

and spiders, particularly of the families Oxyopidae,

Araneidae, Lycosidue and Thomisidae. Of these

species C. livida was by far the most common in

irrigated crops during January-February 1982.

Sixteen of the 41 screened larvae died before they

pupated. After death they became soft, darker in colour

and developed faint spots undet the integument. These

larvae were later examined and found to contain large

numbers of microsporidian spores and some bacteria

(Bacillus sp. — possibly 8. thuringiensis Berlinger)

in the haemolymph. Nothing is known of the mortality

caused by these pathogens in the field, but B

thuringiensis at least is known to be highly toxic to

E, behrit \arvae- in the laboratory, having an LC,,,

value of 311 spores/mm?, a figure 20 times lower chan

that for Helicoverpa armigera (Hiibner) (Cooper 1983),

Discussion

Irrigated lucerne crops flower and form pods later

than dryland lucerne and volunteer plants which grow

around irrigated crops, along road-sides and in

scrubland. So the number of moths available to attack

irrigated lucerne crops each year should depend on the

namber and suitability of pods available to.the earlier

generations of £. behrii in that season, and on some

Tunction of weather. However, the very low numbers

of larvae in the irrigated crop south of Keith in Jamiary-

February 1982, despite the presence of a large number

of adult moths in the vicinity (Fig. 6) and suitable

oviposition sites, suggests that some variable or

variables, other than the number of moths, may be a

key factor in the amount of crop damage that occurs

from year to year, This variable might be predation

or parasitism on eggs and young larvae, the effect of

pathogens, or some property of the plant which

prevents young larvae establishing and surviving in

pods, either of certain cultivars or in certain years. Such

a variable might also account for the unpredictability

of attack of lucerne by &. behrii, In most years damage

is so slight as to go unnoticed, but occasionally there

is avery heavy infestation in one season, Such outbreak

seasons appear from the available records to be

infrequent, but when they do occur, outbreaks are

widespread. To date, further research aimed at

examining the factors responsible for heavy infestations

of E. behrii in seed lucerne crops in South Australia,

has been hindered by the infrequency of such events.

In recent ycars there have been only low or moderate

numbers of this pyralid pest reported in seed crops in

the Keith area, possibly because of the more

widespread and effective use of pesticides since the

early 1980's.

TABLE 3. Furasitolds kriown fo attack various stages of Etella behrii in South Australia (records collected 1959-82),

Parasitnid Stage from which Primary (P) or hyperparasitoid (H)

species parasitoid emerges solitary (S) or gregarious (G)

Tchneumonidae

Temelucha cycnea Kerrich pupa P;S

Enynis sp. P;S

Braconidae

Iconeila alfalfae (Nixen) Sth instar larva P;S

Phanerctoma sp. pupa PS

Bassas sp. | Sth instar larve Ps

Bassus 4p, 2 Sth instar larva PS

Bracon sp. thaturé larva PS

Elasmidae

Elasmus: sp- pupa H:S or G

Eurytomidac

Eurytoma sp. pupa Ps

Tachinidae

? Siphonini (gen. & sp. indet.) mature larva PSE

tis) A. D. AUSTIN, T. C. R. WHITE, D. A, MAELZER & D G. TAYLOR

Acknowledgments

We wish to thank Mr A. Campbell for access to his

property, ‘Brecon, Dr David Cooper for assaying

pathogens of E&tiella, Dr Jenny Gardner for

identification of native plants, and Drs Brian Cantrell

and Mike Sharkey for identification of dipteran and

hymenopteran parasitoids, respectively. We gratefully

acknowledge the Commonwealth Special Research

Grants Scheme and the lucerne seed growers through

the Seed Section of the United Farmers and Stock

Owners of South Australia Inc. for funding this project.

We also thank Paul Dangerfield for providing technical

assistance in the preparation of the manuscript.

References

Aput-Nasr, 8. & AWADALLA, A.M. (1957) External

morphology and biology of the bean pod-borer, Friella

zinckenella Treit. (Lepidoptera: Pyralidae). Bull. Sac. ent.

Egypte Al, 591-620,

AANon, (1987) Bicnnial Report of the Waite Agricultural

Research Institute, The University of Adelaide, 1986-1987

(University of Adelaide, Adelai ie).

Austin, A.D., DurHAM, M. & Bamey, P. (1986) Etiella

Moth. Department of ‘Agriculture. South Australia, Fact

Sheet, FS 51/85, 4pp.

Common, £.F,B. (1990) "Moths of Australia.” (Melbourne

University Press, Melbourne).

Coorer, DJ. (1983) The susceptibility of Etiella behrii to

Bacillus thuringiensis. J. Aust, ent. Soc. 22, 93-95.

DurHam, M.S. (1984) Commercial monitoring of insect

pests and beneficials of seed lucerne in south east South

Australia. pp. 121-127 Jn Bailey, P. & Swincer D. (Eds)

“Proceedings of the 4th Australian Applied Entomological

Research Corference™ (South Australian Government

Printer, Adelaide),

Harpstack, A.W., Henpricks, D,E,, Lopez, J.D.,

STADELBACHER, E.A., Priuups,J.R, & Wirz, J.A, (1979)

Adult sampling. pp. 105-131. In Sterling, W.L. (Ed.)

“Economic Threshholds and ling of Heliothis Species

on Cotton, Com, Soybean and Other Plants.” Southern Co-

operative Services’ Bulletin No, 231. (Texas A & M

University, College Station).

Maenzer, D.A., Prynock; D,E., BAILEY, P-T, & MADGE,

P.E. (1982a) Pollination and pest management of lucerne

seed crops in South. Australia. pp. 121-141. /n Cameron,

P.J., Wearing, C.H., & Kain, W.M. (Eds) “Proceedings.

of the Australian Workshop on Development and

Implementation of I.P.M.” (Government Printer, Auckland).

Mapce, P.E, & Bailey, PT. (1982b)

Pollination and arthropod pest management of lucerne seed

crops in South Australia. pp. 385-392. J Lee, K.E. (Ed.)

“Proceedings of the 3rd Australasian Conference on

Grassland. Invertebrate Ecology.” (South Australian

Government Printer, Adelaide),

McMuLLen, R.D. & Jona, C (1970) The biology and

influence of pesticides on Campylomma verbasci

(Heteruptera: Miridac), Can. Ent. 102, 1390-1394.

Perrin, R.M. (1978) Varietal differences in the susceptibility

of cowpea to larvae of the seed moth, Cydia ptychora

aeytick) (Lepidoptera; Tortricidae), Bufl. ent, Res. 68,

Snorey, H.H. & HALE, D.L. (1965). Mass rearing of the

larvae of nine noctuid spp. on a simple artificial medium.

J. econ. Ent, 58, 522-524.

SincH, H, & DuooriA, M,8, (1971) Bionomics of the pea

ped Brees Fiiella zinckenella (Treitschke). Ind. J. Ent. 33,

Stons, M. (1965). Biology and control of lima bean pod-

purer in southern California. %ch. Bull. U.S. D.A. 1321,

TuRNER, JW. (1980) Insect pests of peanuts in eastern

Queensland. Qld Agric. J. 106, 172-176.

Wuatiey, P.E.S, (1973) The genus. Etiella Zeller

(Lepidoptera: Pyralidae); a zoogeographic and taxonomic

study. Bull. Br. Mus. (Nat. Hist.) Ent. 28, 1-21.

INCABATES HAMMER AND SETINCABATES GEN. NOV.

(ACARIDA: CRYPTOSTIGMATA: HAPLOZETIDAE) FROM

SOUTH AUSTRALIAN SOILS

BY D, C, LEE*

Summary

Incubates Hammer is commented on and an allied genus, Setincabates gen. nov., established. Three

new species are described from South Australian soils: . macronudus sp. nov., I. punctatus sp. nov.

and S. hypersetosus sp. nov. (type). . angustus Hammer, previously known from New Zealand, 1s

newly recorded from Australia and 1. medius Hammer is newly regarded as its junior synonym.

A key is given for the adults of these four species. This is the first record of Incubates from

Australia. An African species previously grouped in /ncabates is newly combined as Muliercula

longisaccula (Mahunka).

KEY WORDS: Incabates angustus Hammer, Incabates macronudus sp. nov., Incabates punctatus

sp. nov., Setincabates hypersetosus gen. noy., sp. nov., Haplozetidae, Acarida, soils,

South Australia.

Treasuerions of the Rawal Society of S. Aiesr., (1993), IIW(23, 77-85,

INCABATES HAMMER AND SETINCABATES GEN. NOV. (ACARIDA: CRYPIOSTIGMATA:

HAPLOZETIDAE) FROM SOUTH AUSTRALIAN SOILS

by D. C. LaB*

Summary

Lee, D. C, (1993) Incabates and Setincabates gen. mov. (Acurida: Cryptostigmata: Haplozetidae) from South Australian soils,

Trans. R. Sac. S. Aust. 117(2), 77-85, 4 June, 1993.

Incoubales Haxamer is commented on and an allied genus, Setincabates yen. nov., established. Three new species are described

from South Australian soils; /. macronudus sp. nov,, 1. punctatus sp, nov. and S. Aypersetosus sp. nov. (type). I. angustus

Hammer, previously known from New Zealand, is newly recorded from Australia and J. medius Hammer is newly regarded

as its junior synonym. A key is given for the adults of these four species. This is the first record of Incabaies from Avstralia.

An Affican species previously grouped in Incabates is newly combined as Muliercula longisaccula (Mahunka).

Key Worns: Incabates angustus Hammer, Incabates macronudus sp. nov., Incabates punctatus sp. nov,, Setincabates

hypersetosws gon. nov... sp. nov., Haplozetidae, Acarida, soils, South Australia.

Intraduction

The genus /ncabates Hammer, 1961 and a similar

undescribed genus are examined as part of an ongoing

study of sarcoptiform mites sampled from nine florally

diverse South Australian sites, and for which Lee (1987)

provided an introduction to the relevant work on the

advanced oribate mites, The relevant mites wre from

soil and plant litter or mass under either savannah

woodJand, sclerophyll forest, mallee or coastal

vegetation at only four of the sites.

Incabutes is grouped in Haplozetidae Grandjean,

1936, which has been discussed by Lee & Shepherd

(1990) in considering Magnobaies Hamamer, 1967.

Relationships between Jncabares, the new genus and

some other haplozetid genera are discussed. J. angiastate

Hammer, 1967 is newly recorded from Australia, and

two new species of Incabates and the new genus

together with one new species are described.

Material and Methods

New material examined here, collected by the author,

is deposited mostly in. the South Australian Museum

(SAMA), but also in the. Natural History Museum,

London (BMNH), the Field Museum, Chicago

(FMNH) and the New Zealand Arthropod Collection,

Lam! Care Research, Auckland (NZAC), whilst

previously described material is deposited in te

Zoological Museum, Copenhagen (ZMC). The

morphological notional system follows Lee (1987),

the somal chaetotaxy of which is.summarised in Figs

* South Australian Museum, North Terrace. Adelaide, S.

Aust_ 5000.

2 and 3, with the total setae present in cach file (e.g.

6Z) indicated by number Coming first, whilst a

particular seta (e.g. Z6) would have the number last.

The venter and legs have been described only for

Incabates punctatus because of their uniformity within

the genus, The abbreviations for zoogeographical

regions follow Lee (1970, Fig. 427), Descriptions of

eggs are based on those still within the female soma,

All material was examined using a Nomarski

interference contrast device. All measurements ate in

micrometres (zm) and were made using un eyepiece

micrometer at x250 magnification.

Key to Australian Incabates and Setincabates species

(adults)

|. Thirteen pairs of hysteronotal seta. Hysteronotal foramen

FS sacculate without narrow duct oo. eee

ety seed, s2brneadinedbeas Setincabates hrypersetomis sp. nov,

Ten pairs of hysteronotal seta or ulveoli. Hysteronotal

foramen F3 sacculate with narrow duct ...., éncabates.

Hysteronotal setae represented only by alyeoli. Tutorium

ts

absent. ......2..... saves pelle e's T, macronudus ap. nov,

At Jeast one pair of hysteronotal setae ae Tutorium

PRONE. Nyy ereesdthgtarateas wheartbeaeedtyaices 3,

3. Six posterior small hysteronctal fetal pat present

(JS, J6; ZS, 26; $5, §6). Anterior soma

punctate ose. Wirestewassg tig veuese T. punctatus sp. Nov.

One posterior small hysteronotal seta] pair present (/6).

Anterior soma not punctate -..... I, angustus Hammer.

Systematics

Incabates Hammer

Incabates Hammer, 1961, p. 108 (type species by monotypy:

Incabates nudus Hammer, 1961); Coeter, 1968, p. 25; Balogh

ot ai 1984, p, 274: Loxton, 1985, p. 67; Corpuz-Raros,

1980; 174.

78 DE

Definition: Haplozetidae. Hysteronmtem with |}

pairs (2/, 62, 25) of short setze, microsetac or alveoli

without thair setae. Hysteronotal forarnina with sarcule

bag-like, either lapermg gradually to pore or with short

narrow duct near pore, Dorsosejugal furrow entire.

Translamella and prelamelia absent, costate or lineate

tutorium (between setae jl-z2) sometimes present.

Rostral eta {jl} directly in front of tamellar sera (21).

Picromorph movable, with clear weakly sclerotized

basal Hne. Discidium triangulate. Tibia | with large

solenidium (s02) on tubercle. Tibia I without

disodorsal spur. Genu I and 0 with two setae (vy

ubsent), Trochanter [Y usually with distodorsal process

exunded over femur IV (exception! [ncabares

eugustus), distodorsal crown with only anterior

angulate lobe, broad {lange extending along entire

venter of caput, Pretarsus with three claws, slimmer

fateral claws with pointed tip,

General Morphology of Austratian species; Samal

length range for adults: 239-445 (for all species

239-496). Sowal chaetotaxy: 2/, 22, ls; U, 62, 23;

32, WT, Bil, 31% AlZg, 18g. 2 Za, 3Sa. Leg chaetotaxy

{solenidia tn parentheses): |— 1, 5, 2¢1), 442), 20(2);

D — 1, $, 2(1), 40), 16(2); I — 2, 3, 10), 300), 15;

TV — 1,2, 2, 3¢]), 12, Alveolrof somal setae bonnded

by refracule ring, canal short and tapering without

internal refractile ring. Pteromorph may lic close to

pleural surface or be lifted away from it, Sub-bothridial

flange present, Apodemes.1, ll, ventrosejugal and Il

present, Ventrosejugal gap not wider than genital

orifice, Subpedal and circumpedal ridges merged into

single continous line. Slitdike pore Saf nearly

longitudinal (less than 45° from longitudinal axis).

Proximovectral spur on ferur f without caput collar.

Conspicuous yentral flanges on femora I, Hi, and TV,

wn femur Il margin curved, not angulate.

Distrituaion: Peru (NTe}, Japan (Pc), Philippines

(Om), Australia (Aa), New Zealand (An), Fiji (Ap),

All species from outside South Australia are from moist

localities, often in moss, cither in high altitude

grassland or in forest or mangroves, and are sometimes

arboreal. In South Australia ali three species. were

collected from the svlerophyl! forest site, which bas

the highest rainfall (annual mean in the range of

1150-1200 mm, moxily in winter}, but Jnowbates

punctatus sp. nov, was also collected from the two

mallee sites Which are dry (mean annual rainfall in the

range of 350-500 man).

Remarks: Incabates was considered allied to

Protaribates Berlese, 1908 when it was established.

More recently it has been allied to Scheloribates

Berlese, 1908 (Corpuz-Raros 1980; Laxion 1985),

which it is similar in having a tangulate discidium,

hysteronotal sacculate foramina that gradually taper te

4 pore, and, for some species, in having no tutorium

fin species wilh a cutarium, it only cosinte). In comtrast,

Incabates tins a Gerived character state, a hinged rather

jEE

than a fixed pleramorpl as in scheloribatid genera, on

the basis of which, Cnetter (1968) suggested that

Incabates shwuld be transferred to the Haplovetidae

Grandjean, 1936. Because /nonbates also has a derived

trochanter LV (the distoventral crown with only an

amterior lobe that extends along entire vemer af caput),

and if there is a lateral proteronotal ndge it is a tulariam

rather than a prelamelia ridge, this grouping in lhe

Haplozetidae is followed here:

ly current classifications of the Onpodoidea, the

Haplozetidae are wit closely allied to the Protonbatidue

Balogh & Bslogh, 1984, because of the importance

given lo the former family haying derived sacculare

hysteronutal foramina, tather than jultiporose

foramina: out both Iacabates anG Protoribates have

derived hinged pteromerphs. The significance of

particular character states and so the accepted

relationships amongst oripodoid taza, may well change

considerably: In this paper it is considered that, since

Incabates shares some character statés with mest

genera of the more primitive Scheloribaudae

Grandjean. 1933, characler states thal are derived in

many members of ihe Haplozetidae (the discidium is

réctangulate, the tutorium ia canspicuously laminate

and hysteronotal sacculate foramina often have narrow

ducts leading to their pores), it is more primitive than

Magnobaies Hanmer, 967 and the genera closely

allied to it (Lee & Shephard 1990).

in distinguishing Incabates from other genera, the

following scheloribatid genera must be considered:

Muliercula Coetzer, 1968, Nennerlia Coetzer, 1968 and

Styloribates Jacot, 1934. These three genera can be

regarded as derived within the Schelortharidiae because:

of character states of the proteronotum and trochanter

IV. In general, members of the Scheloribatidae fold

feg 1 against the soma so that the tarsus pomls

downwards behind seta 1, lying behind the prelamella

tidgs, also trochanter [V is simple, with the crown

contined to the distal end, In contrast, haplozetids and

the three scheloribatid genera listal above, have a

tutorium of subtutorium. rather than a complete

prelamells and, when the leg is folded, tarsus I can

paint forward between setae zl-jh, whilst on truschanmer

IV, the crown extends as a broad flange along the enture

vemter of the caput. Therefore, these three scheloribatid

genera are wnly distinguished from the faplozericd

Jncabates by laving fixed pteromorplis, which ts

sometimes a difficy]| character state to assess. Furiber

studies may require that these four genera dre cither

grouped into a new family or that the Scheloribatidae

and Haplozetidae are merged.

The naining of a species, that is very similar to Ihe

type species of Mulierculz, as Uncabates longisaccutus

Westrates this confusion. It is here combined as

Muldiercida longisaccula (Matiunka, 1984) comb, now.,

despite its lack of a tutocium and the presence of 2

partial prelamellar ‘This is partly because these

INCABATES AND SETINCABATES, HAPLOZETID MITES 7

character states are variable in Jncalenes, aud therefore

the presence of a tutorium may not he diagnostic of

Muliercula, but it is mainly because the pteromerph

is only partially delineated from the hysteronotnm by

a Clear furrow as described for Muliercula, where it

is regarded as fixed, although ambiguously described

by Coelzer (1968) as “pteromorphae immovably

hinged”. Because of the similarity between Incahares

and Muliercula, despite their grouping in: separate

families, only the more extensive wing-like expansions

of the lamellae of M. fongisaccula and its occurrence

in Afriva support the pew grouping.

Tn comparison te other haplozelid genera such as

Haplozeres Willmnann, 935 and Magnabates Hammer,

1967, Incabates ts regarded as prinufive in having a

triangulate disctdium, as on oribatulid and

scheloribatid adults, Similarly it has only two setae

on gen I apd the alyeolar canal of te somal setae

is short and simple,

facabates includes seven species: Lb angustus

Hammer, 1967 (= 7. meafus Hammer, 1971, syn, nov.)

(rom Australia (Aa), New Zealand [An) and Fiji (Ap);

{. mactrenudus sp. nov. fron Australia (Aa); 1 major

Aoki, 1970 from Japan (Pe): 2 mudus Hammer, 1961

from Peru (NTc) and Philippines (Om); |, pahabaeus

Corpuz-Raros, 1980, from Phillppines (Om); /

punctatus sp. noy. from Australia (Aa); J. strianas

Cormiz-Raros, 1980, from Philippines (Ons).

Incabates angustus Hammer

fncabates angusnss Hammet, 1967, p. 43, 44, Fig. 57, S72.

facutates medias Hanuner, (971, p. 42, Fig, 49. 49a, syn, noe,

Type material examined: J, angustus, lectorype 9

{labelled “type” us vial of alcohol, three specimens

recorded in Orginal description) examined (ZMC}.

liverworts and sewall ferns, Gead tree trunk, native

forest, Waitakere National Park. North Island, New

Zealand, M. Hammer, 1962. f. madius, lectotype @

(labelled type in vial of alcohol, two specimens

recorded |n original description) examined (ZMC),

withered leaves, river bank, above mangroves,

Corolevu, Vita Levu, Fiji Islands, M. Hamner, 962-

Female: Soma oval, brown. Idiosomal length

(original description from New Zealand: ‘about

0,35 mmm, from Fiji; ‘about 0305 mm), 335 (n = 1,

New Zealand) of 318 (n = 25, Sclerophyll forest,

306-332), Leg lengths (femur-tarsus, Sclemphy!l

forest. for 320}: 1-142, 0-127, 1-108, I'V-137. Tibial

maximum heights (for 320): 1-22, (1-17, 1-14, [V-14,

Proteronotum with mediuny width fostruin,

Integument smocih, ‘Tuionum. present, sometimes

indistinct level] with lameflar seta (21). Sensory seta (z2)

with globular caput longer than exposed stalk. Sub-

hothridial Mange inconspicuogs, Posterior margin of

bothridium raised to form tooth-like spur. Imerlamellar

seta (72) longer than distance j2 ~ zl, Hysteronotum

with only one pair (/6) of noticeable small setze, nine

pairs (J5, 21-6, S5 and 6) of alveoli, possibly with small

Mucrosetae, Alveolus $5 posterior m ZS (South

Australian specimens) or anterior to ZS (New Zealand

Specimens). Foramen F3 conspicuously lazer and

sometimes sacculus bilobed (South Australian

specimens) or shghtly larger, mot bilobed (New

Zealand specimens),

Idjosternal setae fine and short, conite seta 3, J

and J¥3 (not Ulustrated as present by Hammer, 1967:

Fig. 37a) longest. Integument smooth excepi for

indistinct reticulations around ventrosejugal apodeme

and apodeme TIT. Discidium height less than 05x

height of pedotectum MT, coxite seta JV3 néar discidium

base, Eggs subellipsoidal, exochorion smooth, mean

size 151 x 72 (n = 7), length 48% of somal length,

eges per female — | (n = 3), or 2 (mn = 2).

Legs short (median fermur-tmarsus length: 40% of

somal lengeh) with stout girth (mean maximum tibial

height 50% of mean length). Trochanter ['V anterior

margin parallels posterior margin of femur fY¥ caput,

angulate distodorsally but without process extended

over femur I'V.

Male: Similar to ferale but idigsoma shorter, mean

length, 293 (7, Sclerophyll forest, 283-303) or 319 G1,

Sayatinslt woodland)

Referred material: 12 females (SAMA NIQ9iL —

NI99112), seven males (SAMA NI99113 — NI99119),

plant litter, sparse moss and sandy soil, under

sclerophyllous shrub arnongst messmate stringybark

(fucalyptus obliqua), dry scleropbyll forest, near

summit of Mt Lofty (34°59°S, 138°45'B), Cleland

Conservation Park, 9.v.J974,

One male (SAMA N199120), grass, moss, plant litter

and loamy soil, under manna gum tees (Eucalyptus

vininalis), savannah woodland, Chambers Gully

(34°S8’S, 138°41’E), Cleland Conservation Park,

12.¥i, 1974.

Remarks; Incabates angustis was ane of four species

that have only a single pair of bysteranotal seta (/6),

but one of the other three species, | medius from Fiji,

is here synonymised with it. The other two species,

I. striatus with a stviated intégument and the stouter

1, nudus with a much broader lamella, are easily

distinguishable as separate species, The slim /,

angustus and f, medius are very similar with the new

material from South Australia having intermediate

character states, /. angustus is 350 wim long, has

hysteromotal seta 96 anterior to $$ and the imerlamellar

Seta (f2) & about 0.75% the length of the lamellar seta

zi). £ medius is 305 xm long, has hysteronotal seta

56 level with $5 and the interlamellar seta is about

O.5x the length of the lamellar seta. The specimens

from South Australis are similar in size ta J, medins,

have seta 96 posienor to 55, und the relative sizes of

the interlamellar and lamellar sete are as J. angusuce.

80

D, C. LEE

Fig. 1. Incabates macronudus sp. nov., female soma, notum.

1004um

INCABATES AND SETINCABATES, HAPLOZETID MITES BL

These differences are here regarded as intraspecific

variations within a species distributed across Australia,

New Zealand and Fiji, and comparable with

differences within J nudus from Peru and. the

Philippines (Corpus-Raros 1980).

Incabates macronudus sp. nov.

Fpe material

Holotype female (SAMA NI99121), plant litter,

Sparse moss and sandy soil, under sclerophyllous shrub

amongst messmate stringybark (Eucalyptus obliqua),

dry sclerophyll forest, neat summit of Mt Lofty

(34°59’S, 138°45’E), Cleland Conservation Park,

9yv.1974.

Female: Soma oblong, brown. Idiosomal length 446

(n = 1). Leg lengths (femur-tarsus for 446): 1-178,

II-163, II-142, TV-166. Tibial maximum heights (for

446): 1-22, I-17, I-14, IV-l4.

Proteronotum with narrow rostrum. Integument

smooth, Sublamella obscured in dorsal aspect by

laminar lamella, Tutorium absent. Sensory seta (z2)

3

100,.um

Figs. 2, 3. Incabates punctatus sp. nov., female soma. 2, notum; 3, idiosternum

Fig. 4. Incabates punciatus

setae illustrated (vy = ventral).

clavate (caput may appear globular if viewed end on),

caput subequal in length to exposed stalk. Sub-

bothridial flange conspicuous. Posterior margin of

bothridium raised to form tooth-like spur. Interlamellar

seta (j2) longer than distance j2-zi. Hysteronotyum with

no obvious setae. Alveoli form clear refractile rings,

mictoseta recognised in Z2, may be present elsewhere.

Extra alveolus between Z2 and Z3 on right side only.

Foramina with conspicuously refractile saccule, F3 oval

and larger than other saccules.

Idiosternal sctae fine and short, setae /3 and Sal the

longest, setae 113 and I'V3 not located. Integument

smooth except for indistinct reticulations around

ventrosejugal apodeme and apodeme IIT. Genital seta

JZg2 midway between JZg] and JZg3, rather than

closer to JZg1, Discidium height less than 0.5x height

of pedotectum Il, coxite seta /V3 not located. Egg

. noy., female right legs I-IV, femur-pretarsus, also trochanter IV, posterior aspect. Only two

subellipsoidal, exochorion smooth, mean size 166 x

70 (n = 4), length 42% of somal length, four eggs

in single female.

Legs short (median femur-tarsus length: 36% of

somal length) with stout (mean maximum tibial height

39% of mean length). Trochanter IV with blunt process

on angulate distodorsal margin which extends over

femur LY.

Remarks: The name macronudus is from the Greek

for ‘large’ and the Latin for ‘naked’ and refers to its body

size and the absence of recognisable setae on the

hysteronotum. I. macronudus is the only species known

in Incabates with no easily observable hysteronotal seta

(I. nudus has one pair) and with J. major it has a somal

length greater than 440 um. Because there is only one

specimen, it was not dissected, which may explain why

some coxite setae (7/3 and IV3) have not been located.

INCABATES AND SETINCARATES, HAPLOZETID MITES &3

Incabates punctatus sp. nav.

Fype material

Types: Holotype female (SAMA N1987645), soil,

plant litter ami sparse moss under ridge-fruited mallee

(Eucalyptus éncrassata) clumps amongst broombush

shrubs (Mefalewcu wneinata), Fetries-McDonald

Conservation Park (35°1S’S, 139°O9'E), 20vi, 1974,

Paraty pes, 45 females (SAMA NIG87646 — NIGRTES4,

NI99122 — N199142, five — BMNH, fve — FMNH,

five — NZAC) and 55 males (SAMA N1987655 —

NIS87671, NI99I43 — NI99165, five — BMNH, five

— FMNH, five — NZAC}), as holotype.

Female: Sama oval, brown, Idiosomal lengta 259

(n = 25, Mallee broombush, 249-267), 257 (n = 2,

Mallee heath, 256, 258) or 260 (n = 3, Sclerophyll

forest, 252-267), Leg lengths (ferwir-tarsus, Mallee

broombush, fioy 262): 1-113, 1-98, 111-82, PY-105, Tiblal

maximum heights (for 262): 1-16.5, 1-13, M-11.5, TV-13,

Proteronotum with medium breadth rostrum.

Integument punctate anterior to sela 72, Sublamella

mainly obscured in dorsal aspect by laminar lamella,

Costale tutortum present. Sensory seta (22) clavate,

caput subequal in length to exposed stalk. Sub-

bothridial flange conspicuous. Posterior margin of

bothridium raised to form small single or double tooth-

like sour, Interlamellar seta (j2) shorter than distance

2-21. Hysteronorum without obvious setae anteriorly

(Zl, Z2, Z3, ZA) but microsetae recognised for ZI and

Z2, whilst posterior small setae (/5, J6, 25, Z6, 55,

S56) present, Integuiient punctate anterior to

fteromorplis. Foramina with conspicuously refractile

saccule, £3 spherical and Larger than other saccules.

Idiosterna! setac fine and short, variable relative

lengths, but JA, (7fi, JZg3 always shortest. lotegument

with indistinet reticulate sculpturing and punctate in

coxistemnal region. Genilal seta JZg2 closer to JZyl

than JZ23. Thscidom height more than 05x height

of pedotectum I, coxite seta /¥3 level with, and close

to apex, Egg subellipscidal, exochorion smooth, mean

size 132 x f4(n = 13), length 49% of somal length,

usually one egg per femats (n = 14), two egys in single

fernale-

Legs short (medion ferur-tarsus length: 38% of

somal length) with very stout girth {mean maximum

tibial height 51% of mean Jength). Trochanter ['Y with

sharp process on angulale distodorsal margin which

extends over femur IV,

(dale: Sienilar to female but idiosomal sherter, mean

length, 239 (25, Mallee broambush, 226-252), 249 (5,

Mallee feath, 238-250), 251 (3, Coastal, 250-253) or

247 (3, Sclerophyll forest, 244-252),

Referred material. Three females (SAMA NIOS7672,

Ny99174, NIG9I7S) and five males (SAMA NIS9I76 —

Ni99180), sand, plant litter, wnder Banksia shrubs

(Banksig omata), amongst other sclerophyllous shrubs

and sparse brown stringybark mallee (fucalypis

bzxteri), Tainboore Homestead (35%57'S, 140°29'F),

ar Mt Rescue Conservation Park, 4,vii.1974.

Three males (SAMA NI99T71 — NI99173), soil,

plant litter and sparse grass under coastal wattle

(deacie.rophorae), Piccaninnie Ponds Conservation

Park (39°03'S, 140°S7'B}, 3.vii.1974.

Three females (SAMA NI99166 — NIY9168), two

males (SAMA NI99169, NI99170), plant litter, sparse

moss and sandy soi), under sclerophyllows shrub

amongst messmate stringybark (Eucalypnes obliqua),

dry sclerophyll forest, near sunnmit of Mt Lofty

(34°S9'S, 138°45'E), Cleland Conservation Pazk,

9yv, 1974.

Remarks: The name punctanes is From the Latin for

‘dotted’ and refers to what may be small pits on anserior

Parts of the soma, which wre more extensive than in

olher species, Also /. punctatus is unique within the

genus m lacking four anterior purrs of frysteronatal setae

(microsetae may be present), whilst the posterior six

pairs have small setae, that are relatively long for

Incabates. J. pahabacus Corpus-Raros, 1980 is similar

but without such 4 clear difference in size between the

two proups of setae. Also, on /. pahabeeus, the

interlamellar setae are much longer and the hysterosotal

alit-shaped pore A/S is short and not as long as seta J6,

Genus Setincabates gen, mi,

Type-species: Serincabates hypersetesus sp. now,

Definition; Haplozetidae. Hysteronotum whh 3

pairs (4J, 6Z, 35) of miedium-sized setose setae,

Hysteronotal foramina with saccule bap-like, gradually

tapering (0 pore, Dorsosejugal furrow entire.

Translamella, prelamella and tutorium ahserni_ Rostral

seta {fl} directly in front of Jamellar seta (z)).

Pteramorph movable, with clear weakly sclerotized

basal Ine. Discidium triangulate, Tibia I with large

solenidium {se2) on tubercle. Tibia Uf withour

distodorsal spur. Genu | and Tl with peo setae {vy

absent), Trochanter IV with distodorsal process

extended over femur IV, distadorsal crown with only

anterior angulale lohe, broad flange extending along

entire venter of capul. Pretarsus with three claws,

slinumer lateral claws with pointed tip.

Distribution; Australia (Aa). Only recosd from South

Australia, ai site which amongst those sampled has the

highest rainfall,

Remarks: The name Serincabates has the prefix Ser,

an abbreviation of the Latin seta for ‘bristle or hair’,

und refers to the presence of more hysteronatal setae

on the single included species than on members of the

samilar /ncabates and other baplozetine genera with

short or medium-sized setae. The haplozetine

Flagellobates Mahunka, 1973 with long hysteronotal

seme has 14 ppits of hysteronmal setae. Some other

subfamilies included an Haplozetidae by Balogh &

Balogh (1984) have similarly numerous hysteronotal

84 D.C. LEE

setae, Members of the Peloribatinae, all of which have

long hysteronotal setae, have either 14 pairs or, in

Acutozetes Balogh, 1970, 13 pairs of hystetonotal setae

(possessing six not five setae in file J). The only other

haplozetid species with more than 10 pairs of

hysteronotal setae are members of the Pilobates Balogh,

1960 (Pilobatinae) with 14 paits and six pairs of genital

setac and a straight, transverse complete ventrosejugal

apodeme and one species of Rustrozeres Sellnick, 1925

(Rostrozetinae) with 74 pairs and a short fissure-like

pore 43 similar in length to the hysteronotal foramina.

The character states of Setincabates recognised here

as distinguishing it from Mcabetes are probably

primitive.

Setincabates hypersetosus sp. Nov,

Type material

Types: Holotype female (SAMA NI99181), plant

litter, sparse moss and sandy soil, under sclerophyllous

shrub amongst messmate stringybark (Eucalyptus

obliqua), dry sclerophyll forest, near summit of Mt

Lofty (34°59'S, 138°45'E), Cleland Conservation Park,

9y.1974. Paratypes, 21 females (SAM NI99182 —

Ni99196, two — BMNH, two — FMNH, two —

NZAC) and 21 males (SAM NI99197 — NI99IIIL, two

BMNH, two — FMNH, two — NZAC), as holotype.

Female: Soma oval, brown. Idiosomal length 264

(22, 254-275). Leg lengths (femur-tarsus for 267);

1-22, 11-104, 111-86, [V-110. Tibial maximum heights

(for 267): 1-19, 11-13, 0-13, [V-14.

Proteronotum with broad rostrum. Integument

smooth except for punctuations on gnathosternum

between postoral setae. Sublamella mainly obscured

in dorsal aspect by laminar lamella. Tutortum not

present, although short ridge present around base of

rostral setae (jl). Sensory seta (22) clavate. with long

ellipsoidal caput more than twice length of exposed

stalk, Sub-bothridial flange conspicuous. Posterior

margin of bothridium rounded without tooth-like spur.

Imerlamellar seta (j2) about x2 distance j2-zl.

Hysteronotum with I4 pairs of small setae (5/, 62, 35),

Foramina with small oval refractile saccule without

parallel sided duct, all similar in size.

Idiosternal setae fine and short, peripheral setae

longer, 72 and JZa3 longest. Integument with indistinct

reticulate sculpturing in coxistemnal region. Genital seta

JZgi closer to JZgl than JZg3. Discidium height less

than ).5x height of pedotectum I, coxite seta /V3 near,

but posterior to apex. Egg subellipsoidal, exochorion

smooth, mean size 132 x 65 (n = 4), length 49%

of somal length, one egg per female.

Legs short (median femur-tarsus length: 40% of

somal length) with very short girth (mean maximum

tibial height $4% of mean length). Trochanter I'V with

short sharp process on angulate distodorsal margin

which just extends over femur IV.

Male: Similar to female: but idiosoma shorter, mean

length, 239 (21, 244-285).

Remarks; The name Aypersetosus is the Latin for

‘bristly’ or ‘setose’ with the Greek prefix for ‘beyond’

or ‘over’ and refers to the unusually extensive

hysteronotal setation compared with members of genera

similar to Sefincebates.

10940

Fig, 5. Setincabates kypersetosus gen, & sp. nov., female

soma, notum,

INCABATES AND. SETINCABATES, HAPLOZETID MITES 8S

Acknowledgements

I am indebted to Ms C. M. Birchby for the

illustrations and to the Australian Biological Resources

Study for funding her salary. Thanks are also due to

Dr Henrick Enghoff (Zoological Museum,

Copenhagen) for making types available, and Mrs D,

Lowery for typing the manuscript.

References

Aokt, J. (1970) Description of Oribatids Mites collected by

smoking of trees with insecticides. I. Mt Jshizuchi and Mt

Odaigatiara. Bull. nam, Sci. Mus,, Tokyo 13, 585-602.

BALoGu, J. (1960) Descriptions complementaires d’Oribates

(Acari) d’Angola et du Congo Belge. Publ. cult. Co. Diam.

Ang. Lishoa 51, 89-105.

(1970) New oribatids (Acari). from New Guinea. II

Acta zool. hung. 16, 291-344,

—____ & BaLoau, P, (1984) Review of the Oribatuluidea

Thor, 1929 (Acari; Oribatei). Acta zool. hung. 30, 257-313.

ee FT: A, (1908) Elenco di generi e specie nouvi. Redia

15,

Coerzer, A. (1968) New Oribatulidae Thor, 1929 (Oribatei,

Acari) from South Africa, new combinations and a key to

the genera of the family. Mems Inst. Invest. cient. Mocamb.,

Sér, A, 9, 15-126,

Corpuz-Raros, L. A. (1980) Philippine Oribatei (Acarina)

V. Scheloribates Berlese and related genera (Oribatulidae).

Kalikasan, Phil. J. Biol. 9, 169-245,

Granpigan, F, (1933) Etudes sur le development des

Oribates. Bull. Soc. zool. Fr. 58, 30-61,

———— (1936) Observations sur les Oribates (10e serie). Bull.

Mus, natin Hist. natur., Paris (2) 8 246-253.

HAMMER, M, (1961). Investigations on the oribatid fauna of

ue snes Mountains, Part Il. Peru. Biol. Skr, 13, 1-157,

43 pls.

——_— (1967) Investigations on the oribatid fauna of New

Zealand, Part II. Ibid, 20, 1-70, 40 pls.

(971) On some oribatids from Viti Levu, the Fiji

Islands. Ibid. 16, 1-60, 35 pls.

Jacot, A. P. (1934) Some Hawaiian Oribatoidea (Acarina),

Bull. Bernice P. Bishop Mus. 121, 1-99, 16 pls.

Lee, D. C, (1970) The Rhodacaridae (Acari: Mesostigmata);

classification, external morphology and distribution of

genera, Rec. S, Aust, Mus. 16, 1-219.

(1987) Introductory study of advanced oribate mites

(Acarida: Cryptostigmata: Planofissurac) and a

redescription of the only valid species of Constrictobases

(Oripodoidea), Ibid. 16, 35-42,

——— & SwEPHERD, K. J. (1990) Magnobates (Acarida:

Cc igmata: lozetidae) from South Australian soils.

Thons. Ri Soe St 14, 179-186.

LuxTon. M. (1985) Cryptostigmata (Arachnida: Acari) — a

concise review. Fauna N,Z. 7, 1-106.

Mauunxa, §, (1978) Neue und interessante Milben aus dem

Genter Museum XXXIV. A compendium of the Oribatid

(Acari) Fauna of Mauritius, Renunion and the Seychelles

Is. LL. Revue suisse Zool. 90, 709-724.

——— (1984) Oribatids of the Eastern Part of the Ethiopian

Region (Acari). V, Acta zool. hung. 30, 87-136.

WILLMANN, CC, (1935) — Faunistisch-kologische

Untersuchungen im Anningergebiet IV. Die Milbenfauna.

I, Oribatei. Zool. Jb. Syst. 66, 331-344.

BRYOZOA IN COORONG - TYPE LAGOONS, SOUTHERN AUSTRALIA

BY MARGARET SPRIGG & YVONNE BONE*

Summary

Unilaminar, multiserial colonies of living Bryozoa (Conopeum aciculata) are widespread

throughout the Coorong Lagoon, South Australia. The bryozoan is an opportunistic species that is

able to tolerate variable salinities. It establishes itself early in spring and reaches maturity prior to

the drying-out of its nearshore environment in late summer, whereupon it dies. It is intimately

associated with serpulid growth in the northern Coorong Lagoon. C. aciculata has also been found

growing in the hyposaline waters of Lake Clifton, Western Australia, where it is found within

clotted, calcareous thrombolites. The age of the Lake Clifton bryozoan colonies is unknown.

Sub-Recent buildups of C. aciculata are extensive along the north-eastern side of the Coorong

Lagoon, between Magrath Flat and 2.5 km south of Salt Creek. The colonies are multiserial and

multilayered. The architecture of these buildups differs, with those in the northern lagoon being

densely packed, flat to globose coalescing mounds whereas those in the more saline southern

lagoon are loosely packed, highly contorted and convoluted "pavements". Serpulid association with

these Sub-Recent bryozoans is minor.

KEY WORDS: Bryozoa, Coorong, hypersaline lakes, buildups, serpulids.

Transactions of lie Royal Society of S. Aust. (1993), 117(2), 87-95.

BRYOZOA IN COORONG + TYPE LAGOONS, SOUTHERN AUSTRALIA

by MARGARET SpPRIGG & YVONNE BONE*

Summary

Spricc, M. & Bone, Y. (1993) Bryozoa in Coorong-type lagoons, southern Australia. Trans. R, Soe 5. Aust.

U7(2) 87-95, 4 June, 1993

Unilaminar, multiserial colonics of living Bryozna (Conopewmn aciculala) are widespread throughout the Coarang

Lagoon, South Australia. The bryozoan is an opportunistic species that is able to tolerate variable sallnilies 1

establishes itself early in spring and reaches maturity pelor to the deying-~ut of its nearshore environment in

late summet, whereupon it dies, It is intimately associated with serpulid growth in the northern Coorong Lagoon,

C. aciculara bas alse been found growing in the hypogaline waters of Lake Clifton, Western Austrafia, where

it is found within clotted, calcareous throtnbolites, The age of the Lake Clifton bryozoan colonies is unknown.

Sub-Recent buildups of C. aciculata ure extensive along the north-eastern side of the Coorong, Lagoon, berween

Magrath Flat and 2.5 km south of Salt Creek. The colonies are imulliserial and multilayeréd. The architecture

of these buildups differs, with those in the northern lagoon being densely packed, flat to globose coalescing mounds

whereas those in the more saline southern Jagoon are loosely packed, hiphly contorted and convoluted “pavements”,

Serpulid association with these Sub-Recént bryozoans is minor,

Ker Wonns. Bryozca, Coorong, hypersaline lakes, buildups, serpulids.

Introduction

The soothern Australtan continental shelf and its

coastal inlets (Fig. la) have provided an tdeal

environment for prolific bryozoan growth and

preservation throughout the Cainozoic Era,

Accumulations of their calcareous skeletons attain great

thicknesses over tens of thousands of square kilometres

of both open water and protected embayments. Their

deposits extend from high-tide levels about protected

and open cousts to beyond the edge of the continental

slope (James & Bone 1992; Bone & James 1993).

The extension of bryozoan growth into sub-coastal

lagoons is less well known. Today, an encrusting

chetlostome anascan, Coneperm aciculata

(MacGillivray, 1891), ts found in South Australia's

Coorong Lagoon (Fig, 1b), where it tolerates salinities

that range from well below sea-water through to

hypersaline. This bryozoan was formerly reported as

Membranipora aciculata (Bone & Wass 1990: Bone

1991), but has now been confirmed as C. aricudata.

A millenium ago it thrived in the Coorong waters when

they were only marginally more saline than the sea.

Another occurrence of whal is believed to be the

same species (Bock pers. comm.) has recently been

discovered in hypesaline portions of Lake Clifton,

abour 100 km south of Perth, Western Australia (Fig,

ley, The sub-Recent history of the Lake Clifton

example is not known.

Interestingly, both occurrences ure found in sub-

coastal interdunal lagoons facing open oceans in mid-30

degree south latitudes, Both lagoonal systems ans

* Department of Gellegy, and Goophysics, University of

Adelide, Adelaide, S, Aust. 5005,

subject to a degree of ground water drainage via highly

porous and permeable, semi-consolidated , aealianitic,

Pleistocene back-shore beach dunes, Both lagoons are

similar in that significant salinity differences are

manifested along their length, and these fluctuate

according to seasonal water influxes. Holocene

dolomite is found in both systems (Rosen & Coshell

1992), Lake Clifton was probably once a continuation

of the present day Harvey Estuary (Moore et al. 1984)

into Which River Murray waters from the adjacent

Darling Ranges flow at its northern, Peel Inlet end,

but it is now cut off from the sea.

The Coorong still receives influxes of water directly

{rom both the sea and its own Murray River, Although

the Coorong and Lake Clifton environments possess

these basic similarities, they alsnm have significant

differences in detail and salinity fluctuations which

affect bryozoan establishments and growth.

Tn this paper, the Bryozoa and their environments

are compared and summarised, with the emphasis

placed on the Coorong example.

Geological Settings

The Coormng linear lagoons and Lake Clifion are

products of repeated oscillation of Quaternary sea level

and regional tectonic uplift. This has resulted in a

sequence of abandoned sea beaches across a width

suitable to create significant separation of respective

sea coasts and their fossilised backshore dune deposits,

both oo and off-shore. This is more apparent in the

South Australian example.

The present Coorong Lagoon lies ih the latest

interdunal corridors of successive strandings of parallel

Ocean-beach and backshore dunes of the Bridgewater

88

Formation. This is a highly calcareous cross-bedded

medium to coarse grained aeolianite. (Sprigg 1952).

Similarly, Lake Clifton lies between Jinear shore-

parallel Pleistocene ridges of calcareous sandy Tamala

Limestone. These ridges show large scale cross

! GREAT

| BARRIER

j i REEF

AUSTRALIA

\ F

| |

SHABK WESTERN :

BAYS AUSTRALIA. — |

| Sou

AUSTRAL 1A

LAKE

CLIFTON

“STUDY AREA

eke

\Prertoal Th Bunbaty

Fig. 1. la: Map of Australia showing the locations of the Coorong Lagoon and the Lake Clifton area.

Location Map. Ic; Lake Clifton Location Map,

M. SPRIGG & Y.BONE

bedding, and are composed of fine to coarse grained

skeletal-fragment calcarenite with variable amounts of

quartz sand (Playford & Leech 1977),

In both the Coorong and Lake Clifton regions fossil

soils with rhyzoliths have developed at various levels

VQ aa fe Milang

RIVER s ee Lake Alexandrina

M Y

ai ‘hag HINDMARSH ISLAND

Pelican Pt, —\

Mark Pt A

4

a -

S\Na MENINGIE

a\t

i

6

7" Bonney

Oy Reserve

,

Magrath

Hells Gate

<

9 ua

a Zz

< é

=: §

>

Pollcamans Pt.

* Gemini Downs

lisalt Creeh

2 aS a

BUL BUL BASIN As

a q

J LACEPEDE

x STROMATOLITE

am «LAKE

5 SHELF

_)

et

2

SCALE: kms

0 5 10 15

mmm Barrage

Ib; Coorong Lagoon

BRYOZOA IN COORONG sy

within the dunes. These dunes in places include

calcretised layers, if the soil forming periods were of

sufficient duration. The Lake Clifton Tamala

Limestone has calc rete-lined solution pipes (Semeniuk

é& Searle 1985) that are similar lo those in the Coorong

(Bone & Wass 1990). These solution pipes are features

of calerete profiles, and are formed tube-like, by

calcium carbonate precipitation around regions of net

vertical water percolation (Scholle et al. 1983) through

limey dunes, In the Coorong area, erosion has removed

much of the original dunes so that many of these large

solution pipes are free standing by the water's edge,

where they appear like the remains of petrified forests.

The Coorong and Lake Clifton interdunal areas were

both inundated by the sea during the Holocene. and

in euch case, the consolidated aeolianites or their

calcrete cappings provide excellent bases for bichermal

buildups. (Fig, 2).

The Lake Clifton example

The rise in Sea Jevel during the Holocene caused

some of these shore-parallel limestone ridges in the

southwest of Western Australia to become inundated

and form a lakeland system of linear barrier estuaries

and lagoons, including Lake Clifton (Fig, le; Searle

& Semenivk 1985), The ridges, which enclose the

lagoon and separate it from the Indian Ocean 1.5 km

awity, dip below sea level just north of the lake and

run across the narmow bryozoan-rich sediments

A BRYOZOA HORIZON

(Memoranipara acicuiald)

spray ore

Aray zane rhyzalilhs

splast) sone

SuUPTANdal gone

ertiaal Zone

OW Water. he

orey muddy

sand

fool clasts

centoned layet highly foneyeamped

of Bryozpa region

(forming bionarns)

ra ~s*,

Puan of + ~ oS,

Gemini Pont py Te

pron ecaley i “a

Jt

at ,

i

Q Pp

Se

a rg

cantoites 74 y

bryozoai bed a

i)

> 10mm thick

limit of

\ Bryomge \

rool moulas & chantels

Sandy sol)

Seg TF

of the Rottnest Shelf (Collins J988) ro re-emerge at

Rottnest Island (Playford & Leech 1977),

The 21.5 km long, 1.0 km wide lake is replenished

by winter mains falling directly on to it and by

underground water [rom un extensive aquifer along its

eastern shore. The lake is shallow, much of it being

Jess than 1.0 m and a maximum of 3.0 m deep (Fig.

3). Water levels seasonally Muctuate up to a metre,

periodically exposing and desiccating the large areas

of thrombolites growing there.

A bryozoan, tentatively identified as C. aciculata,

intimately grows over and through the clotted

caleareous thrombolites (Fig. 4), The largest colony

found was 2 cm in diameter, The thrambolites (Burne

& Moore 1987) outwardly appear similar to

stromatolites and are constructed in part by the trapping

and binding of detrital sediment by cyanobacteria. As

Well as supporting Bryozoa. the lithoherms provide

shelter for communities of fish, amphipods, isopods,

decapods, nenid worms and various insects, The

spaces between the clots often become filled with

sediment rich in small gastropod (e.g. Cariella sp.)

and ostracod shells.

The salinity of Lake Clifton remains less than that

of sea water throughout the year, though other lakes

in the same system range from) hyposaline to

hypersaline. Their salinity and water chemistry, and

that of the regional ground water, were investigated

by Moore (1987) to elucidate why thrombolites were

restricted to Luke Clifton. She found the waters of

x

karst Suflace ‘ cn

OHalky 2are

endonline es

nenens

y

speleathion

HARUPAN -

Plaly “calerete"

ae xy) > an

4 £52 woh

t shallow lagoon & sandspe

( 9990)

ABN Quart’ caroonate Marert 7990)

{shelly sands)

~%

Fig. 2. Gemini Downs, Coorong Lagoon. Section A-A, through headland, showing relationship of bryooun bioherms to

caleretised acolianites.

90 M, SPRIGG & Y.BONE

Fig. 3. Lake Clifton shallow water, hyposaline environment

- reeds at margins in contrast to Chenopodiaciae rimming

the Coorong Lagoon. Figure (holding a thrombolite) for

scale.

all the lakes had ionic compositions proportionally

similar to sea water, but that Lake Clifton, which

receives virtually no surface run-off, obtains a large

volume of its annual water from low salinity (1-2 %)

ground water. This water is enriched in calcium and

bicarbonate (Moore 1984) which locally modifies the

chemical composition of the sediment-water interface.

Moore (1987) found the calcified thrombolites were

consistently associated with the aquifer’s discharge area

along the lake's eastern shore, It follows that this, then,

must also apply to the Bryozoa. Her study suggested

the debouching ground waters provide a chemical

environment conducive to the formation of the

thrombolites that the Bryozoa encrust.

The Coorong example

Geological Setting

The Coorong (Fig. Ib) is a modern artefact of an

ongoing process of successive strandings of ocean-

beach backshore dunes straddled across the northern

limb of the regionally uplifting Mt Gambier Arch. This

uplift resulted in Tertiary seas withdrawing from the

Fig. 5. Sub-Recent laterally extensive buildup of Conopeum

aciculata in the Coorong Lagoon. Width of photo is 3 m.

Fig. 4. Clotted, calcareous thrombolite from Lake Clifton.

The bryozoan Conopeum aciculata is most densely

intertwined in the central portion of the thrombolite.

Murray Basin. Relatively rapid sealevel oscillation in

response to dramatic climatic change left sub-parallel

coastlines which can be tied to Milankovitch insolation

signatures (Sprigg 1979; Idnurm & Cook 1980). These

roughly coincide with the perturbation cycles in the

earth's orbit during the late-Quaternary.

A sequence of “palaeo Coorongs” exists between

ocean-beach backshore dunes to the east of the present

Coorong system. These dunes have been preserved by

uplift and low surface drainage over the Tertiary karst

topography.

Sediment cores taken from the Coorong suggest that

initially the Coorong was a_ protected-marine

environment, becoming progressively more restricted

and subject to harsh salinity fluctuations (Gostin e¢ al.

1988). Stable 80 isotope studies of the sub-Recent

buildups of C. aciculata in the Magrath Flat area

indicate that the Coorong was only marginally more

saline than sea water when the Bryozoa grew there

around a millennium ago (Bone & Wass 1990).

Later low stands of sealevel have resulted in the

t

~~

as \

Fig. 6, Colonies of Conopeum aciculata encrusting bottle

found at Magrath Flat in early 1990.

BRYOZOA IN COORONG S|

exposure of earlier formed carbonate flats (von der

Borch 1976), These for terraces around both major

lagoons and also around the extensively eroded and

modified beach that forms an intervening string of

shoals and islands in the southern Lagoun. These latier

lie between the Younghusband Peninsula barner beach-

dune system and the earlier consolidated backshore

calcareous dunes.

Hydrology

The elongate nature of the Coorong estuary in an

aca of low, winter raintall and high, sumujier

evaporation, and the restriction of walertlow between

its hwo major lagoons, results ina longitudinal salinity

gradient, increasing southwards (Noye 1974). ‘This 1s

compounded by slow mixing of less dense river water

from the Murray, flowing over denser Coorong waters

when levels between the sea and lagoon are appropriate

(Geddes 1987). Tide, wind direction, currents,

irreyzular bottom topography and barrage openings all

contribute to the south-eastern portion remaining

hypersaline year round (Noye 1974),

Unlike Lake Clifton, however, groundwater seepage

via aquifers to the Coorong, although probably

significumt, is minor Compared to the high evaporation

rate (Jensen et al, 1984; Noye 1974), Although drainage

earlier this century arund Salt Creek introduced some

freshwater into the Coorong for a short period. water

quality in the carly 1980s was probably similar to pre-

drainage (Jensen er al, 1984), from the time of early

exploration of the ares through unui 1864. Tf so, it

seems thut for at least a century salinities affecting

bryozoan growth have been similar to those of today,

and that even weir and barrage constniction (1920s and

1930s respectively) may not have radically altered

conditions. Prior to this, in periods of drought when

the Murray dried up, the Coorong would haye becume

saline, and major floodwaters reaching the mouth

would have freshened the lagoons, just as happens

today,

Salinity peaks have been noled to gradually. increase

over a penod of years (Jensen er al, 1984). Such

increases arc terminated by the release of large

quantities Of Murray waters during the spring of years

of heavy flooding upstream. Some of these waters reach

aod freshen the southern reaches of the Coorong before

the cycle begins again Vensen ef al. 1984),

Bryezoa

in 1987 Bone discovered laterally extensive sub-

Recent multi-laminar buildups (Fig. 5) of encrusting

anascan cheilostome Bryozoa. Individual buildups. are

up to 40cm in diameter and 30 em high. The most

extensive bioherm growth visible centres on Muaprath

Flat (Fig, Ie). The bryozoan was identified as

Membranipora agiculata (MacGillivray 1891), but 1

is now stivpested chat mt belongs w the genus

Conapeutn.

From miid-1987 to miil-1989 rare small scattered

colonies (1 cm dianteler) were found living intergrown

with the serpulid, Ficopamatus enigmaricus (Fauyel),

along the casters margin of the Coorong from the

Murray River to 2 km north of Policemans Point. The

range of (he living serpulids, however, was found to

extend ty Sait Creek (Bone & Wass 1990). In Spring.

1989, conditions were upparently more advantageous

lor the bryozoan. ft was found at Magrat Plat thriving

on almost every hard surface, from the consolidated

acolianite of relict sand dunes, to bottles (Fig. 6) and

tyres. Colonies up to 6 cnr in diameter were found,

bur only as unilaminar forms (Bone 1994). Later the

same year, serpulids out-competed the Bryozoa to

completely cover and conceal all trace of sub-Recent

bryozoan buildups in this locality. The same occurred

in 1991 with serpulid banks building on those of the

previous year (Fig. 7). Thus, locatmp the sub-fossil

bryozoan buildups in this area can be difficult even

when their position is known because of the thick

covering of serpulids, Searches during 1990-91 for new

locations of the sub-Recent bioherms in the northern

Lagoon proved fruitless because of the extent and

thickness of these serpulid “banks” In 1991 in

particular, for more than 20 km each side of Long

Point, serpulids were prevalent im the shallows (Pig.

8). They were found in cauliflower-shaped proups of

worm tubes attached to almost any potm ol hard

substrate piercing the carbonate-rich muddy sands.

They were even found colonising the handle attachment

hole of an old pick, growing out from it in Opposite

directions (Fig. 9).

Modern serpulid mounds range to >LO m in

diameter, with multi-seasonal growth making some up

to 0.5 m high. They form biohernns (Figs 7 & 8) with

many other animals additional to the Bryozou, such

as crabs, isopods, copepods, amphipods, decapods,

neriid worms (e.g, Cerdtonereis aequisetis Augener,

1913) and occasionally relatively large fish.

Fip, 7. Living serpuhd mounds encrustng Sub-Recent

bryozoan buildups in the northern Coorong Lagoon, 199},

Figure for scale.

7 M SPRIGG & Y.BONE

Methods

Mupping of the areal extent of sub-Recent buildups

of the bryazoan, C acicalata in the Coorong was

undertaken in concert with observation of its present

day range, The easter side of the Coorong Lagoon

was mapped in detail, fram Pelican Poin| to.8 km south

of Salt Creek (Fig. tb). A boat was used on several

occasions to cantinge mapping around reefs and islands

inthe centeal part of the lagoon, and along the western

shore,

Salinity measurements and observations of colony.

growth were made on a monthly schedule over two

years, from pdints along the eastern coastline. Water

saunples were collected for analyses at the same time.

These were analysed for their concentrations of

sodium, potassium, magnesium and calcium by the

atumic absorptiun method The salinity of the same

samples were tested by a Kent clevtrolyne conductivity

meter, temperature corrected to 25°C, and converted

ta % (total dissolved salts), Samples of very high

sulinity were diluted and the salinities re-scaled . us the

meter used was poorly scaled at very high values.

Results

The water salinity analyses confirm (he trends of

earlier studies (Noye 1974; Geddes J984, 1987; Botting

1990). Data is biased, as sumple sites had to be along

the eastern shoreline, Analyses of Na, K, Mg, and Ca

showed the elements were roughly in proportion to

seawater, but that usually Cas K. 4 water sample [rom

the River Murray (S.A.) had Ca>Kk.

Flooding of the Coorong by water released through

the barrages stirred Up the fine carbonate mud on the

floor of the Lagoon, reducing visibilty to almost zero.

Ths, plus the greater water depth, hampered the

mapping aspect of the study, Location of the Bryozaa

was offer by touch and not by sight, so only a Small

number were found. Howeyer, these were sullicient

io indicate that the buildups are widespread in the

southern Lagoon.

Sub-tossil boildups were Jound at three separate

levels, The highest and most altered level lies just below

the high watermark, tiniming the terraces previously

mentioned, The middle level is generally found 20-30

cm lower, These become exposed in summer, lying

about IS cm above the lowest water level noted, ‘The

lowest level is found approximately 50 cm lower down.

In the noptherm Lagoon the sub-Reeent © aeieulata

buildups form densely packed semi-flat layers (Fig. 10)

The buildups inthe southern Lagoon haye layers which

are convulute, which result in spaces occurring belween

the more laminar layers (Pig. 11). These spaces often

contain Coxiella confusa and other small gastropods,

minute bivalves and many different ostracods, as well

as foraminifera, The latler include Ammonia beceuril,

Kiphidium articulamm and Discorbis dimidiarus,

which aré marine fauna, and so may be midiecative ot

much less saline water being present in this part of

the Coorong at the time of bryozoan growth,

Altermatively, they may have been re-worked fram sub-

Recent sand dunes (Can pers. commn.), Qogonia, the

fruiting bodies of non-niarine charophytes (c.g. Ruppia

sp., Which is tolerant to a wide range of salinities from

fresh to hypersaline in the Coorong today ~ Wamersley

1984) are alsa found within these spaces.

In both 1989 and 1990 the bryazoan Cyphonaute

larvae had settled and begun ta bud asexually by the

beginning of Oetober. In 1990 at the end of the first

week in Oclober colonies 10 cm mm diameter were

common and widespread, growing in the central

southern Coorong, both aleng the shore and around

islands, and on “reefs” The most southerly discovery

of hying €. acteulara was on the edge of the shallow

hypersaline Bul Bul Basin, 4.5 km south of Salt Creek,

These rare colonies Were. Small encrustations on the

sides of consolidated terraces belonging to earlier

carbonate flats of a higher water level (Table lL.

Colony establishment time is not known for either

year at che northern end of the study area at Pelican

Point. At the torthern end of the Coorong, small

colonies (Table 1) were common both years in the fresh

waters adjacent to the barrages, where lake waters are

known io seep through,

A-sketch and cross-section over the Point adjacent

tu the Gentini Downs Boat Ramp illustrate 4 typical

southern Lagoon environment showing relationships

batween features and the sub-fossil accurnulations (Pig.

2). These deposits are extensively intergrown around

solution pipes. and some contorted bands (5-10 em

thick) contam teepec-like structures and possible

fractures. They are underlain by grey, semi-plastic

mud,

Tasie l. Size of colontey al madern Conopeum aciculats on

north-eastern margin of Coareng Lagoon, Size differences

were transitional between adjacent locations, “Warnecke

Point’ is ath unnanwl] feature lying seth of Hells Gare, on

the Warnecke property,

Location Masiimum Sia

(em)

Bryazoan Colonies = —

989 1990)

Pelican Paint 2

Mark Point 2.

Lop Pot -

Bonney Reserve —

Magrath Plat 6

“Warnecke Point” 7

Poltcertans Povat 3

2uuwo

We} Fp

it] 0

Boat Runip

Gemim Downs 2.0 trare) 2.75 (rave)

Boat Ramp

4.5 km south of 16

Salt Creeks

15 (rare)

BRYOZOA IN COORONG 93

Fig. 8. Close-up of living serpulid mounds, showing (heir

similarity to stromatolite growth form, The destruction of

the upper, central portion is caused by fishermen dragging

their boats across the bioherms, Individual mounds average

1m in width

Discussion

Due to conditions prevailing in the northern Lagoon

favouring prolific growth of thick serpulid colonies in

1989 and 1990, searches of this area did not reveal new

locahions of sub-fossil Bryozoa- In the southern

Lagoon, however, conditions were Jess beneficial to

both serpulids and Bryyzoa. In 1990 serpulid growth

was less abundant than in the northern Lagoon, but

1991 brought even poorer growth. Consequently

ubiquitious sub-fossil sub-Recent bryozvan buildups

were often exposed in the southern Lagoon.

The convoluted nature of many of the layers. of the

sub-fossil bryozoan buildups found jn the southern

Lagoon were unusual in that instead of being parallel

to each other like those found in the north, they were

often highly contorted. Some twist upwards, only to

turn sharply and grow back on themselves, thus leaving

“holes”. Others were regularly laminated similar w

those found in the northern Lagoon where one layer

is laid down directly on the last, The growth of

bryozoan colonies is largely affected by changes in their

environment and this can cause changes in colony

morphology (McKinney & Jackson 1989), ‘Thus it may

be possible to trace parucular periods-of environmental

change over Jarge areas of the lagoons through

examination of the growth patterns of different

buildups, if the contorted Iayers do indeed have a

similar pattern at a particular time.

The three separate levels of buildups found in the

southern Lagoon may have grown concurrently, bul

it is more likely that.as water Jevels are known to have

changed in the lagoons that each might belong to a

different time period. High water Jevels prevented clear

mapping of these ditferent levels. Radiocarbon dating

analyses currently being undertaken may elucidate

these and other timing uncertainties.

Microfossils found within the convoluted layers of

bryozoan buildups include Oogonia from algae

characteristic of marginal marine and non-marine

saline lakes, The Foraminifera are all extant, benthic

species, Cann (pets. comm.) suggested (i) that they

could have become encased by bryozoan growth alter

being washed in after eroding out of the surrounding

aeolianite, or (ii) that having crawled there, the

Foraminifera were trapped and died, as the fast

growing, C. aciculaia entombed them.

New colonies found a week into October 1990 were

common and widespread in the southern Layoon. They

were all about 1.0 cm in diameter. Colonies. 0.2 cm

larger or smaller than this were not present, Thus, a

switching signal for the beginning of asexual budding

must have occurted almost concurrently over a large

area. Actual growth rate was not determined, as a

precise settlement timing is not known, but growth

certainly appears to have been rapid compared to that

Fig. 9, Serpulid colony growing through the handle-hale of

a pick-axe, found in the northem Coorong Lagoun in 1991,

O4 HRYOZOA IN COORONG

Fig. 10. Common growth form of sub-Recent Conopeum

aciculata in the northern Coorong Lagoon, The

multilayered habirhas flat layers densely packed and relief

is low,

of most Bryozoa (McKinney & Jackson 1989). Growth

ceases and the colony dies once it is exposed.

The rate of multilaminar layer formation is also

unknown. Only one example of multilaminar growth

was found in present day colonies. The colony is

located on the internal walls of a solution pipe, where

iis well protected from high energy wave and storm

damage. Tt may be that mullilayer growth occurred in

(he one season, or that growth of the second layer

occurred ina later season coincidentally on the earlier

skeletal remains,

Today, the living Bryozoa are only found growing

on solid objects whereas many af the Sub-fossil

buildups form pavements (5-8 cm thick) that appear

to have grown out laterally from the terraces, over the

soft carbonate muds. Other bands, up to 20 em thick,

appear to have grown out laterally from consolidated

acoljanite, so that now they are completely

unsupported, other than for the hard rock they rim.

ft appeurs. they avay have originally grown over

unconsolidated sediment, since eroded away.

Observations, coupled with analyses, show that very

high salinities are lethal to C. aeiculara . These high

Fig. 1. Common grown: foro: of sub-Recent Conopeum

acied/ata in the southern Coorong Lagoon. The

multilayered habit is convoluted and distorted, and results

in many buildups becoming fragmented.

salinities occur early in summer in the southern

Lagoon, resulting in maximum colony size being

relatively small, especially compared ta those in the

northern Lagoon. This can be seen from Gemini

Downs southwards to Salt Creek, where colony size

ranges. from 2.75 cm down to 1.5 cm, suggesting 4 very

Short growth period, particularly in 1990. The switch

to the salinity level lethal to the organism could not

be pinpointed, as samples were collected nearshore and

thus did not always allow for local and duily salinity

variations, due to slow mixing. However, even here

daily variations are observed, confirmed by Botting

& Associates (1990).

Conclusions

|. Sub-Recent buildups of the bryozoan Conopeum

acioulata are widespread. in both the northern and

southern lagoons of the Coorong. The buildups are up

to 30 cm high and 40 cm in diameter. Growth form

in the northern Lagoon is multiserial, noultilayered.

gently arcuate and compact, Growth form in the

southern Lagoon 1s multiserial and multilayered but

the layers are highly convoluted.

2. Modern colonisation of hard. substrates in the

Coorong by €. aciculaia is seasonal, occurring in the

spring; and even more widespread than the sub-Recent

occurrences. Colony size increases rapidly, utilising

multiserial, unilaminar growth form. Maximom colony

size 16 6 cm in diameter, occurring in the northern

Lagoon, Serpulids frequently outcompete the Bryozoa,

thereby concealing evidence of annual bryozoan

presence.

3. Salinity in the Coorong increases southwards and

increases seasonally up to 60%, due to high

evaporation in summer. The higher salinities are lethal

to the Bryozoa but not to the serpulids. Similarly,

exposure duc to summer water-level fall kills the

Bryozoan but not the serpulid colonies. Cyphonaute

larvae are able to survive the high salinities, and re-

colonise when conditions are optimal, Small bryozoan

colonies occur in Coorong hyposuline waters adjacent

to the fresh-water lakes at the northern reaches of the

Coorong.

4. Modern C. aciculara occurs intimately associated

with thrombolites in fresh water on the eastern margin

of Lake Clifton, Western Australia.

BRYOZOA IN COORONG us

Acknowledgments

We would like to acknowledge R. Sprnigg, A. Chivas

and P. MacDonald for continued support and

suggestions; P, Cook, P. Bock and the Jate S. Parker

for assistance in identification of C. aciculata: I, Cann

for identification of Foramintfera; L_ Moore for

assistance with the Lake Clifion work; P. Hutchings

for identification of serpulids and neriid worms; S.A.

Dept of Fisheries for boat loan; S.A, Engineering and

Water Supply (J. Whitbread and A. Jessup particularly)

for their assistance; technical staff of the Dept of

Geology and Geophysics, University of Adelaide.

References

Bane, ¥. (1991) Population explosion of the bryozoan

Membranipora aciculaty in the Coorong Lagoon in late

1989, Aust. J. Earth Science 38, 121-123,

—____. & Wass, R, (1990) Sub-Recent bryozoan-serpulid

buildups in ihe Coorong Lagoon, South Australia. ?bid.

37_ 207-216.

——___ & James, N. P. (1993) Bryozouns as carbonate

sedimem producers on the coul-water Lacepede Shelf,

southern Australia. Sea, Geol. (in press),

Borminc, J. & Associates (1990) Coonsng and Murray

mouch working group project: Preliminary assessment of

rok data base, (Engineering & Water Supply,

aide)

Burngé, R. Y. & Moors, L. S. (1987) Microbialites:

Organosedimentary deposits of benthic microbial

cofmumunities. Palatos 2, 241-254

Co.iins, L. B. (1988) Sedimems and history of the Rotinest

Shelf, southwest Australia; A swell-dominated, non-tmpical

carbonate margin. Sed. Geol. 60, 5-49,

Esvewan, M, & Kiarea, C. F, (1983) Subserial exposure

pp.l-5S4, in P. A, Scholle, D. G. Bebout & C. H. Moare

(Eds), “Carbonate Depositional Environments’. 44?G

Memoir 33.

Geppes, M CC, (1987) Changes in salinity and the

distribution.of macrophytes, macrobenthos and fish in the

Courong Lagoons, South Australia, following a period af

Rivee Murray flow. Trans. R. Soc. S. Aust. 111, 173-181.

—____ & Burner, A. J. (1984) Physicochemical and

biological studies on the Coorong Lagoons, South Australia,

am) the effect of salinity on the distribution of the

macrabynthos. Ibid. 108, 51-62.

Gastin, V. A., Becperro, A. P & Cann, J, H. (195%) The

Holocene tton-trepical coastal and shelf carbonate province

of southern Austrilia. Sed. Geol. 60, 51-70.

Tonukm, M. & Cook, P, J, (1980) Paleomagactism of beach

ridges in South Australia and the Milunkovitch theory of

ice ages, Nanue 286, 699-702.

Jamas, N. P, & Bone, Y. (1989) Petrogenesis of Cenozair

temperale wuler calcarenites, South Australia; A model for

meteoric/shallow burial diagenesis of shallow water calcite

sediinents, J. Sed. Pet, 59, 191-203.

JENSEN, A., Hokv, P., Korii, Bh, Siemumep, &., ‘UiLL, M,

& WEINER!, M. (1984) The effects of drainage on

rovectesior behayiour in countries Cardwell and

uckenghara and the effect on the Coorang: A report for

ihe Minister of Water Resources 1981-1983. S.A. Dene

Enviranment and Planning 66, 8-65

MacGituivray, PH, (1891) Description of new or Litnle

known Polyzoa, Trans. R. Soc. 5. Aust. 3, 77.83.

Moore, L. S. (1987) Water chentistry of the coastal saline

lukes of the Clifton-Preston lakeland system, south-western

Australia, and ils influence ou stronutolite formation. Aust:

J. Marine & Freshwater Res. 38, 647-660,

Kwort, B. & Stancey, N. (1984) The stromiatolites

of Lake Clifton: Living structures ropresént the origins of

lite. Search 14, 309-314

—__—& Turner, J, V. (1989) Stable isatopic,

hydrogeochemical and nutriont aspects of [ake-groundwater

relations at Lake Clifton, pp, 201-213, /n G, Lowe (Ed,)

“Proceedings of Swan coastal plains groundwater

Management conference” (W. Aust. Water Resources

Council, Perth).

Noye B,J, (1974) The Coorong - An introduction. pp.1'79

Jn J. Noye (Ed.) “The Coorong”, Dept. Adult Education,

Onty, Adelaide Prob, 39,

pote rh E. Besa R E. J uy) Geology and

logy tinest Island, Geol. Sury, West, Aust,

oe

Rosen, M. R. & Cusner., L. (1992) A new location of

Holocene dolomite foomation, Lake Hayward, Western

Australia, Sedimentology 39, 161-166.

SEARLE, D. J. & Semeniuk, V. (1985) The natiral sectors of

the Inner Rottnest Shelf coast adjoining the Swan coustal

plain Jown, RX, Soc. W Aust. 67, 109-115

SEMENIUK, V. & SEARLE. D. J, (1985) The Becher Sand, a

new stnitigraphic unit for the Holocene of the Perth basin,

thid, 67, 109-115.

SpaiaG, R. C. (1952) The geology of the South-east Province,,

South Australia, with special reference lo Quaternary

coastline migrations and modem beach developments, Depr.

of Mines, South Aust. Bull. 29.

(1979) Stranded and submerged sea-beach systems of

southeast South Australi and the aeolian desert cycle, Sed.

Geol. 22, 53-96.

voN DER Borcu, C. C. (1964) The distribution and

preliminary geochemistry of modem carbonate sediments

af the Coorong area, South Australia, Geochim.

Cosmochim, Acta 29, 781-7199.

(1976) Stratigraphy and formation of Holocenc

dolomitic carbonate deposits of the Coorong area, South

Australia, J. Sed. Per, 46, 952-966.

WomersLey, H. B, 3, (1974) Plant life in the Coorong. Ja

J. Noye (Ed,) “The Coorong.” Depr Adult Edunation, Univ,

Adelaide Pub. 39, 81-88.

FEMALES AND IMMATURES OF THE AUSTRALIAN CADDISFLY

HYALOPSYCHE DISJUNCTA NEBOISS (TRICHOPTERA), AND A NEW

FAMILY PLACEMENT

BY A. WELLS* & D. CARTWRIGHT}

Summary

The female, larva and pupa of the northeastern Australian Hyalopsyche disjuncta Neboiss, 1980, are

figured and described for the first time. Comparisons between Hyalopsyche disjuncta and several

North American species of Phylocentropus Banks, 1907 (Family Dipseudopsidae Ulmer, 1904),

show that both genera share apomorphous larval features. Transferral of Hyalopsyche Ulmer, 1904,

from the family Hyalopsychidae Lestage, 1925, to the family Dipseudopsidae and supression of

Hyalopsychidae are proposed.

KEY WORDS: Trichoptera, Dipseudopsidae, Hyalopsyche, larvae, pupae, females.

Transactians af the Royal Saciety of §. Aust. (1993), 417(2}, 97-104.

FEMALES AND LIMMATURES OF THE AUSTRALIAN CADDISELY HYALOPSYCHE

DISJUNCTA NEBOISS (TRICHOPTERA), AND A NEW FAMILY PLACEMENT

by A. WELLS? & D. CARTWRIGHT}

Summary

Weuis, A. & Caxrwricur,, D. (1993) Females and immatures of the Australian caddisfly Hyalopsyche disjunen

Neboiss (Trichoptera), aiid a new family placement, Trans. R. Sec, S. Aust, 107(2), 97-104, 4 June, 1993,

The fernale, larva and pupa of the northeastern Australian Hyalopsyche dixjancta Neboiss, 1980, are figured

and described for the first time, Comparisons between Hyalupsyche disjuncia and several North American species

of Phylocentropus Banks, 1907 (Family Dipseudopsidac Ulmer, 1904), show that both genera. share apomorphous

larval features. Transférral of Ayalopsyche Ulmer, 1904, from the family Hynlopsychidae Lestage, 1925, to the

family Dipseudopsidae and supression of Hyalopsychidac are proposed.

Key Woros: Trichoptera, Dipseudopsidae, Hyalopsycke, larvac, pupae, females.

Introduction

The history of unstable taxonomy of the group of

polycentropodid-like caddisfly genera including

Dipseudopsis Walker, 1852, Protodipseudopsis Ulmer,

1904_ Phylocentropus Banks, 1907, and Hyalopsyche

Ulmer, 1904, is discussed in detail by Weaver &

Msalicky (in ms), These authors:present strong support

for the familial status of Dipseudopsidae, including the

first three of the above-mentioned genera, and possibly

also Hyalopsyche, an African-Oriental genus, which

was placed by Schmid (1980) with Phylocentropmus in

the family Hyslopsychidae. Convincing evidence for

the monophyly of Dipseudopsidae Ulmer, 1904, sensu

Weaver & Malicky, derives from a consideration of

features of the female abdomens and larvae.

Although reference bas been made in the past to

females of Hyalopsyche (Ulmer (1915) identified three

New Guinean female specimens as H. rivalis (Betten,

1909)). no descriptions of females are available, and

unti] now no larvae have been associated,

A single Australian species of Hyalopsyche, H.

disjuncta Neboiss, was described hy Neboixs (I980)

from two adult males from northern Queensland,

Repeated efforts to collect more specimens, mcluding

larvae, failed; H, disjuncta appeared to be rare,

Recently, however, during an intensive collecting tip

of two week's duration in the Jardine River urea of fr

northern Cape York (as part of the Royal Geographic

Society of Queensland’s “Wet Season” Cape York

Peninsula Scientific Expedition), the authors collected

* Northern Territory Museuin of Arts and Sciences, PO Box

4646, Darwin, NT: O8Ql. Present address: Australian:

peabeie Resources Study, GPO Box 636, Canberra, ACT

260)

a Bileey Laboratory, Werribee Treatrnent Complex, Private

Rag 10, Werribee, Victoria 3050,

numerous adults of both sexes, and just prior to

departure, made the exciting find of several larvae and

pupae. Female and Jarval and pupal features of this

species aré here described and illustrated, and the larval

niche as described. For comparisons, some larval

features of Phylocenirams sp., and the female genitalia

and head showing the tentorium of the North American

P flucidus (Hagen) are also figured.

Confirmation of sharing of the specialised larval

niche and other synapomorphous larval character

states, firmly supports the monophyly of

Phylocentropus — Hyalopsyche, and we present an

hypothesis for the derivation af the specialised female

penilalia af Hyalopsyche from those of Phylocentropus,

We argue that Ayelopsyche should be placed in the

family Dipseudopsidae sensu Weaver & Malicky (in

ms), and the name Hyalopsychidae is here-supressed,

Ross (1965) and Ross & Gibbs (1973) discussed the:

phylogenetic relationships of Dipseudopsinae (in the

Psychomyiidse and subsequently in the

Polycentropodidie), but with more data available,

phylogenetic analyses of higher taxa of Trichoptera now

in progress (Weaver & Frazer pers. comm.) should

provide a more definitive assessment of the

relationships of this family.

Materials and Methods

Adak Hyalopsyche disjunct were taken mainly at

12v or By black or UY lights; adults and larvae were

collected into and stored in 70% ethanol. Comparative

material of the North American Plyocentropus tuciduy

(adults) waa provided by Dr J.S. Weaver, and adults

of P placidus and Dipseudopsis indicus.and larvae of

Phylocentropus sp. were provided on loan by Dr J.

Motse.

Specimens were prepared for study by treatment in

caustic potash, or using a lacto-phenol preparation.

Drawings were made using camera lucidas on a Wild

98

A. WELLS & D. CARTWRIGHT

m 4

0.1 mm

Figs 1-3. Hyalopsyche disjuncta Neboiss, female: 1, dorsal head showing tentorium; 2, mesothorax, dorsal view; 3, genitalia,

lateral view.

Figs 4-6. Phylocentropus lucidus (Hagen), female; 4, dorsal head showing tentorium; 5, mesothorax, dorsal view; 6, genitalia,

lateral view.

Scale bars = 1 mm, unless otherwise indicated.

FEMALES AND (MMATURES OF THE AUSTRALIAN CADDISFLY 99

M5 dissecting microscope and a Wild M20 compound

Microscope,

Material ts deposited in the following institians:

Australian National Insect Collection, Canberra

(ANIC), Museum of Victotia, Melbourne, (NM);

Northern Territory Museum of Ans and Sciences,

Darwin, (4TM); and Queensland Museum, Brisbane,

(QM).

Systematics

Hyaloprehe disiuncta Neboiss, 1980

PIGS 1-3, 7-13, '6, 18-21

Hyalopsyche disjuncoa Neboiss, WH 357-361; Figs 1-8,

Holotype male, N, Quecnsland, 29.iv.1979, NMV.

Material examined: N, Queensland: 1 o, it? 50°S

142° 30°F, Bertic Creek, at Telegraph Crossing,

5.11.1992, D, Cartwright ond A. Wells, NTM; 2 > or,

2 99, UP 50'S 142° 30'E, Dulhunty River, a

Telegraph Crossing, 8-9,i7,1992, 1D, Cartwnght and A

Wells, NMV; 2 oo, 1 9, I" 39°S 342° 28'E,

McDonnell-Cockatoo Creek jn, 14,7i,1992, D,

Cartwright and A. Wells, NTM; DB oo, § 9 9, i"

44'S 142° 29'E, Gunshot Creek, at Telegraph

Crossing, 17.11.1992, D, Cartwrighi and A, Wells, (M-

ioe,’ & 9, same loc. and collectors, 4-15.1L1992.

QM; S oc, 15 9 ©, same loc. and collectors,

18-19,71, 1992, NMY; 21 ovo. 4 9, same loc.,

$-5,iV.1992, M. Crossland, ANIC: 2 oa, 19 9G,

same loc, and collector. 10-11.1v.1992, NM; | pupa,

same loc., D. Cartwright and A. Wells, 18.13.1992,

NTM; 2 larvae, satne loc and collectors, NTM; 7

larvac (L reared to adult 9), J pupa, same Joc. und

collectors, 19,7i,1992_

Neboiss (1980) illustrates the general body features

of male Hyolopryche disjancia, and gives anterior wing

length as 6 mm, but in our samples male anterior wing

length ranges from 4.3-6:2 mm. Females conform with

males in general features, including reduced maxillary

Palpi.and warts on the mesothoraa (Fig. 2), but are

considerably larger. An additional feature. the

temtorum, is. figured (Fig. lt: its arms are well

developed, with inner lateral Manges posteziarly, and

the posterior bridge complete

Female (Figs 1-3). Anterior wing length 0-80 mm.

Genitalia im the form of a slender, elongate, non-

Tetractile ovipesitor (Fig. 3). Sternité VIII not

subdivided, exterrial gonopod ot segment VIII slender,

laperead, reachmg almost in tip of segment [X.

Postenoriy segment X is divided to form pair of distal

lobes. each with apical cercus and one papilla partially

fused on inner margin, the second projecting venteally

from the outer Margin at about the level af division

of segment X.

Larva (mature) (Figs 7, 8 12. 15, 16, 38-20).

Campodeiform, slender, elongate, (Fig. 7). Mend,

pronotum, limbs and anal claws brown-yellow or pale

brown, dorsal margin of prothorax and strengthening

tidges of thoracic pleury black, rest of body pale. Body

length 6.0-14.8 mm, head capsule width 0604.88 mm-

Head round Ja dorsal view (Fig, 12); fronteclypeus

sub-tnangular, extending to posterior of head capsule,

anterior margin rounded, wntorial pits situated

posteriorly; labrum broadly rounded with few setae,

anterior margin finely divided. Maidibles (Fre. 16)

stout, dissimilar, each with differing numbers of

denticles apically and on inner distal margin, and with

a dense penicillus medially om inner margin, antero-

lateral margins crenulate; right mandible ragose antero-

dorsally. On ventral head (Fig. §3), anterior ventral

&potome (gula) short, forming an isosceles triangle;

submenium rounded anteriorly, paired setae situated

subapically; maxillary palps relatively slender, lobus

intertius broad, obliquely truncate in dorsal view, and

bearing a brush of haiz apically; labium modified to

form a slender, elongate spinneret, about 34 as long

as head capsule, labial palpi absent.

Prathorax only with sclerotised tergite. Fore

trochantin fused to pleuron (Fig. 18). Mesostemum

(Fig. 8) with a pair of elongate sclerites posteriorly,

Legs (Figs 18, 19) short, flattened, tarai with dense

Fringes of setac, claw on fore limb strongly curved,

Fore tarsus with an udpressed row of modified setae

on the mesial margim.

Abdomen elongate, with a lateral fringe of Fine setae,

without gills, but with a group of anal papillae (Fig.

7}; anal prolegs clongate, with cluster of lang setae

anterior to the slender strongly curved anal claw (Fig,

20).

Pupa (Figs 9-11, 21). Length 7.5-10.3 sun Labrum

(Fig, 9) short, with a smal) marginal papilla apico-

mesally. Mandibles (Fig. 10), simple, slender, curved.

Hook plates (Fig, 21) with 3-5 demicles, on abdominal

segments IT-VIIT. posterior plates on segment V only,

developed ints hand-like processes, Posterior abdomen

(Fig, Lf) divided into two simple setose lobes.

Biology. Larvae and pupae were extracted from

fragmented silken tubes. The exact architecture of the

larval dwelling is unknown, but the branched portions

of rabeg appear to have all the features of the tubes

of Phylocentrapys placidus (see Wallace er al 1976)

and FP carofinus Carpenter (see Wiggins 1977, Fig,

19.4D). (Such features are also shared by

Protedipsendopsis sp. (uh-named) as jllustrale! Sy

Gibbs (1968, Fig, 3)). The architecture of the

Hyalopsyche disjuncta dwelling is probably similar:

a blunt-ended vertical branch with a small wpening

Projected into the flaw, a series of connected cross

tunnels, and a dilated net chamber.

Habient, Larval tubes were collected from a sinall

atream with a rather unstable base of coarse sand, The

dwelling tuhes were found attached to logs amongst

deeply packed sand and litter, usually in situations

Where the Utter, trapped against logs embedded in the

100 A. WELLS & D, CARTWRIGHT

Figs 7-8. Hyalopsyche disjuncta Neboiss, larva: 7, whole animal, lateral view, 8, mesosternum.

Figs 9-11. Hyalopsyche disjuncta Neboiss, pupa: 9, labrum; 10, mandibles, 11, posterior abdomen.

Scale bars = 1 mm, unless otherwise indicated.

FEMALES AND IMMATURES OF THE AUSTRALIAN CADDISFLY 101

13

Figs 12, 13. Hyalopsyche disjuncta Neboiss, larval head, dorsal and ventral views.

Figs 14, 15. Phylocentropus sp., larval head, dorsal and ventral views.

Scale bars = 0.1 mm.

102 A. WELLS & D. CARTWRIGHT

Figs 16, 17. Hyalopsyche disjuncta Neboiss and Phylocentropus sp., larval mandibles, dorsal view.

Figs 18-20. Hyalopsyche disjuncta Neboiss, larva. 18, foreleg and proleuron; 19, midleg; 20, anal proleg.

Fig. 21, Hyalopsyche disjuncta Neboiss, pupal hook plates, right side,

Scale bars = 0.1 mm.

FEMALES AND IMMATURES OF THE AUSTRALIAN CADDISFLY wy

sand, was begmming to decompose. Collections of

adults were obtained at lights beside smull and larger

streams; the smaller streams generally were almost

completely shaded by the canopies of riparian drees,

the larger 4treams were open,

Discussion

Comparisons of features of latval Hyalepsyche

disfuricra With those of Phyloceniropus placidas, P

fecidies and P. carolinus (fram Ross & Gibbs 1973 Fig,

2; Wiggins 1977 Fig. 15.41, and this work, Figs 4,

15) reveal extraordmarily ¢lose similarities between

species mm the two genera, In derived or specialised

features, la general form the heads are similar

(compare for example Figs (2, 13 and Phylacentrapus

sp., Pigs 14, i): mandibles of all species are stout,

with rugose sculpturing dorsally, albeit on only the

right mandible in 4, disjurcta; ventral sclerites differ

slightly, with the submentum of H. dis/uncie rounded

rather than subquadrate; the iobus tjrternus of the

maxillary palp of 4, disjuncta is broad and apically

truncate rather than tapered and rounded: labia are

modified te form virtually identical slender, elongate

spinnerets. Thoracic features, too, show close

resemblance and the highly specialised legs differ only

in the exact shape of the specialised setae adpressed

to the mesal margin of the fore tarsi. A. disfwreta Jacks

gulls but conforms with Phylocentropus species in fonn

of anal prolegs.and their claws, The larval niches

appear to be wentical,

Pupae are relatively uninformative: mandibles of

Hi, disjuncte resemble those illustrated by Ross (1944,

Fig. 204) for P. placidtes. Little information is available

On pupye of Phulocentropus.

Some of the above features decur in other caddis

larvae. For example, specialised clongate spinnerets

are seen in larvae of some Psychomyiidae and some

Ecnomidae, although tn these families the spinnerets

are proportionally shorter than in PhWoeentropas and

Hyalopsyche. Simple anal claws also occur elsewhere,

but are probably plesiomorphous. The highly tnodified,

flattened limbs, the form of the mandibles and the larval

niche are found only in Hyalapsyehe, Phylocentrepies,

Protedipseudopsis and Dipsenedapsix.

The probability of such a suile of resemblances

evolving by converyence are slim. The simplest

hypothesis is that these taxa anvse from the sante

specialised ancestral stock.

Adult characters do not concur with those of the

larvae. No clear synapomorphies can be recugnised,

which ms the very reason Hrat placement of these genera

has been confused for so long. Aniong adult features

several small differences in general body features can

be observed benwecn Hyalopsioke and Phyocentropus,

such as the arrangement of setac an the meta-scutellum

(see Figs 2. 5) and the absence of several crossveins

in the Wings, a8 well as other more major anes, such

as shape of head, position of antennae, reduction of

maxillary palpi and form of genitalia, These differetices

probably represent autapomorphies for the particular

genera: The wings of Hyalopsyche can be derived from

those of Pkylocentrapus, which closely resemble

polycentropodid wings, simply by loss of several

crossveins; those of Phylocentropus probably represent

the plesiamorphous form. Similarities such as form

nf the tentormin (Figs |, 4) are probably also

plesiomorphous,

Superficially, the female genitalia of Hyaiopsyche

disjurcta (Fig, 3) appear to be strikingly different from

those. of Phylocentrapus placidas (Fig, 6), Yet upon

close’ examination, similarities ase apparent,

particularly in the cerct and papillae, Derivation of the

Hyalapsyche torm from that of Pirylacensropus tequires

only simple elongation of the arrangement stern in

Phylocentropus, probably adapting Hyalopsyche

females for oviposition into sand af crevices,

Investigation of the female of Dipseudopsis indtoa

McLachlan during this study, indicated that an the basis

of female genitalia and wing form, Phylocensropus and

Hyalopsyche are closer than Dipseudepsis-

Hyalopsyche.

Association of females and larvae of a species in

Hyalapsyche has helped to stabilise the higher level

taxonomy of this genus, placing it in the family

Dipseudopsidae. On the basis of studies reported here

and examination of tite literature, we suggest that

Hyalopsycke and Phylacenrrapus tepresent sister

groups.

Ross & Gibbs (1973) suggested on the basta oi

similarities between FPhylocentrepus and

Polycentropodinae that Phylocensrapus is closer fo the

ancestral stock than other Dipseudopsinae

(=Dipseudopsidaz) and, given its occurrence in the

Oligocene Baltic ambers. they suggested a Eurasian

origin for ihe subfamily (=family). We have not found

evidence with which to dispute this contention — the

single north-eastern Australian species in Hyalopsyehe

is undoubtedly of Oriental origin. Careful phylogenetic

analyses in the future will elucidate tntrafimilial

relutignships, bul al last ite family Dipseudupsidae is

reunited,

Acknowledgments

We thank the Royal Geographical Society of

Queensland for providmg the opportunity and financial

support for field work at “Heathlands” and the

surrounding Jardine River area of Cape York, and for

the zenerous help and assistance given to us by

members of the Socicty durimg aur stay ut “Heathlands"

D, Cartwright’s work was undertaken with backing

from his family, and A. Wells’ with backing from the

Northern Territory Museum of Arts and Selences,

Darwin. We are grateful w beh partics.

104 A, WELLS & D. CARTWRIGHT

Dr J.S. Weaver of the University of New Hampshire,

USA, kindly provided material of Phylocentropus

lucidus and Dipseudopsis indica from his collection for

comparisons. Adults of Phylocentropus placidus and

larvae of Phylocentropus sp. were made available by

Dr J. Morse, on loan from Clemson University, USA.

References

Banks, N. (1907) Descriptions of new Trichoptera. Proc.

Ent. Soc. Washington 8, 117-133.

Betten, C. (1909) Notes on the Trichoptera in the collection

of the Indian Museum. Rec. Indian Mus. 3, 231-242.

Gisss, DG. (1968) The larva, dwelling-tube and feeding of

a species of Protodipseudopsis (Trichoptcra:

Dipseudopsidae). Proc. Ent, Soc, London 43, 73-79.

Lestace, J.A. (1925) Notes trichopterologiques. VIII, Etude

du groupe polycentropidien et catalogue systématique des

genres et espéces decrits depuis 1907 dans Genera

insectorum. Bull, Ann. Soc. Entomol, Belg., 65: 93-120.

Nesoiss, A. (1980) First record of the subfamily

Hyalopsychinae from Australia (Trichoptera:

Polycentropodidae). Arch. fiir Hydrobiologie , 357-361.

Ross, H.H. (1965) The evolutionary history of

Phylocentropus (Trichoptera; Psychomyiidae). J. Kansas

Ent. Soc. 38, 398-400.

____ (1944) The Caddis Flies, or Trichoptera, of Mlinois.

Bull. Illinois Nat. Hist. Survey 23, 1-326.

___ & Girss, DG. (1973) The subfamily relationships of

the Dipseudopsinae (Trichoptera, Polycentropodidae). J.

Georgia Ent. Soc. 8, 312-16.

Scumip, F. (1980) “Les Insectes et Arachnides du Canada,

partie 7”. (Agriculture Canada). 296 pp.

ULMER, G. (1904) Uber Westafrikanische Trichopteren.

Zool, Anz, 28, 353-359.

_____ (1915) Trichopteren des Ostens, besonders von Ceylon

und Neu Guinea. Disch. Entomol. Z., 1915, 41-75.

WALKER, F. (1852) Catalogue of the specimens of

neuropterous insects in the collections of the British

Museum, Pt, 1, 192 pp. (London).

WaLtace, J.B., WoopalL, W.R., & Staats, A.A. (1976)

The larval dwelling-tube, capture net and food of

Phylocentropus placidus (Trichoptera: Polycentropodidae).

Ann. Entomol. Soc. America 69, 149-54.

Weaver, 1S., WI., & Macicxy, H. (In ms) The genus

Dipseudopsis from Asia (Trichoptera: Dipseudopsidac).

Wiccins, G.B. (1977) “Larvae of the North American

Caddisfly Gencra (Trichoptera)”. (University of Toronto

Press: Toronto and Buffalo).

OBSERVATIONS ON THE BREEDING BIOLOGY OF A MICROHYLID

FROG (GENUS OREOPHRYNE) FROM NEW GUINEA

BY G. R. JOHNSTON & S. J. RICHARDS

Summary

Transactions of the Royal Society of S. Aust. (1993), 117(2), 105-107.

BRIEF COMMUNICATION

OBSERVATIONS ON THE BREEDING BIOLOGY OF A MICROHYLID FROG

(GENUS OREOPHRYNE) FROM NEW GUINEA

The conquest of terrestrial environments is a major theme

in vertebrate evolution. Classically the Amphibia have been

viewed as an intermediate stage between the aquatic and

terrestrial groups!?. However, the standard view of the

amphibian life-cycle involving eggs hatching into aquatic

tadpoles which subsequently metamorphose into terrestrial

frogs belies the true diversity of reproductive modes in the

Amphibia.

The diversity of reproductive modes is greater among

amphibians than in other vertebrates*. All orders of the

Amphibia include species which deposit eggs in terrestrial

environments, and terrestrial development has evolved several

times within the order Anura®.

Deposition of eggs out of water is a major step toward

terrestriality among amphibians. Therefore, it is not surprising

that most variation in reproductive modes among anurans

occurs in tropical environments where there are high levels

of atmospheric moisture’. A second major step toward

terrestriality occurs in those groups which exhibit direct

development. The evolution of direct development has been

important in the successful invasion of montane environments

by amphibians*. Direct development occurs in all

Australopapuan microhylid frogs‘.

Despite their abundance, little is known about the breeding

biology of New Guinean microhylids. Small numbers of large,

heavily yolked eggs are concealed in leaf axils, hollow stems,

moss clumps or beneath the ground*. They may be attended